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PHYTOLOGIA

Designed to expedite botanical publication

1. 23 March, 1972 No. 1

b CONTENTS

BOIVIN, B., Flora of the Prairie Provinces. Part III (continued) ......++-+-

RUDD, V. E., A new variety of Poiretia latifolia and a brief resume of

: the genus Poiretia Vent. (Leguminosae) ....-+++++++55> 141
ROBINSON, H., The status of the genus Cryphidium (Bryophyta: Musci) .149
BING, R. M., & ROBINSON, H., Studies in the Eupatorieae
j (Asteraceae ). LXV. A new genus, Neocabreria ....-.+++:+: 151
“KING, R. M., & ROBINSON, H., Studies in the Eupatorieae
a (Asteraceae). LXVI. The genus, Pachythamnus ......--+-> 153
OGIMI, C., & KORF, R. P., Discomycete flora of Asia, Precursor IV:
' A new species of Bifusella (Rhytismataceae, Hypodermateae)
< on Cumminghamia in Okinawa... 1. ++ 620s eee reer trees 155
REVEAL, J. L., Descriptions of new species and combinations in
Eriogonum (Polygonacedé) ......++++++000errrreertenees 163
HALE, M. E., Jr., Parmelia jamesii, an unusual species in Section
‘ Imbricaria (Lichenes) from Australia and New Zealand ....... 179
MOLDENKE, H. N., Novelties from Brazil, Peru, Venezuela, and Thailand .180
MOLDENKE, H. N., Additional notes on the genus Verbena. Ed ARAN ne 181
197

MOLDENKE, A. L., Book reviews .....-..0 see e eect ters r reeset

oe

Published by Harold N. Moldenke and Alma L. Moldenke

303 Parkside Road
Plainfield, -New Jersey 07060
US.A.

Price of this number, $3.50; per volume, $7.50, in advance,
or $8, at close of volume

LIBRARY

FLORA MAR 9 19!-

OF THE PRAIRIE PROVINCES

NEW YORK

Bernard Boivin BOTANICAL GARDE

Fact ic.

(continued)

113. COMPOSITAE
(COMPOSITE FAMILY)

Floret lacking an involucel and typically with
5 anthers fused into a ring around the style. The bulk
of the species with flowers in heads belong in this fami-

Lys

a. Heads radiate.
b. Pappus of capillary bristles.....cccececcee eo Group I
bb. Pappus lacking or different, of awns,
chaff, scales or minute bristles.....c.ee.e.eeGroup II
aa. Florets all ligulate or all tubular.
Ge Hees! GLACOLC jeckclcles clo Suecicceciss covcieneeeisOLOUp ELE
CEs PlOre tS AL VelaGWLa te! terwsleere wisle'ciete'e viols c's cfeeGrOUp LV

Group I
Heads normally with a ring of ligulate flowers.
Central flowers all tubular. Casuals sports may be
double or discoid. Species that are normally discoid
are also included in Group III. Pappus of numerous fine
bristles usually at least as long as the achene.

a. Ligules white, pink, blue or purple..........Group l1-A
SLO I LOW: 1s clafacitisiae sia eile wie es vies tele ete ees Siccs maeaGrOoup L=5

Group 1-A
Ligules not yellow.

ea. Annual with a taproot........l14. Machaeranthera p. 126
aa. With a rhizome or caudex, rarely perennial
with a taproot.
b. Tegules narrow and numerous, all of the
same length or a few of the outer ones
much shorter... ..-ssseccecccscsceslds Brigeron p.127
bb. Broader and unequal, usually imbricated
and the outer gradually shorter, sometimes
the outer larger than the inner and + folia-
ceous.
{1] 85 COMPOSITAE

Peay TOLOG IA Vol. 23, no. 1

c. Ligules coloured and/or the
inflorescence not corymbiform.
d. Stemless....+.cceceseels. Townsendia: pelos
dd. Stem present.
e. Heads many or rarely single
Qnd small. ie csc cece eels AStenEpemLOs
ee. With a single large head.
f. Disk 2-3 cm across when
dried; perennial from a
taproote.s..e. 12. Townsendia p. 108
ff. Somewhat narrower; peren-=-
nial from a branched
CAUDEXK sess -ccecoee 13. ABStenimn tos
cc. Flowers white; heads in a corymb.
ge Pappus of dimegueth bristles, the
outer about 1 mm long..... 13. Aster p. 109
gg. Much longer and isome=

GUSth so wo ccsseccsesc ce GO. SOlIdecome ee
Group 1-B
Ligules and florets yellow.
Bie LEGVER OPPOSMES sss ..0 dvi vals s os epee 47. Arnica pee

aa. Leaves alternate or all basal.
b. Monocephalous plants.
c. Tegules of uniform length..15. Brigeron p. 127
cc. Imbricated, the outer gradually
shorter.........++++eeee04 9. Haplopappus p. 106
bb. Heads normally numerous.
d. Tegules nearly all of the same
length, a few of the outer ones
many times shorter......... 48. Senecio p. 183
dd. Much imbricated,.
e. Perennial from a rhizome or
caudex... .....+.2.05-2+28. Solidago p. 98
ee. Taprooted perennials.
f. Pappus bristles dimegueth,
the outer much shor-

teLee.s-esee++eee0 7. Chrysopsis p. 98
ff. Bristles of uneven length

but not sorted out in two
serieS........e0 9. Haplopappus p. 106

Group II
As in Group I but the pappus not of bristles,
sometimes lacking, or of scales, or awns, or chaff,
the latter often setaceous-tipped.

&. Ligules not yellow, mostly white, but

OLECENGCOLOUEE go \tarransejanate v, sidierere, Hath ccsher erate Group-II-A
aa. Ligules yellow.

COMPOSITAE 86

1972 Boivin, Flora of Prairie Provinces 3

b. Receptacle chaffy or bristly.......... Group-II-B
bb. Receptacle naked... ....cccecceseeessee Group=lI-C
Group II-A

Radiate in white, pink or purple, not in yellow.

a. Lemves opposite ....+++-++e+++e+++33- Galinsoga p. 161
ea. Alternate or all basal.
b. Leaves entire or nearly so.
c. Rays 2 cm long or more...26. Echinacea p. 154
cc. Much shorter.
d. Stemless or monocepha-~
OWS <ieensiele selce sie eebtlae) Lownsendia: p. 108
dd. Stem present; heads nume-
TOUSis ss csls esislelsiclsasisia, ove BOLtonia p. Los
bb. Dentate to much dissected.
e. Closely or coarsely dentate.
f. Ligules deep red
DrOwWNeceescccccecce 39. Gaillardia p. 164
ff. White.
g.- Ligules +tlmm long 41. Achillea p. 165
gg. Larger, 3-5 cm
long........ 43. Chrysanthemum p. 169
ee. Finely and repeatedly dissected.
h. Ligules 3 mm long
Or eésSic nc enniewe ce Sh. Achillea ip. 165
hh. Larger, 4-20 mm long.
i. Receptacle naked...
Dit wiiiddistddetente, Matricanin ps 167
ii. Bach floret subtended
by a bractlet (= chaff)...
eee cescccecececoe 40. Anthemis p. 165

Group II-B
Heads radiate in yellow and the receptacle chaffy
or bristly, that is with the tubular florets individual-
ly subtended by bractlets or bristles.

a. Leaves opposite, becoming alternate in the inflores-

cence.
b. Rays formed by the larger inner
EegGulesivinuw. «ips ols -Sisictse <csiatvohee es OCs, Bidens p. 160

bb. Rays formed by the prolongation of
the corolla of the peripheral flowers.
c. Leaves narrowly pinnatifid to
bipinnatifid..........e. 30. Coreopsis p. 159
cc. Entire to coarsely lobed.
d. Head center much higher than
wide; receptacle conical; peri-
pheral florets fertile .......

Ore ee ee eeeeeeeeecese 24, Heliopsis p. 152
87 COMPOSITAE

Pn Yh OL 0'e Fa Vol. 23, no. 1

dd. Head not so high; receptacle
flattish; peripheral florets
sterile .tsccccceces 296 Helianthusipeplon
aa. Leaves alternate.
e. Ligules bicolour, deep red-brown towards
the Mbasescrers'<.1- Jotdcciclelicicicee G9. Glan Larciem pen or:
ee. Yellow.
f. Head cylindric with 4-5 drooping
IAL GuLes «.iueeieddedeeccas 27s Ratibide peas
ff. Head shorter and the rays much more
numerous.
g. Leaves mainly basal, the cauline
ones only 1-3 and much reduce ...
div stele sis letetetelets’ots eo! 26 Balsanorhi.za pews
gg. Stem quite leafy.
h. Leaves narrowly dissected ..o.
ces siteuscleeselsewe 40. Anthemia peas
hh. Entire to dentate.
i. Disk flattish to somewhat
convex..... 29. Helianthus p. 155
ii. Hemispheric to oblong
and very protuberent..
Weisteicieieles ele) COS eRUd DECK al peelos

Group II-C
Heads radiate in yellow; receptacle naked, i.e.
neither chaffy nor bristly.

Bie) LEAVES [OPPOSE te sciclcle cistclsleicislcleclocieeis CO. Bahvaupelog
aa. Alternate or basal.
b. Leaves all basal or remotely pectinati-
PALtite 2... esse eeerecccces 37. Hymenoxys p. 163
bb. Stem leafy, the leaves entire or less
narrowly divided, dentate to pinnatifid
with broad lobes.
ce Somewhat shrubby at base; heads small
and numerous .......e-... 6 Gutierrezia p. 98
cc. Stem herbaceous to the base; heads
larger or longer.
d. Heads rather inconspicuous, the
rays only about 2 mm long and only
l-3 per head.....eseceoe.. 34. Madia p. 162
dd. Ligules more numerous and much
longer.
e. Heads very sticky; tegules
strongly squarrosSe.......e.

esc crescseseceeee Oe Grindelia p. 97
ee. Tegules neither sticky nor
squarrose.

f. Ligules cumeate, coarsely
trilobed at summit ....
eosseecceee--e 38. Helenium p. 164

COMPOSITAE 88

1972 Boivin, Flora of Prairie Provinces 5

ff. Ligulate or slightly
wider near the middle
and finely 3-toothed at
summit «... 49. Calendula p. 192

Group III
Heads discoid.

a. Leaves and/or heads spiny or catchy by hooked
DETCK LAS) eres aise, « diols tlele vole side cisislelaiecewaeien Group III-A
aa. Not spiny.
b. Leaves opposite or verticillate .... Group III-B
bb. Alternate.
c. Various special types: semishrubby,
or the heads all or mostly in termi-
nal racemes, or the heads globular
with the outer florets pendant .. Group III-C
cc. More run-of-the-mill types.
d. Pappus of capillary bristles .Group III-D
dd... backing, or, Chatty, o-s00<se.000 GLOUD) LLL—E

Group III-A
Tegules and/or leaf lobes ending in stiff sharp
spines or with hooked tips.

a. Head catchy, the tegules ending into spines hooked
at tip.
Be Fruits, ax Liarys wiied «5s aswiess ony REDEEM p, 151
DOME HeGaemeLermi nAls ics isistslameswisiekias Olean hccet il Pp. 19S
aa. Spines not hooked but stiff and sharp at tip.
c. Only the heads spiny.
d. Heads all terminal and spiny .....

RR sath. coven miserews DOs. CONERUTSR: Pp. 200
dd. Some heads axillary and spiny ....
ace eh eo cpaateanGeees oh5 5 ae Wiseerey jae se,
cc. The leaves also spiny-lobed.
e. Pappus plumese 2... .ccssees. 54. Cirsium p. 196

ee. Barbellate or glabrous.
f. Barbellate; tegules gradually
tapered from base to tip .....
tinininininpnieceisivivln (cho sirysinie OOM OCOGGULSID. BLS
ff. Pappus bristles glabrous; tegu-
les constricted towards the
middle ...«svccsseccee 55. Silybum p. 200

Group III-B
Leaves opposite or verticillate.

a. Achene devoid of pappus.
b. Head with an involucre very different
BrQ@e the LSAVSS dsicncewae eves vanes (Clu cee 2a0

89 COMPOSITAE

Pye. T) Ovl OG. DA Vol. 23, no. 1

bb. No involucre, but the head subtended by
a few folliage leaves .... 16. Psilocarphus p. 139
aa. Pappus present.
c. Pappus of 2-(4) terminal awns or horns.
d. Inner tegules connate, about twice
as long as the outer.. 31. Thelesperma p. 160
dd. Free and petaloid or shorter than
thesouter We. .scacsssescene Gon, Didenar spare
cc. Pappus of capillary bristles.
e. Leaves becoming alternate in the
inflorescence ......ecscses 3. Brickelinmpe eee
ee. All opposite.
£. Florets yellow?...ssc.. 47. Arnicamp. 275
ff. White or purplish ... 2. Eupatorium p. 94

Group III-C
Various unusual types.

a. Heads globose, terminal on very long pe-
duncles ...... Foon AGS seevecsiecsees SOs. Echinops p.toe
aa. More numerous and in racemes or panicles.
b. Inflorescence a raceme; herbaceous plants.
c. With a taproot or corm; heads

jemegole) Bogoggac eer csseisceesne 8. LLAtELS peo
cc. Stoloniferous; heads white ......
Sb ocdoaDGDoOaC Mekiclelitielel lif. ANntennarlemp amos

bb. Heads nennentera: shrubby at base.
d. Leaves opposite, heads white ....

S @ okeuelis oe iaeie-eieks eereresreee@ave S36 Brickelia Peo 96
dd. Leaves alternate and narrow, heads
yellow.

e. Leaves spinulose-serrate ....
elelois elelsisleleicicieiaician io) Machaerantheral palao
ee. Entire ....0-+.0. 10. Chrysothamnus p. 108

Group III-D

Not spiny; heads discoid; leaves alternate; papus
of capillary bristles.

Bye ConclVast yellow? teleicl- «1s cle ce vlsiclslaleie Oo) OENCC1O! peLoo
aa. Not yellow, but white, pink, etc.
b. Pappus plumose .....+....+.. 52. Saussurea p. 194
bb. Barbellate or glabrous.
c. Stem leaves reduced to large foliaceous
petioles; normal leaves large and all
baeeal., .ytelerey on. calsicsle’s ee eKtGndRetasiteslp. G75
cc. Stem with normal leaves.
d. Herbage copiously white-wooly;
leaves entire.

COMPOSITAE 90

1972 Boivin, Flora of Prairie Provinces

e. Flowers all perfect; taprooted
herbs ....-+seecee 19. Gnaphalium p.148
ee. Some flowers staminate only;
mostly with stolons or a rhizome.
£f. Flowers dioecious ......
eocsecseeees 17. Antennaria p.139
ff. Central florets staminate,
the peripheral pistillate ..
ecoccccscecee 18. Anaphalis p.147
dd. Pubescence different; leaves mostly
dentate.
g- Pappus, like the corollas,
purplish ....-cece.s. 1. Vernonia »p. 94
gg- Pappus white to tawny.

h. Tegules narrow, numerous and
long-tapered, all of the same
length or a few of the outer
ones many times shorter ....
ee OL 15. Brigeron p.127

hh. Tegules uneven, the outer much
longer or more commonly imbri-
cated in gradually shorter
rings.

i. Tegule tips flabellately
enlarged, or flabellately
lobed to pectinatifid, or
flabellately spiny .eccee
eccvecees 06. Centaurea p200

ii. Tegule tips entire and not
enlarged.

j. Fibrous-rooted and
usually stoloniferous..
esevnvesie'eis) Loe Aster pa09

jj. Tatrooted; not stolo-
NLESCTOUGS! ~ clelc sis » sels ees
214. Machaeranthera p26

Group III-B
As III-D but the pappus lacking or chaffy.

B. Mein leaves opposite! ssve.cessccsccscese Zl. Iva pla9
aa. Alternate.
b. Leaves entire to coarsely toothed.
c. Leaves large, deltoid-ovate ...
So. seew Teel Sie. ws 6550208. Adenocauion: pao
cc. Smaller and narrower, elliptic to linear.
d. Involucrakt bracts in one series and
uniform in length -o-e...- 34. Madia pJ62
dd. In many series and the outer ones
successively shorter.

91 COMPOSITAE

Pi YT OP LiO+G: Tok Vol. 23, Hoss

e. Receptacle bristly; heads few,
large and mostly terminating
elongated branches......-ceecc
jie Soba eiee 56. Centauzesiia. 2a5
ee. Receptacle naked; heads many
and small, on pedicels often
shorter than the heads.
f. Leaves closely and finely
crenate..43. Chrysanthemum p. 169
ff. Entire or with a few rather
coarse and remote teeth or
lobes...oece- 45. Artemisia p. 171
bb. More dissected, narrowly pinnatifid to
tripinnatifid.
g. Pappus of scales + 1 mm long; leaves
narrowly pectinatipartite .....-ee-c.

sis aisle aivinte Ste'eleta sltdleid’ste 35. Hymenopappun ae =m
gg. No pappus.
h. Inflorescence paniculate to
SPACULOEM geicielcistels ele) CO. AGteMaswaN pl. Lt
hh. Inflorescence not so elongate but
corymbiform.
i, Annual; receptacle conical ..o..
dia ldttbcliiicle sisleces 426 Matricaniaspewem
ii. Perennial; receptacle flattish
to slightly raised towards the
maddie .....ce,e. 44. Tanacetum p. 270

Group IV
Flowers all ligulate. Stem and leaves commonly with
a milky juice.

ln MSeENIIES) Gogond0c aiahehele el) <1 elle elle) «ican elsiiel slefeletereve/one COULD mealies
aa. Stem + leafy.
b. Pappus of minute scales.
c. Flowers blue ........... 57. Cichorium p. 201
Cc. NELUOW ‘w.e's-c OAT OC OL en SIS . Lapsana p. 202
bb. Pappus of bristles.
ds) Bristiles plumoSeeces..%.+-.ccecoes Group LV=s
dd. Merely barbellate or scabrous .... Group IV-C

Group IV-A
Scapose herbs. Head solitary and borne directly
on the rhizome, the peduncle naked or merely with small
bracts, not leafy. Florets all ligulate.

a. Achene beakless,
b. Leaves tomentose-ciliate; achene about 8 mm
Wome teenies «e's ee secccecsee 59, Microseris p. 202

COMPOSITAE 92

1972 Boivin, Flora of Prairie Provinces 9

bb. Leaves not ciliate or at least not tomentose-
ciliate; achene much smaller.72. Hieracium p. 214
aa. Pappus borne at the end of a long beak.
c. Pappus bristles plumose; receptacle
CHELLY» 6: <0, 5:uinivieyeianeravte deran’s Gl, Hypeochaeris p. 203
cc. Bristles smooth or slightly scabrous;
receptacle not chaffy.
d. Achene becoming spinulose-muricate
towards the tip cceacoe. 65. Taraxacum p. 205
dd. Achene uniformly tuberculate on
the ridges .....e-cececcace 59. Agoseris p. 210

Group IV-B
Stem leafy. Otherwise as IV-A. Pappus of
branched (i.e. plumose) bristles.

a. Ligules pink .......seeeee0e0 63. Stephanomeria p. 204
aa. Yellow or sometimes orange to deep red.
b. Involucre not calyculate, the tegules
isomegueth ...-..e+sesee+eeee00 64. Tragopogon p. 204
bb. Tegules dissimilar, the outer ones
many times shorter or much broader.
c. Leaves all basal or near basal ..

= oct wcvcccccccccocce Ol. Hypochaeris p. 203
cc. Stem leafy to the inflorescence ..
Se ee et se 62..Picris p. 203
Group IV-C

Pappus of simple and smooth or scabrous bristles.

a@. Pappus double, the outer 5 units small and scale-
like, the inner bristle-like ...... 60. Krigia p. 203
aa. Pappus of bristles only.
b. Achene compressed, at least twice as
broad as thick.
ce Seed without beak or disk ....
Siota/ stsice ntaite leks Velie rn ekekatafataya « wefan Omm COONCHUSE Dist 206
cc. Pappus borne on a@ disk at the
end of a + obvious beak... 67. Lactuca p. 207
bb. Achene terete or polygonal, little
if at all compressed.
d. Ligules yellow or orange to deep red.
e. Herbage long=pilose at least in
PACE: 6 afisn catherines aay EOLA, Das oot
ee. Glabrous or the pubescence very
short or farinose,.
f. Stem leafy towards the base
only ...cee+++ee 59. Microseris p. 202
ff. Leafy throughout or at least
in the upper part.. 70. Crepis p. 211

93 COMPOSITAE

10 Peg OG Dk Vol. 23, no. 1

dd. White or creamy or pink.
g. Florets 5, pink ... 68. Lygodesmia p. 209
gg. More numerous and usually white.
h. Inflorescence a panicle or
branched raceme .oceeccoeces
Sib ee cate eldveeee 71 sPrenanthss pe aie

hh. Heads corymbose ...---cecee
see eee te eT! .WHieracmim pene

1. VERNONIA Schreber
IRON-WEED
Style branches filiform, short-hirsute on the
outer side only.

1. V. fasciculata Mx. var. corymbosa (Schwein.)
Schub. -- Numerous discoid heads with purplish florets,
pappus and tegule tips. Tufted virgate herb. Lanceo-
late leaves alternate, serrate. Heads numerous, mostly
corymbose. Tegules conspicuously imbricate. Second
half of summer. Along Rat and Red rivers at Otterburne
and Morris; rare. --scMan, US.

Doubtfully reported from "Weyburn Prairie, Sask."
by Rydberg 1932, Russell 1937 & 1944 and Breitung 1957
on the basis of a collection by Sanson (NY). We share
Breitung's doubt and consider that the locality or the
label is almost surely incorrect. Not only is this spe-
cimen disjunct by 300 miles to the nearest locality of
the species, but it has never been confirmed and it is
also irregular on a phytogeographical basis as it belongs
to var. fasciculata and not to var. corymbosa as it could
be expected in the northwest corner of the range of the
Species. For another doubtful Weyburn report see under
Desmodium canadense in Part IV.

2. EUPATORIUM THOROUGHWORT
Style branches filiform like the above, but finely
puberulent all around. Leaves opposite or verticillate.

a. Leaves verticillate ..........c2e2-+. 1. Be purpureum

6a. Opposite eos ee ee ee ee ee eee ese orere oe 2. E. perfoliatum
1. E. purpureum L. var. maculat (L.) Darl. (E.
maculatum L., var. Bruneri (Gray eit., var. foliosum

Fern.) -- Joe-Pye-Weed -- Tall herb with large verticil-
late leaves. Leaves lanceolate, serrate. Heads discoid,
pinkish purple, in a terminal corymb., Second half of
summer. Marshy and semi-open spots in galerie-forests.
-- NF-SPM, NS-S, BC, US -- F. Faxonii (Fern.) Boivin --
Flowers white. Local: Nipawin. -- NF, O, S, (US) -- F.
tegulosum Boivin -- Floral parts modified into so many
scales. Local anomaly: Otterburne -- se Man.

VERNONIA 94

1972 Boivin, Flora of Prairie Provinces 11

F. Faxonii (Fern.) stat. n., EB. maculatum L. f.
Paxonii Fern., Rhodora 97: 195. 1945,

F. tegulos (Boivin) stat. n., E. maculatum L.
ff. tegulosum Eoterice Svensk Bot. Tidskr. 53: 431. 1959.

Plants with more densely pubescent herbage have
been called var. Bruneri, but this may be only an eco-
logical variant. Its greater frequency in the western
part of the range is probably climatically conditioned.

The group of E. maculatum has had a checkered his-
tory and old records are not always readily interpreted.
It started with Linnaeus as a group of three species:
maculatum, purpureum and trifoliatum. By the time of
the 1908 edition of the Gray's Manual these had been re-
duced to a single species and three varieties. But the
pendulum has swung back and Barratt's 1841 classifica-
tion was revived after being revised as to taxonomy by
Wiegand 1920, amended as to nomenclature by Wiegand and
Weatherby 1937. The proposed arrangements finally found
their way into major eastern manuals with 4 species and
one variety keyed out and described. We have always
found this too complex arrangement unsatisfactory and
others seem to have had the same trouble, judging from
the many erratic.identifications in the various herba-
ria visited. There is a general lack of morphological
cleavage between the taxa and too much sympatry when the
diagnostic criteria are applied rigorously. We have
been able to recognize only two weak geographical varie-
ties as follows.

The more eastern var. purpureum (=_E. trifoliatum
L.) has larger leaves, ovate-lanceolate, the main ones
4-8 cm wide and often 2 dm long or more; stem solid and
purplish or speckled in purple; heads commonly pinkish,
usually with 5-7 flowers each.

The widespread var. maculatum (= E. dubium W.; E.
fistulosum Barratt) is more deeply coloured and its
leaves are smaller by about half. Stem fistulose or
solid, greene or glaucous to purplish or purple-speckled.
Main leaves mostly 2-4 cm wide and usually around 1 dm
long. Heads darker in colour and bigger, usually with
8-12-(15) florets each.

Reports of var. maculatum from west of our area may
possibly relate to introduced plants only. At any rate
the only two B.C. collections we know, Lulu Island (DAO)
and Chilliwack (CAN, DAO), seem to represent introduc-
tions.

2. BE, perfoliatum L. -- Boneset, Thoroughwort
(Herbe & Souder) ~~ eaves triangular-lanceolate and
connate across their bases into perfoliate pairs. Lea-
ves broadest at the base. Heads small, discoid and
white in rounded terminal corymbs. Mid summer. Shores.
-- NS-sMan, US.

95 EUPATORIUM

12 PHY TOL, 0'G PA Vol. 23, HOseL

Eupatorium rugosum Houtt. (EB. urticifolium Rei-
chard) was doubtfully reported from "Weyburn prairie,
Sask." by Rydberg 1932, Russell 1937 & 1944 and Brei-
tung 1957. The comment under Vernonia fasciculata
seems equally applicable here. The habitat is quite
wrong for an eastern forest species that would reach
its limit of range some 400 miles further east. Nor
is it likely that the specimens were collected in Ma-
nitoba as speculated by Breitung.

3. BRICKELIA Ell.

Like Eupatorium, with very long filiform styles,
etc. Tegules strongly imbricate and strongly costate.
Pappus bristles minutely barbellate.

1. B. grandiflora (Hooker) Nutt. -- Large white
discoid heads somewhat drooping in terminal clusters.
Leaves opposite, petiolate, deltoid-ovate or cordate,
serrate, becoming alternate in the inflorescence. Te-
gules deep green, costate in white, the inner whitish
at tip, the outer long caudate. Mid to late summer.
Wet rocky places in the mountains: Waterton. -- swAlta,
wUS, (CA).

The first canadian collection was by Dawson in
1874 at the Kootenay Pass (MIMG) while surveying the
49th parallel. A provincially ambiguous location,it
has led to the species being reported from both Alberta
and B.C., but more modern collections would restrict its
Canadian range to Waterton Park only. Indeed modern
collecting indicates that a fair proportion of Dawson's
specimens labelled Kootenay Pass probably came from
within Waterton Park as defined to-day.

4. LIATRIS Schreber BLAZING STAR
Like the last two, but the pappus plumose and
the tegules not striate.

a. Heads hemispheric to subglobose... 1. L. ligulistylis
aa. Much narrower, cylindric ........e0.. 2. L. punctata

1. L. ligulistylis (Nelson) K. Schuman (L. aspera
AA.; L. scariosa AA.) -- Showy and stiffly erect herb
with a terminal raceme of a few large heads, the termi-
nal one largest. Root a woody irregular and subglobose
corm. Leaves lanceolate, the upper many times smaller.
Heads purplish. Tegules tips concave, squarrose, erose,
purplish and membranous-margined. Mid summer. Frequent
in draws and around groves southward, in sandy woods
northwards. --sMan-Alta, US --F. leucantha Shinners --
Flowers white. Local. --Man-S, (US).

BRICKELIA 96

1972 Boivin, Flora of Prairie Provinces a |

Two sheets of this species at MIMG deserve special
mention. Both were collected in 1855 by Jas. Anderson,
Both specimens look like twin brothers, the way speci-
mens from the same collection do. But they carry wide-
ly different locality data: "Mackenzie River" on one case,
the other: "Found on the plain of Saskatchewan and
North", In view of the known distribution of L. ligulis-
tylis, the locality data of the first specimen would seem
to be erroneous and should presumably be changed to read
like the second.

There are a number of other apparent range exten-
tions among the Anderson specimens, mostly of prairie
plants. But there are too many such extensions and qui-
te a few appear improbable. We therefore have ignored
them all unless confirmed by later collections and the
localities given on his labels are considered to be gene-
rally questionable,

2. L; punctata Hooker var. punctatg -- Taproot with
a globular enlargement at some distance below the surface.
Smaller than the last and the leaves linear and conspi-
cuously scabrous-ciliate. Tegules ciliate, cuspidate.
Mid summer. Frequent and showy on steppes and hillsides.
-- Man-Alta, US -- F. albiflora (Sheldon) Boivin --
Flowers white. Teele ton greene rare: Souris, Rock-
glen --Man-S, US.

F. albiflora (Sheldon) stat. n., Laciniaria punc-
tata (Hooker) Ktze. £. albiflora Sheldon, Quart. Bull.
Univ. Minn. 1: 26. 1892; L. punctata Hooker f. albiflora
Sheldon. nomen invalidum ex Scoggan, Fl. Man. 514. 1957;
L. punctata Hooker. f. alba Horr & McGregor, Trans. Kan-
sas Ac. Sc. 54: 216, 1951. The only U.S. collection seen
of the white-flowered form was from Les Genoux, Montana
(DAO).

Var. punctata is 0,.5-4.0 dm high and its leaves are
3-5 mm wide. To the southeast it gives way to a var.
nebraskana Gaiser, taller, 4-8 dm high, and with narrower
and eciliate or barely ciliate leaves, 2-3 mm wide.

5S. GRINDELIA W. GUMWE ED
Pappus of 2 or more deciduous awns. Heads large,
radiate in yellow. Otherwise much as in Aster.

1. G. squarrosa (Pursh) Dunal var. squarrosa (var.
quasiperennis Lunell, var. serrulata (Rydb.) Stey.; G.
perennis Rydb.) -- Gumweed, Gumplant (Epinette de prairie)
-- Very sticky heads, tangling readily by their very
strongly squarrose tegules. Leaves serrate, resinous-
punctate in darker green. Involucre very resinous. Mid
to late summer. Somewhat saline soils, especially if dis-
turbed. -- sMack, (NF), QBC, US.

Middle and upper stem leaves ovate to oblong-lan-
ceolate and auriculate-clasping. The more western var.

97 LIATRIS

14 1s; Ge: ag at 6 a OO ee a Vol. 23, no. 1

integrifolia (Nutt.) stat. n., G. nana Nutt. var. integri-
olia Nutt., Trans. Am. Phil. Soc. 7: 314. 1840, has nar-
rower leaves, at least the middle and lower ones oblan-
ceolate, long cuneate at base, sessile and neither auri-
culate nor clasping.

Other segregates listed as synonyms have the range
of the species and appear to be part of the normal range
of variation of the typical phase.

6. GUTIERREZIA Lag.

Similar to Solidago, though the pappus is of ci-
liate scales.

1. G. Sarothree (Pursh) Britton & Rusby (G. diver-
sifolia Greene) -- Broom-Weed -rLike a diminutive Soli-
dago graminifolia. Tufted, semi-shrubby perennial with
a taproot. Leaves linear, very narrow, scabrous. Heads
somewhat glutinous. Late summer. Mainly eroded step-
pes and badlands. --Man-Alta, US, (CA).

7. CHRYSOPSIS Nutt. GOLDEN ASTER

Like Solidago, with dimegueth pappus-bristles, the
outer many times shorter.

ie Cc; yillosa (Pursh) Nutt. (C. angustifolia Rydb.;
Cc. hirsutissima Greene; C. hispida (Hooker) Nutt.) --
Like a yellow-flowered Aster. Tufted perennial, densely
and stiffly pubescent throughout. Stem very long hirsu-
te. Leaves oblanceolate, entire, very long ciliate to-
wards the base. Tegules purple tipped. Summer. Common
on light or disturbed soils. --Man-BC, US.

A rather variable type, often subdivided into ma-
ny varieties or species. While we would hesitate to be
too positive about extralimital segregates, within our
area this species appears to form a single population,
hence the consolidation.

8. SOLIDAGO L. GOLDENROD
Like Aster, but the flowers yellow (white in two
species). Heads rather small and the ligules short.

a. Stem leaves entire and long-linear; inflorescence
corymbiform.
b. Ligules white............... 13. S. ptarmicoides

bb. Yellow; plants caies
co. Leaves straight and flat, uniform
in length .............. 14. S. graminifolia
cc. Conduplicate, falcate, larger and
the upper many times shorter ....
Cet e ee ce ecw oer ec eceeeeee 12. S. Riddellii
aa. Stem leaves serrate.

CHRYSOPSIS 98

1972 Boivin, Flora of Prairie Provinces 15

d. Leaves fairly uniform in length, the upper
at least half as long as the lower.
e. Leaves obovate to broadly oblanceolate,
typically oblong........+-e-ecece 8. S. mollis
ee. Rather narrowly lanceolate.
£. Stem pubescent ......... 9. S. canadensis
ff. Glabrous except in the inflores-
CONCE issn esoccervesscee LO. Ss gigantea
dd. Upper stem leaves gradually shorter...... Group A

Group A
Leaves serrate and gradually shorter upwards,
the lower ones at least twice as long as the upper.
a. Leaves densely puberulent on both faces.
b. Heads in @ rounded corymb.......... ll. S. rigida
bb. In a panicle or a thyrse.
c. Heads in an elongated thyrse .. 1. S. bicolor
cc. In a panicle. z
d. Panicle + secund; middle leaves
less than 1 cm wide and narrowly
oblanceolate ...+.eseeees 7e S. nemoralis
dd. Not secund; middle stem leaves
broader and tending to elliptic ..
Secs cccecccoscccosceucesesee§ 8. S. mollis
aa. Glabrous or merely ciliate.
e. Inflorescence broadly pyramidal, over
ERMAN WAS ialsladais. fee olale elon cveretetsler else ates O's
ee. Narrower and often elongate.
f. Lower leaves long ciliate towards
the base; heads rather few and
LATGS ccvccccccsccceesss 4 S. multiradiata
ff. Leaves not ciliate or merely finely
scabrous-margined.
g.- Achene pubescent.
h. Tufted; heads glutinous ..

- juncea

|n

eee e reese cece ce ceeee So Se Spathulata
hh. Stoloniferous; not gluti-
NOUS e.ceeeeceeree De Se missouriensis

gg. Achenes glabrous.
i. Lower leaves at least 2 cm
wide and the petioles somewhat
sheathing at base.... 4. S. uliginosa
ii. Usually much narrower and
tapered to a narrow base ..

ere ee cescccccecce Oe Se missouriensis
1. S- kisoler L. var. bicolor. -- Silverrod, Sil-

verweed -- Herbage hispid throughout. Perennial from
a short rhizome. Inflorescence an elongated thyrse,
cylindric or rarely narrowly paniculate. Ligules white

99 SOLIDAGO

16 Pe yel CULO 1g TA Vol, 235 maser

Tubular flowers tending to yellowish. Mid summer.

Dry, semi-open places. --NS-Man, US -- var. concolor

T. & G. (S. hispida Muhl., var. lanata (Hooker) Fern.)
-- Ligules yellow when fresh, often fading or drying
white. Green midnerve of the tegules tending to be
narrower, narrowly oblanceolate, and more sharply con-
trasted with the larger paler and whitish margins. Dry
open woods, frequent. --NF, NS, NB-S, US.

Var. concolor is commonly treated as a species
(=S. hispida) but the difference amounts to only this
single colour character, hence the rank of variety adop-
ted here. The colour difference is quite obvious in
the ligules when fresh, but there may be some fading in
drying. In the tegules the colour is more variable and
the difference is not susceptible of being sharply defi-
ned.

Var. lanata is an extreme of pubescence of spora-=
dic occurrence.

ae Ss. ultiradia Aiton (var. scopulorum Gray S.
scopulorum (Gray) Nelson) -- Heads rather large, the in-=
volucre (6)-7-(8) mm high, and not very numerous, mostly
10-20 per inflorescence. Puberulent in the inflorencen-=
ce and the lower leaves long ciliate towards the base,
otherwise glabrous. Stem leaves, especially the upper,
tending to be oblong-oblanceolate. Thyrse rather short,
sometimes merely round-corymbose, but usually a little
bit taller and the lowest branch + drooping when fresh,
tending to be a bit remote. Mid summer. Northern and
montane or alpine meadows. -~(F)-K-Aka, L-NF, NS, NB-
BC, US.

3. S. spathulata DC. var. spathulata (var. nana
(Gray) Cronq., var. neomexicana (Gray) Cronq.; S. decum-
bens Greene, var. oreophila (Rydb.) Fern.; S. oreophila
Rydb.) -- Tegules glutinous, marking the paper in yellow
in drying, remaining t shiny. Tufted, 1-5 dm high. Fi-
nely puberulent on the stem and in the inflorescence.
Basal and lower leaves spathulate-oblanceolate. Inflores-
cence an elongate and cylindric thyrse. Mid summer.
Common prairie species. --Mack-Aka, (NS), NB-O-(Man)-S-
BC, US.

Often subdivided into a series of poorly defined
and commonly intergrading varieties: the more common and
more widely distributed type with narrower and more elon-
gate leaves is var. neomexicana; smaller plants of higher
altitudes are var. nana; here and there, especially in
the mountains and closer to oceans or large bodies of
water, one will find broader-leaved plants which may be
termed var. spathulata westward or ssp. Randii (Porter)
Cronq. in the east.

Around the Great Lakes there is a somewhat more
clearly defined var. Gillmanii (Gray) Cronqg., taller,

SOLIDAGO 100

1972 Boivin, Flora of Prairie Provinces gy

(4)-5-8-(10) dm high; leaves larger, the lower often
up to 3-4 cm wide and more coarsely serrate; heads
somewhat larger.

4. S. uliginosa Nutt. var. yliginosa (S. Purshii
Porter; S. neglecta T. & G.; S. uniligulata (DC.) Por-
ter) --Lower leaves rather large, 1.5-3.0 dm long; in-
cluding the sheathing petiole, and mostly 2-3 cm wide.
Fairly tall, but less than 1m high. Inflorescence elon-
gate, a thyrse or a very narrow panicle. Late summer.
Bogs. -= seK, L-SPM, NS-seMan, US.

In younger and smaller plants the branches are at
first more strongly ascending and the heads not obvious-
ly secund. This is sometimes distinguished as A. Purshii.

Around the Great Lakes one finds smaller plants
with narrower leaves 1 cm wide or less. These may be
segregated varietally as var. jejunifolia (Steele)
Boivin.

5S. S. missouriensis Nutt. var. missguriensis --
Long stoloniferous and common prairie species usually
2-4 dm high. Glabrous or somewhat puberulent in the
inflorescence. Leaves narrowly oblanceolate, less than
1.0 dm long, all or mostly less than 1 cm wide, the lower
often scabrous-margined. Inflorescence very variable,
short and usually less than 5 cm wides heads not stron-
gly secund. Involucre 3-4 mm high. Mid summer. Common
in prairies and open places. --wO-sBC, US -- Var. extraria
Gray -- Heads larger, the involucre 4-5 mm high. Rocky
ridges in the mountains and foothill prairies. -- swAlta,
wUS -- Var. fasciculata Holz. (S. glaberrima Martens) --
Larger throughout, the inflorescence up to 1 dm wide and
broadly pyramidal. Lower leaves up to 1.5 dm long.

Plant commonly 4-5 dm high. Lower branches recurved at
tip and their numerous heads all or mostly borne on the
upper side only. More common in lighter or disturbed
soils. -- wO-Alta, US.

Our 3 varieties intergrade freely and the last is
somewhat transitional to S. juncea.

6. S. juncea Aiton -- Similar to the last variety,
but larger throughout. Loosely tufted. Basal leaves
1.5-3.0 dm long, mostly 2-3 cm wide and forming large and
conspicuous rosettes. Glabrous throughout. Inflorescence
broadly puramidal, commonly 1.5-2.0 dm wide, the lower
branches elongate, recurved and bearing numerous heads
turned upwards. Mid summer. Dry semi-open places: San-
dilands. -- NS-seMan, US.

7. S- nemoralis Aiton (var. decemflora (DC.) Fern.;
S. pulcherrima Nelson) -- Inflorescence usually stron-
gly secund and mostly facing southward on sunny days,
with the tips strongly arching in the opposite direction.
Herbage densely puberulent throughout and somewhat sca-

101 SOLIDAGO

18 Poly Yer O1L/O0G Tek Vol. 23, mosct

brous on the leaves, Loosely tufted and mostly 2-4 dm
high. Leaves oblanceolate. Mid summer. Hillsides and
dryer prairies. --wNS-eBC, US.

The more western plants are commonly segregated
as a var. decemflora on the basis of narrower and less
dentate leaves, of larger heads and sericeous achenes.
Our specimens do not conform readily to this dichotomy.

8. S- mollis Bartl. -- Stem leaves numerous and
commonly elliptic, varying from narrowly obovate to
broadly oblanceolate and obtuse to rounded at tip.

Upper leaves many times smaller. Lower leaves slightly
shorter than the middle ones but usually deciduous. Com-
monly 3-4 dm high, long stoloniferous, without basal
rosettes but with short sterile shoots. Herbage densely
puberulent and scabrous. Leaves with 3 stronger and
nearly parallel nerves. Inflorencence thyrsiform to
paniculate. Second half of summer. Dryer prairies and
steppes. --swMan-sAlta, US.

9. S. ganadensis L. var. canadensis (var. fallax
(Fern.) Beaudry, var. salebrosa (Piper) Jones; S. lepida
AA., var. elongata (Nutt.) Fern.; var. fallax Fern.) --
Conspicuous virgate herb, often + 1 m high, with numerous
small yellow heads. Long stoloniferous and without ro-
sette but producing shorter sterile stems. Stem finely
pubescent at least in the upper half, the hairs incurved.
Leaves numerous, fairly uniform in length, mostly around
1 cm wide, narrowly lanceolate, with 3 much stronger ner-=
ves, + puberulent below, glabrous above to slightly pube-
rulent, especially along the nerves. Inflorescence pyra-
midal. Mid summer. Open woods and moist meadows. <=-
Mack-(Y-Aka), L-NF, NS-BC, US -- Var. gilvocanescens, Rydb.
(S. dumetorum Lunell; S. gilvocanescens (Rydb.) Smyth}

S. Lunellii Rydb.; S. pruinosa Greene) -- More pubescent,
especially the leaves densely puberulent and scabrous
above. Leaves often broader. Common in prairies. --
swMack, L-NF, NS, Q-BC, US. -- Var. scabra (Muhl.) T. &
G. (S. altissima L.) -- Stem densely pubescent with
longer, crip-flexuous, and + spreading hairs, usually
around 0.5 mm long. Leaves often similarly pilose, es-
pecially on the lower face. -- wNB-swQ-seMan, US.

It has been customary to restrict the application
of S. canadensis to the plants with smaller heads, the
involucre 2-3 mm high, its tegules mostly + 0.3 mm wide,
and narrowly lanceolate leaves + 1 cm wide. Plants with
larger heads and leaves have been distinguished as var.
salebrosa, or var. fallax, or S. lepida, but the distinc-
tion does not appear to be a significant one in our area
and we are dubious about its value elsewhere.

If the distinction is accepted, var. salebrosa may
possibly be the correct name for the coarser plant, but,

SOLIDAGO 102

1972 Boivin, Flora of Prairie Provinces 19

as pointed out by Cronquist 1955, S. serotina Aiton
var. minor Hooker 1834 could be an earlier and valid
name. This name needs checking as to its exacts mea-
ning; unfortunately its type could not be located at
K in 1969. A more recent var. fallax (Fern.) Beaudry,
Nat. Can. 95; 37, 1968 has also been proposed for what
appears to be essentially the same entity.

The use of S. lepida for our slightly larger-
headed plants is erroneous. S. lepida DC. is a Paci-
fic Coast species with a very narrow panicle of much
fewer and much larger heads, the involucre 5-7 mm high.

9X. S, canadensis var. gilvocanescens X gigantea
var. serotina -- With the broadly Tanceolate leaves,
about 2 cm wide, of either parent. Intermediate as to
pubescence. Stem rather coarse as in S. gigantea, but
puberulent towards the top as in S. canadensis. Some
leaves glabrous below, some puberulent on the main ner-
ves. Upper leaf surfaces partly or sparsely scabrous-
puberulent. Heads rather large as in S. gigantea, the
involucre 4mm high. Pincher Creek. --swAlta, (US).

10. S. gigantes Aiton var. ggrotina (Aiton) Cronq,
(var. leiophylla Fern.; S. serotina Aiton) -- Closely
resembling the last but larger and less pubescent. Stem
coarser, mostly 1.0-1.5 m high. Glabrous except the
inflorescence. Leaves rather lanceolate and commonly
2-3 cm wide, glabrous on both faces, scaberulous at mar-
gin. Heads tending to be large, the involucre commonly
3.5-4.0 mm high, the tegules often 0.5 mm wide or larger.
Second half of summer. Mostly near watercourses and
shores. --Mack, NS-eBC, US.

The more eastern var. gigantea is pubescent along
the main nerves on the lower leaf surfaces, usually sca-
brous above. Old reports of the latter variety from
Western North America are apparently to be discounted
as pointed out by Cronquist 1955. The exact basis of
Macoun's 1884 western report could not be readily deter-
mined; the original specimens are presumed to have been
revised to other taxa. All western Canadian material
found under var. gigantea at CAN, DAO and MT has been re-
vised, mostly to var. serotina, but some also to S. cana-
densis and S. missouriensis.

ll. §, gigida L. var. humilis Porter (var. canes-
cens (Rydb.) Breitung; S. parvirigida Beaudry; Oligoneu-
ron canescens Rydb.; O. rigidum AA.) -- Leaves conspi-
cuously and pinnately veined, the lower leaves many times
longer, oblong-lanceolate and long petiolate. Tufted,
2-7 dm high, with conspicuous rosettes 1.5-3.0 dm high.
Herbage densely puberulent and somewhat scabrous. Inflo-
rescence 3-10 cm wide, round-corymbose, its branches almost
always ebracteate. Mid summer. Common on steppes. --
O-Alta, US.

103 SOLIDAGO

20 PBeYeT Or Lc0eG TA Vol. 23, "ise

Our variety is generally smaller and with fewer
leaves, 5-23 per plant, the basal ones shorter. The
more eastern var. rigida is (6) -10-(17) dm high; lower
and basal leaves + lanceolate, 2.5-5.0 dm long inclu-
ding the petiole; stem leaves 20-30 per plant; inflo-
rescence mostly larger, 7-16 cm wide, its lower branches
more or less bracteolate in their lower half. We are
keeping the two taxa at varietal rank because there is
some degree of morphological overlap. Both varieties
were reported (as species) from our area by Rydberg 1932
and Russell 1937, 1944 and 1954; but all western speci-=
mens examined, including duplicates of collection by
Fraser (DAO) and by Russell (DAO), proved to be refera-=
ble to var. humilis.

It was recently discovered that some specimens of
var. rigida were tetraploid (2n=36) while others of var.
humilis were diploid (2n=18). On that basis var. humi-
lis was promptly elevated to specific rank as S. parviri-
gida. Such a procedure is considered to be unsound on
two counts.

First it is an attempt to express the concepts of
one speciality (genetics) in the terminology of another
(taxonomy). This abuse of terminology can only create con-=
fusion for both specialities. That the geneticists have
not yet provided themselves with their own naming proce-=
dures, as some other specialities have done (e.g. culti-
var, forma specialis), may be a handicap to the geneti-
cists, but it is not a justification to take over the
terminology of taxonomy for genetic purposes.

The attempt at a take-over has gone quite far in
some cases. Witness the folowing quote from a geneti-
cist: "Plants which belong to different levels of ploi-
dy are best considered, from a theoretical standpoint,
as different species, even if they are morphologically
identical, because the difference in the number of chro-
mosomes constitutes a strong enough reproductive barrier
to keep the populations separate under conditions of sym-
patry". -- Can. Journ. Gen. Cyt. 5: 167. 1963. -- Obvious-=
ly the author of the quote would define the species in
essentially chromosomian terms.

Second, it is based on a confusion between cause
and effect. A species is first and foremost a disconti-
nuous morphological unit. And the discontinuity implies
the existence of a barrier to hybridization. Without
such a barrier normal reproductive events would rapidly
obliterate the discontinuity, and any taxonomic distinc-
tion would become impossible. However, the barrier itself,
be it geographical or genetic or other, is not a taxono-
mic character in its own right, it is only the mechanism

SOLIDAGO 104

1972 Boivin, Flora of Prairie Provinces 21

that makes a morphological discontinuity eventually pos-
sible. And from this discontinuity arises the taxonomic

character. The discovery of the exact nature of a barrier,
or the discovery of an additional barrier -- a second le-
vel of ploidy in this case -- represents progress in our
understanding of a taxon, but it is not the discovery of

a new character and is not in itself a justification for
changing the rank of said taxon. See also Boivin 1960.

Further it must be pointed out that in this parti-
cular case the ploidy levels exhibit only partial concor-
dance with the morphology. Some specimens apparently
quite typical of var. rigida proved to be now tetraploid
(Beaudry 57-14-1 from Bloomfield, Mich.) now diploid
(Beaudry 57-472 from Manchester, Tenn.). It is not un-
common to find more than one ploidy level within a spe-
cies without corresponding morphological differentiation,
or with only a weak differentiation, as in this case.

11X. S. Maheyxii Boivin -- Hybrid with S. Riddellii
and generally similar to the latter, but the leaves are
densely scabrous-puberulent and not quite so elongate.
Inflorescence very broadly paniculate or round-corymbose.
Rare: Kleefeld. --seMan.

12. §. Riddeldi Frank -- Leaves conduplicate and
falcate. Stoloniferous with rosettes up to 6 dm high.
Herbage glabrous except for the very scabrous leaf mar-
gins. Inflorescence broad-corymbose. Late summer. On
chernozems, rare: Kleefeld, Sainte-Geneviéve. --O-seMan,
US.

12X. S. Bernardii Boivin -- Hybrid with S. ptarmi-
coides and resembling mainly the latter, but the ligules
at first pale yellow, eventually turning white. Leaves
mostly about 5 mm wide and the heads not quite so large
as the above. Rare: Kleefeld. --seMan, (US).

Hybr. n., Verosimiliter hybridus S. ptarmicoides
X S. Riddellii. Ad. S. ptarmicoidem vergens sed flori-
bus minoribus luteolis et albescentibus. Folia tantum
latiora. Involucrum 6-7 mm alt. Type: Boivin, Bernard &
Perron 12942, Kleefeld, 1k milles au sud-est, prairie,

16 aofit 1958 (DAO).

13. S. ptarmicoides, (Nees) Boivin (Aster ptarmi-
coides (Nees) T. & G.; Unamia alba (Nutt.) Rydb.) --
Flowers white and the heads rather large in a flat co-
rymb, resembling an Aster. Mostly + 3 dm high and sca-
brous. Leaves very variable in length, very narrow, less
than 5 mm wide, very scabrous at least at margin. Mid
summer. Frequent on sandier soils in the parkland zone.
-- NB-ecS, US.

Stat. n. Doellingeria ptarmicoides Nees, Gen. Sp.
Ast. 183, 1832; Chrysopsis alba Nutt., Gen. Z: 152.

1818, nec Solidago alba Miller. The frequency and va-

105 SOLIDAGO

22 RYT Orb Ore DA Vol. 23, no. 1

riety of hybrids between this species and various other
Solidago seems to be a clear indication that its rela-
tionships lie with the latter genus rather than with
Aster. It is an atypical species in any case.

13X. S. lutescens, (Lindley) Boivin (Aster ptarmi-
coides (uaeet T. & G. var. lutescens (Lindley)Gray) -
Hybrid with S. rigida var. humilis. Mainly of the habit
of S. ptarmicoides, but the heads smaller, the ligules
yellowish, the leaves broader and the upper ones not so
much reduced, about half as long as the lower. Herbage
densely scabrous-puberulent. Rare: Stoney Mountain,
Bird's Hill, Brandon, Kleefeld, Virden, Red Deer, Indian
Head, Beenuaeds --Man-S, US.

Siceities Mle, Diplopappus lutescens Lindley ex DC.,
Prodr. 5: 278. 1836.

14. S. graminifelig (L.) Sal. var. graminifolia --
Povertywe eed -- Leaves long linear and isomegueth, mostly
15-20 times as long as wide and usually 5 mm wide or less.
Inflorescence a single, terminal, flat corymb of numerous
small heads. Second half of summer. Common on shores and
wet places. --NF, NS-seMan, BC, US -- Var. major (Mx.)
Fern. (var. camporum AA.) -- Leaves broader and usually
shorter, mostly 8-10 times as long as wide and commonly
over 5 mm wide. -- Mack, (NF), Q-Alta-(BC), US.

S. occidentalis is also reported for our area by
Rydberg 1917, 1932, Eastham 1947, Cronquist 1955 and,
doubtfully, Been 1966. Becoeee to substantiate nie
report have been unsuccessful as no relevant specimen

could be located at ALTA, Calgary, CAN, DAO, GH, NY, UBC,
WIU, etc.

9. HAPLOPAPPUS Endl.

Heads yellow as in Solidago, but larger as in Aster.
Bristles somewhat unequal.

a. Leaves mas ae dissected, pinnatipartite ....
Chee ee ee eee ee ee cee eee ee ee ee eeees 1. H. spinulosus
aa. Entire to serrate.

b. Leaves serrate; mostly few=headed ....ce.

Prete eee ee ee ee cece ee ee nero eeee So He lanceolatus
bb. Entire; monocephalous.

c. Subscapose, the leaves nearly all

basal, the 1-2 stem leaves much

smaller ........+.+-+see00e. 2. H. armerioides
cc. Stem leaves numerous and not parti-
cule ly smal Were. skectsete ses ece as He Lyallii

1. H; spinulosus (Pursh) DC. var. spinulosus (Aplo-
pappus spinulosus (Pursh) DC.; Sideranthus STE
(Pursh) Sweet) -- Grayish-tomentose leaves pinnatipartite

SOLIDAGO 106

1972 Boivin, Flora of Prairie Provinces 23

to bipinnatipartite, the lobes abruptly contracted into
white, glabrous, spinescent points. Tegules strongly
imbricate, with a dark green and nearly glabrous subter-
minal patch, and a white point like those of the leaf
lobes. Mid summer. Occasional and scattered on step-
pes and eroded hillsides. --swMan-sAlta, US, (CA).

The more southern var. glaberrimus (Rydb.) stat.
n., Sideranthus glaberrimus Rydb., Bull. Torr. Bot.

Club Z7: 621. 1900, has glabrous leaves.

2. H. armerioides (Nutt.) Gray (H. acaulis AA.;
Aploppappus acaulis var. glabratus AA.; Stenotus acau-
lis AA.; S. armerioides Nutt.) -- Tegules with a conspi-
cuous, dark green, deltoid patch at the squarrose tip.
Cushion-forming perennial from a woody taproot. Basal
leaves linear, erect, entire. Stem leaves few and incons-
picuous, the plant scapose in general habit. Stem mono-
cephalous and usually less than 1 dm high. Late spring.
Eroded hillsides and badlands of the Qu'Appelle Valley
and southwest. --S, US.

3. H. lanceolatus, (Hooker) T. & G. var. lanceola-
kus (var. Vaseyi Parry; H. integrifolius AA.; Pyrrocoma
lanceolata (Hooker) Greene) -- Leaves strongly dimegueth,
the basal petiolate, serrate to subentire, lanceolate,
2-5 times longer than the cauline, the latter sessile and
clasping at base. Herbage glabrous or lightly villous.
Stems stiffly erect from a decumbent base. Heads large,
few or solitary. Mid summer. Saline meadows, frequent.
-- S-Alta-(neBC), US -- Var. sublanatus Cody -- H. uni-
florus (Hooker) T. & G.; Pyrrocoma uniflora (Hooker)
Greene) -- Heavily villous-lanate, especially on the in-=
volucre. More northern: basin of the Peace. --sMack,
nAlta.

The name H. uniflorus is generally misapplied in
current floras and monographs. The type of the species
(K) is quite typical of what we are calling above var.
sublanatus. It is restricted to Canada in its distribu-
tion. But H. uniflorus is currently used to designate
another species occurring from Montana to California,
exclusive of Canada. This U.S. entity was first descri-
bed as Homopappus inuloides Nutt., 1840 and is more cor-
rectly named Haplopappus inuloides (Nutt.) T. & G. The
following three varieties are commonly distinguished:

Haplopappus inuloides var. Howellii (Gray) stat.

————— tl al
n., H. Howellii Gray, Syn. Fl. ed. 2, suppl. part iL: 446.
1886.

H. inuloides var. gossypianus (Greene) stat. n., Py-
rocoma gossipiana Greene, Pittonia 3: 23. 1896.

H, inuloides var. linearis (Keck) stat. n., H. uni-

florus ssp. linearis Keck, Aliso a: 1O3. L958.
107 HAPLOPAPPUS

2h ProyY?T OPLiOvG TeA Vol. 23, no. 1

An Alberta report by Rydberg 1917 of Pyrrocoma car-
thamoides Hooker, was repeated by Cronquist 1955 as Ha-
plopappus carthamoides (Hooker) Gray, and querried by
Boivin 1967. No justifying specimen could be located
at GH or NY in 1965 or at WIU in 1969, etc.

4. H. Lyallii Gray -- Glandular peberulent throug-=
hout. Usually less than 1 dm high. Stem leaves oblan-
ceolate, the larger 5-10 mm wide, nearly as large as the
similar basal leaves if any. Mid summer. Alpine slopes
and shale slides. --swAlta-sBC, (nwUS).

10. CHRYSOTHAMNUS Nutt.

Almost identical to Haplopappus, but shrubby and
the heads discoid.

1. C. nauseosus (Pallas) Britton var. nauseosus
(C. frigidus Greene) -- Rabbit Brush -- Low shrub with
numerous annual branches bearing a small terminal group
of discoid heads. Usually less than 4 dm high and the
new branches longer than the woody base. Branches whi-
tish with a thin tomentum, sometimes inconspicuously so.
Tegules usually tomentose. Mid summer. Badlands, uncom-
mon. --sS-seAlta-BC, US -- Var. glabratus (Gray) Cronq.
(var. graveolens (Nutt.) Hall) --More woody and taller,
4-10 dm high and the new shoots usually shorter than the
woody base. Tegules glabrous. Estevan. --seS, BC, US.

11. BOLTONIA L'Hér,

Resembles Aster but the pappus is partly of minute
bristles and partly of 2 or 4 somewhat longer awns.

1. B, asteroides (L.) L'Hér. var. occidentalis
Gray (var. recognita (Fern. & Grisc.) Cronq.; (B. latis-
quama Gray var. recognita Fern. & Grisc.) -- Inflorescen-
ce leaves more or less decurrent. Perennial and stoloni-
ferous. Stem longitudinally striate in light and dark
green. Leaves narrowly lanceolate, entire, scabrous-mar-=
gined. Heads white, resembling Aster. Tegules acute.
Fall. Shores, sometimes weedy; rare or possibly over-=
looked because of lateness. --scMan-scS, US.

In the ozarkian var. latisquama (Gray) Cronq. the
tegules are round-obtuse at tip.

12. TOWNSENDIA Hooker
Closely resembling Aster; pappus bristles gradual-
ly thickened towards the base, almost awn-like. Monoce-
phalous or stemless.

a. Stem mostly 1-2 dm high .....scesssecess 3. Ts. Panny
aa. Stemless.
b. Tegules gradually less densely ciliated
WoNvebsCle) Boanououdo eisiccles cae ceeiceoace el li exGApal

CHRYSOTHAMNUS 108

1972 Boivin, Flora of Prairie Provinces 25

bb. More densely ciliate and ending ina tuft
OF HAGE PP Pe ere Peels Doe ec dccasce 8. Th Bookeart

1. T. excapa (Rich.) Porter (IT. sericea Hooker) --
Usually a cluster of 3-6 large heads overtopped by the
basal leaves, the latter linear and mostly 2-4 mm wide.
Involucre 12-20 mm high, the tegules around 2 mm wide.
Ligules pink. Spring. Sandy hillsides, infrequent. --
swMan-sS-seBC, US.

2. T. Hookeri Beaman (T. sericea AA.) -- Like a re-
duced version of the last. Leaves 1-2 mm wide. Involu-
cre 8-12 mm high. Ligules white. Early summer. Local
on sandy steppes and eroded badlands: Cypress Hills and
Rockies. --Y, swS-Alta, nwUS.

The range was extended to B.C. in Contr. Gray Herb.-
183: 96. 1957, but this may have been a lapsus calami as
the specimen cited came from Fort McLeod, in Alberta.

3. J. Parryi Baton -- A single large head, 3-6 cm
wide, at the end of a rather short stem. Perennial. Stem
solitary and stiffly erect, or a few together. Leaves li-
near on the stem, oblanceolate in the rosette. Ligules
mauve, drying blue. Early summer, Alpine gravel slopes.
--swAlta-seBC, US.

The range of T. spathulata Nutt. was given by Cron-
quist 1955 as extending north to the mountains of Alberta,
while in 1957 in a monograph of the genus, Contr. Gray
Herb. 183: 120-4. 1957, J.H. Beaman restricts its range
to the mountains of Wyoming. We have not yet ascertained
the source of the Alberta report.

13. ASTER L. ASTER
A basic type with radiate heads, the heads, or at
least the ligules, not yellow. Tegules widely varying in
length, the outer successively shorter and imbricated.
Pappus-of fine capillary bristles.

aReeeenvolucre Qlandwhar, jeje wis ici lela icles siete eitetele oo 5 CLOUD LWA
aa. Not glandular but glabrous or hairy.
b. Heads discoid ..........+...-. 24. A. laurentianus
bb. Ligulate.
c. Ligules white, sometimes pink ...... Group B
cc. Mauve or blue to purplish.
d. Leaves gradually dimorphic, the
VOWS, PSELOLMEG eiclnisicts om cise cw elaeeGLOUp iC
dd. Stem leaves all similar, although
the upper sometimes smaller .... Group D

Group A
Involucre abundantly glandular. Longer and non
glandular hairs sometimes also present. Ligules mauve

109 TOWNSENDIA

26 Pen y°T O'L-O'G FA Vol. 23, no. 1

to blue, except A. alpinus.

a. Leaves large, broadly and deeply cordate ......
Hib ob - cco ce cocccccccccscsosccceee L. A. macrophyllus
aa. Broadly lanceolate to linear.
b. Leaves narrowly lanceolate to long li-
near, entire.
c. Monocephalous; lower leaves larger
and oblanceolate .....-eceec--. 25. A. alpinus
cc. Usually with a few heads; all leaves
narrowly linear.
d. Tufted; inflorescence branches
with many small bracts ....ece
wcrc cc cceccccccscacese 24e A. pauciflorus
dd. Stoloniferous; branches bearing
leaves 1-2 cm long ..... 3. A. campestris
bb. Leaves 1-5 cm wide, narrowly to broadly
lanceolate.
e. Leaves conspicuously serrate; mostly
3-5 cm wide ....e..+seeee44 11. A. conspicuus
ee. Narrower and entire to remotely serrate.
f. Leaves short scabrous on both
faces and broadly clasping at the
DASE 2.-cecaccceaecee 4. A. novae-angliae
ff. Long villous below and cuneate
to a narrowly clasping base ...
cece cece ccc are se ceesecce Se A. modestus

Group B
Ligules white, or sometimes light pink, drying
white or not infrequently pale blue. Involucre gland-
less.

a. Heads in a corymb.
b. Leaves long linear, 5 mm wide or less.
c. Upper leaves nearly as long as the
Lower 2.22. -.ee cece ceeeeecc. 16. A. borealis
cc. Lower leaves many times longer ...
chalielsideleWeiielioKe) otelayerel eters »+e Solidago ptarmicoides
bb. Raneeciace and much larger .... 23. A. umbellatus
aa. In a panicle.
d. Tegules thickish, squarrose, spinu-
lose-mucronate; stem uniformly pubes-
COE atassicysic cw. (en layers co'slels oe sececeeee 12. A. ericoides
dd. Tegules thin, straight and not mucro-
nate; stem pubescent in lines,
e€. Main branches widely spreading,
their heads more or less turned
UPWATOAS: <.crelersvavehetercle: eles stam eloe A. lateriflorus

ASTER 110

1972

Boivin, Flora of Prairie Provinces 27

ee. Branches ascending and their heads

not particularly secund.
f. With a thin stem and few heads,
usually less than 15; leaves
entire and rarely over 5 mm wide ..
inline Lieder kedacveenv & OALGS Av borealis
ff. More vigorous plants with more
numerous heads,
ge Outer tegules larger and
longer than the inner eee.
Mae gvdas Saved’ ates 14e-A. hesperius
gg. Involucre imbricate, the
outer tegules somewhat the
shorter.
h. Main stem leaves typically
1-2 dm long and remotely
SELTrate .eeccecoes 15. A. simplex
hh. Shorter and entire; pani-
cle narrow .....-14. A. hesperius

Group C

Ligules blue, varying from mauve to purple, mostly
drying light to deep blue. Leaves dimorphic, the lower
with a poorly to well defined petiole. Involucre not

glandular.

a. Tegules abundantly puberulent on back; herb 4 dm
high or less.
b. Leaves serrate .....cecceeseeeee LO. A. Sibiricus
bb. Leaves entire ............-+.-c.- 20. A. adscendens
aa. Tegules pubescence consisting mostly or enti-
rely of marginal ciliation; plants usually

taller.

c. Achenes glabrous or nearly so.

de

dd.

Leaves thickish and somewhat glaucous,
the lower cuneate to a winged petiole ..
Soe corn evince enccccsscecccsesee Fe A. lacvis
Leaves not fleshy nor glaucous,
abruptly rounded to a wingless or
winged petiole; tegules less imbri-
cated.
e. Lower leaves narrowly ovate; stem
pubescent in lines at least above
the middle ......+..+++.%. A. ciliolatus
ee. Lanceolate; stem glabrous except
in the inflorescence ... 6. A. MacCallae

cc. Achenes pubescent; herbage + pubescent.

£.

Tegules not regularly imbricated, a
few of the outer at least as long and
as large as the inner ones, at least
longer than the middle ones .......

ee seme ative dmesivinivs: Life sn) SUDSDACeLUS
ila Fab ASTER

28 Poleint Ow iOnG: TA Vol. 23, no. 1

ff. More or less imbricated, the outer
being shorter; leaves usually nar-
TOWSM Wade sv esevieccsestecces “L185 A. occidentauia

Group D
Like the last, but the lower leaves more similar
to the upper, and not petiolate, being clasping to ta-
pered at base. Upper leaves usually somewhat shorter
than the middle ones, not otherwise differentiated.

a. Leaves densely silvery-silky, 4 cm long or

UGE GoGcuct socococr Sslee soccicceaiac sacle) Go Ae SeELCeum
aa. Leaves much longer and much less densely pu-
bescent.
b. Involucre 8-12 mm high; ligules 1.5-2.5
cm long; leaves broadly lanceolate ...

aslo salve sade cece secses@cigiscccce Cle A. Bngelmeanmmin
bb. Heads smaller; leaves lanceolate to
long linear.
c. Stem pubescent in lines.
d. Peduncle lightly pubescent in
lines; outer tegules shorter than
the inner .....-eccoeeee 14. A. hesperius
dd. Peduncle heavily pilose; outer
tegules longer than the inner ..
RO Sea ere eeesecaccee 17. A. Subspicatus
cc. Stem pubescence uniformly distributed.
e. Outer tegules much broader and
longer.
£. Stem coarsely hirsute. 7. A. puniceus
ff. Minutely strigose or pube-
rulent .....ssaeeeecee 19. A. EBatonii
ee. Tegules imbricated, i.e. the
outer ones successively shorter.
g.- Tegules glabrous dorsally;
stem minutely pubescent ...
So00ce te ee eee cecccee 19. A. Batonii
gg. Tegules pubescent dorsally;
stem pubescence coarser ...
cee eeeecceeccascecs 20. A. adscendens

1. A. caepace meas L. -- (Pétouane) -- Leaves cons-
picuously cordate, long petiolate, rather large, mostly
about 1 dm across. Herbage densely glandular-puberulent
throughout, varying to glabrous or hirsute below. Heads
corymbose. Ligules pinkish to pale blue. Late summer
and early fall. Deciduous forests, rare: Whiteshell --
NS-seMan, eUS.

After more than a hundred years of successive
reports, the only tangible and firm evidence for our area

ASTER 112

1972 Boivin, Flora of Prairie Provinces 29

still consists of three rosette leaves collected by H.J.
Scoggan in the Whiteshell Forest Reserve in 1951 (CAN).
Although a sterile gathering, it seems clearly refera-
ble to A. macrophyllus on the basis of size, pubescence,
glandulosity and thickened mucros. Dawson's 1875 report
for the Turtle Mountain (TRT) proved to be based on a
specimen of Aster ciliolatus. Reported for Norway House
by Hooker 1834 and Macoun 1884 but the locality has ne-
ver been confirmed and was questioned by Scoggan 1957.

A Richardson collection labelled Lake Winnipeg (CAN) is
correctly named, but almost certainly did not come from
the locality stated. Richardson's localities are usual-
ly to be interpreted in very broad terms and his specimen
was probably collected in the Whiteshell or more likely
in adjacent western Ontario where the species is common.
Both areas were traversed by Franklin's partly to which
Dr. Richardson was attached.

2. A. giliolatus Lindley (A. cordifolius AA.; A.
Lindleyanus T. G.) -= A common forest species with
blue ligules and dimorphic leaves, the lower ovate ona
long and narrowly winged petiole. Herbage villous to
nearly glabrous, the stem and branches pubescent in li-
nes. Upper leaves + lanceolate. Mid summer. Deciduous
forests. --Mack, NS, NB-BC, US.

There has always been a fair amount of confusion
between A. ciliolatus and the more eastern A. cordifolius
L., the latter being reported for 5 localities in Manitoba.
We have examined the specimens from Winnipeg, Miami and
Grand-Rapide and were not surprised when each proved to
belong to A. ciliolatus. The Swan Lake and Brandon have
yet to be examined, but they are not expected to belong
to A. cordifolius.

Aster sagittifolius Wedemeyer was reported by Liéve
1959 from Otterburne(DAO, MSM) on the basis of what we
estimate to be an exceptionally lush specimen of
A. ciliolatus.

2X. A. giliolatus X simplex -- Or perhaps A. ci-
liolatus x hesperius. Middle and lower leaves lanceo-
late, tapered to base to a winged petiole 1-4 cm long.
About 4 dm high. Pubescence of the herbage varying from
+ villous (Mainly the inflorescence) to very finely
scabrous (i.e. the upper leaf surfaces). Main leaves
mostly 1.0-1.5 cm wide, the upper sessile, shorter and
narrower by about half. Heads conspicuouly bicolour,
the ligules white, fading mauve, the center passing from
light to deep purple. Brokenhead. --sMan.

3. A: campestris Nutt. var. campestris -- Leaves
very narrow and densely glandular-puberulent in the in-
florescence. Stoloniferous. Herbage finely strigose
below. Leaves long linear, mostly 2-3 mm wide, the upper

113 ASTER

30 PHY. T OeL:08G I-A Vol. 23, now. 1

gradually shorter down to about 1-2 cm. Ligules blue.
Late summer. Rolling steppes. --swAlta-BC,wUS.

4. A. novae-angliae L. --Michaelmas Daisy --

rr LAPP IPD :

Stem leaves isomegueth, lanceolate with a broadly clas-
ping base, numerous and closely spaced. Herbage finely
scabrous throughout and glandular-puberulent, especial-
ly in the inflorescence. Stem also + hirsute. Heads
corymbose or paniculate. Ligules reddish-purple. Mid
to late summer. Local in low, open spots. --NS, NB-
swQ-sMan, US.

Sometimes cultivated as an ornamental; not other-
wise known from west of Manitoba. It is however one of
those species that may be expected to escape and become
eventually naturalized in the Aspen Grove zone. Acadian
occurrences are such escapes.

5. A. modestus Lindley (A. major (Hooker) Porter)
-- Habit of the last but the leaves narrowed to a nar-
rowly clasping base. Densely glandular-puberulent in
the inflorescence, but mainly villous below, especially
so on the stem. Heads usually few, corymbose to panicu-
late. Ligules deep mauve. Second half of summer. Wet
to boggy spots. --(Y)-Aka, O-(seMan)-wcS-BC, US.

6. £- MacGallae Rydb. --Rather similar to A. cilio-
latus but the leaves firmer and narrower and the herbage
much less pubescent to glabrous. Lower leaves lanceola-
te, rounded to an asymetrical base and a very long petio-
le, the latter very narrowly if at all winged. Upper lea-
ves linear. Heads few. Late summer. Near mountain
streams. --swAlta-seBC.

7. A. puniceus L. var. puniceus (var. oligocephalus
AA.) --Pitnagen, Tea-Flower -- The coarse stem coarsely
and conspicuously hispid. Usually around 1m high. Lea-
ves numerous, not reduced upwards, long-lanceolate with
broadly auriculate clasping bases. Panicle usually
ample. Tegules variable, the outer either longer or shor-
ter than the inner, often somewhat squarrose. Ligules
bluish. Mid summer. Common in marshy places. --L-
-SPM, NS-Alta, US--F. candidus Fern. --Ligules white.
Local. --sMan, (US). arene:

Var. oligocephalus Fern., the usual phenotype in
the northeastern part of the range of the species, has
fewer heads, these solitary on long peduncles which are
not bracteolate, but leafy to the base of the head. It
was given by Fernald 1950 as ranging to Mackenzie and
Saskatchewan. In 1965 the Gray Herbarium held only one
Saskatchewan sheet classified (but not revised) under
this name: Breitung 866, Wallwort. There was no Macken=
zie sheet, but two sheets from the upper Mackenzie ba-
sin in Alberta were also filed as var. oligocephalus,
although unrevised; they came from the Slave and Embar-
ras rivers. All these specimens were closer to our

ASTER 114

1972 Boivin, Flora of Prairie Provinces 51

concept of var. puniceus and were so revised.

8. £- sexiceug Vent. - Leaves fugaceous, those
below the inflorescence usually gone by flowering time.
Tufted perennial from a short, woody rhizome, 3-4 dm
high. Leaves oblong-lanceolate, entire, sessile. Heads
few on very leafy branches. Tegules sericeous and long,
squarrose, acute. Late summer. Local on light soils.
--wO-sMan, cUS.

9. A; leexis L. (var. Geyeri Gray; A. Geyeri (Gray)
Howell) -=- Leaves thickish, somewhat fleshy and slightly
glaucous. Leaf margins scabrous, herbage otherwise gla-
brous. Leaves ovate to lanceolate, the upper sessile
with a broadly clasping base, the middle ones oblanceo-
late or narrowed to a broadly winged petiole and 4 clas-
ping base, the lower petiolate. Involucre strongly im-
bricated, often with conspicuous, rhomboid, green pat-
ches on the tegule tips. Mid summer. Frequent on better
prairie soils. --Q-neBC, US.

Often subdivided in two varieties or species on the
basis of degree of development of the green subterminal
patch on the tegules. The specimens examined did not
exhibit any morphological discontinuity on this score and
both types appear to have substantially the same range.

Reports from west of us should all be carefully
checked as the species appears to reach only into north-=-
eastern B.C.: Dawson Creek (DAO), etc. Reports from
southeastern B.C. were based on specimens of other spe-
cies: the T. Ulke collection at Horsethief Creek (TRT)
was a specimen of A. MacCallae, etc. Reports from
southern Yukon are apparently based on a collection at
mile 611 on the Alaska Highway. But mile 611 is in B.C.
(see Aster conspicuus), about 16 miles short of the Yukon
boundary.

10. A. sibiricus L. var. sibiricus (A. Richard-
sonii Sprengel) -- Tegules purple-margined around a
green center. Loosely tufted to stoloniferous and 1-4
dm high. Leaves + serrate above the middle, the upper
oblong-lanceolate and sessile, the lower narrowly oblan-
ceolate and petiolate. Heads solitary or few on sub-
naked peduncles. Ligules purplish, usually drying blue.
Mid summer to early fall. Alpine shale slides, descen-
ding to river shores. --Mack-Aka, Alta-BC, US, Eur -- F.
glbinus Lepage -- Ligules white. Local: Ft. Saskatche-
wan -- Aka, Alta.

The more northern var. pygmaeus (Lindley) Cody tends
to be lower and monocephalous, its leaves entire and nar-
rower, not over 5 mm wide, and linear-lanceolate.

ll. A, conspicsuus Lindley -- A coarse and showy
herb, densely glandular throughout, and with rather large

115 ASTER

32 PHY T Only 0:G TyAé - Vol. 23, now. 1

heads. Up to 1m high. Leaves large, narrowly obovate
to broadly oblanceolate, narrowed and rounded to a ses-
sile or subclasping base. Heads few, corymbose to broa-
dly paniculate. Ligules violet. Mid summer to frost.
Light woods. --S-BC, US.

Our species was reported from Yukon by Hultén 1950,
Cronquist 1955 and Boivin 1967. The only relevant speci-
men cited or located was an Anderson collection from mile
62 on the Alaska Highway (S), somewhere in the region of
Fort St. John and about 200 miles south of the Yukon boun-
dary. Occasional distributional reports from Yukon are,
like this one, actually based on B.C. collections, inclu-
ding many Alaska Highway collections, Point O on the
Highway is at Dawson Creek in B.C., at 550 45'N,, 120°
15'W, and the road does not cross into Yukon until mile
627. Then for the next 130 miles or so, the road repea-
tedly crosses the B.C.-Yukon boundary. From about mile
730 on, the road stretches diagonally across Yukon Terri-
tory until it enters Alaska at mile 1221.

12. A. exicoides L. var. commutatus (T. & G.) Boivin
(A. adsurgens Greene; A. commutatus (T. & G.) Gray; A.
crassulus Rydb.; A. falcatus Lindley, var. crassulus
(Rydb.) Cronq.; A. multiflorus AA.;A. pansus AA.; A. poly-
cephalus Rydb.; A. stricticaulis (T. & G.) Rydb.) -- An
obvious and common prairie species with rather small
heads and short white ligules. Tufted from a woody corm
in dense sod, becoming long stoloniferous in disturbed
soils. Herbage scabrous=puberulent to glabrous. Leaves
linear. Tegules squarrose, with a large green tip and
spinulose mucro. Second half of summer. Common and abun-
dant in nearly all kinds of steppe or prairie. --Mack-Y,
O-BC, US.

Two more varieties occur to the east and the west
of us. Both are generally larger plants with more nume-
rous and smaller heads on more heavily bracteolate pedun-
cles. See Nat. Can 89: 67-70. 1962 for a detailed compa-
rison. The eastern var. ericoides is stoloniferous and
its outer tegules are 2 mm long or less. The western var.
pansus (Blake) Boivin is tufted from a woody corm and its
outer tegules are 2-3 mm long.

Within our area, whenever a recently built road cuts
across virgin prairie thus opening part of the habitat to
pioneering activities, one can usually find specimens with
vigorous rhizomes radiating from an old and half disinte-
grated corm. Clearly, the presence of corm or stolons in
var. commutatus is of ecological rather than taxonomic
value. Mentions of A. pansus for our area will be found
to refer the tufted phase of var. commutatus.

13. £, lateriflorus (L.) Britton =-- Similar to the
next two but the heads are secund on the more widely

ASTER 116

1972 Boivin, Flora of Prairie Provinces 353

spreading branches, and the corolla lobes more elongate,
being 1.0 - 1.5 mm long. Leaves + lanceolate, serrate,
glabrous below except for the pilose midnerve. Heads
numerous and smallish, at first white, becoming bicolour
with a purplish center. Late summer. Around bluffs and
in light woods. --NS-sMan, US.

A. praealtus Poiret has been reported from our area
and other parts of Canada, but this could not be confir-
med as all Canadian specimens seen proved to belong to
other taxa. Specimens at DAO were mainly of A. hesperius.
Those from WIN were mostly A. simplex with mauve ligules,
some were A. hesperius, one sheet from Winnipeg Beach was
A. lateriflorus, another was the hybrid A, ciliolatus X
simplex (also at DAO). Those at CAN were mostly A. simplex.

A. praealtus stands largely intermediate between A.
hesperius and A. simplex. It is generally a larger plant
with @ more open panicle in the manner of A. simplex, its
main leaves are around 1 dm long and mostly 1.0 - 1.5 cm
wide. But it resembles A. hesperius by its leaves being
entire and the ligules pale mauve to light blue. The
main criteria are in the leaf nervation.

In A. simplex the primary leaf nerves are readily
traceable, being somewhat stronger and slightly rugose
below. They are often nearly parallel to the midnerve.
The interconnecting network of secondary and tertiary
nerves delimitates small tissue areas that are mostly
oblong and mostly 1.5-2-(6) times longer than broad. On
each side of the midnerve there is a narrow strip of tis-
sue that lies largely outside the reticulum, being merely
traversed by the primary nerves.

In A. hesperius the leaf venation is essentially as
in A. simplex, especially if the leaves are of the broa-
der type. If the leaves are very narrow, the primary ner-
ves may not be so clearly distinct, but the tissue areas
will remain elongate.

In A. praealtus only the midnerve is well defined
and rugose. The rest of the nervation lacks clearly defi-
ned primary nerves but consists in a reticulum which deli-
mitates irregular polygons, most of the latter being about
as long as wide, and this reticulum extends right up to
the midnerve. Canadian reports of A. praealtus will gene-
really refer to specimens of A. hesperius or of A. simplex
with coloured ligules.

14. A. hesperius Gray var. hesperius (A. coerules-
cens AA.; A. Franklinianus AA.; A. johannensis AA.; A.
Oesterhoutii AA., A. salicifolius AA.) -- Perhaps our com-
monest and most widespread species, yet highly variable
and rather nondescript. Stoloniferous and forming large
colonies, often up to 1m high. Leaves entire, lanceola-
te to narrowly linear, less than 1 cm wide. Heads many

7 ASTER

3h Puy Tt ODO G fA Vol. 23, now. 1

in a narrow panicle. Involucre 5.0-7.5 mm high, its tegu-
les narrow and long attenuate, their midnerve green and
slightly broadened above into a narrow and elongated

green tip which is usually less than 0.5 mm wide. Tegu-
les less strongly imbricated than A. simplex, or excep-=-
tionally the outer tegules longer than the inner. Ligu-
les white to mauve, often drying pale blue. Second half
of summer and first half of fall. Open places, usually

in the wetter spots. --Mack, Q-BC, US.

Two variants are worthy of notice. Some transi-
tional specimens have the longer outer tegules of A.
subspicatus but are otherwise similar to A. hesperius by
the size and shape of leaf and/or pubescence of peduncle.
They have been called A. hesperius var. laetivirens in
Western Canada and are sympatric to A. subspicatus. A
check of the eastern material shows that this transitio=
nal form is also present especially in areas where A.
subspicatus occurs. Because of its sympatry with A. sub-
spicatus we estimate that var. laetivirens is not a semi-
autonomous population but merely an extreme of variation,
hence not rating taxonomic recognition as a variety
within our scheme of taxonomic categories, and better pla-
ced in the synonymy of A. subspicatus. Western specimens
previously referred to var. laetivirens have been mostly
placed in A. subspicatus, but the similar specimens from
the east have been versed mainly in A. hesperius; the
choice being made partly on the basis of greater simila-
rity, partly because of what else is known to occur in
the same general area.

In var. gaspensis, the second variant, tegules are
more or less imbricated as in A. hesperius, but otherwise
resemble A. subspicatus in being longer, heavier green
and the outer ones wider, mostly 1.0 - 1.5 mm wide, and
in being green throughout or nearly so. Thus the larger
involucre of var. gaspensis, 8-12 mm high, tends to be
darker green than the foliage. The main leaves are enti-
re and mostly 1.0-1.5 dm long and (0.7)-1.0-1.5-(3.0) cm
wide, tapered below to a narrowly clasping base, and thus
resemble A. subspicatus, but they are not contracted into
a broad petiolar base. The pubescence of the peduncle is
of numerous decurrent lines of pubescence. Originally des-
cribed from the shores of the Bonaventure river, var. gas-
pensis is now known from the shores of the Nottaway (Dut.
& Lep. 35311 & 35342) and from Cabbage Willows (Stirrett
1127 at DAO) on the south coast of James Bay just east of
the interprovincial boundary. Var. gaspensis has also
been reported in Bull. Torr. Bot. Club 73: 143. 1947 as
occurring around Lake Mistassini.

Aster hesperius Gray var. gaspensis (Vict.) stat.
ne, A. gaspensis Vict., Contr. Lab. Bot. Un. Mtr. 20:

ASTER LS

1972 Boivin, Flora of Prairie Provinces 35

3.1932. A. hesperius var. gaspensis f. albiflorus (Vict.)
stat. n., A. gaspensis Vict. f. albiflorus Vict., Nat.

Can. Zl: 209.1944; A. novi-belgii L. f. albiflorus (Vict.)
pret Nat. Can. 94: 646. 1967.

johannensis Fern. was accredited to Manitoba on
the Byer? of a series of specimens (CAN, DAO, LKHD) since
revised to A. simplex or to A. hesperius.

The next three species and A. hesperius are not
always clearly distinct inter se.

15. A. simplex W. var. simplex (var. ramosissimus
(T. & G.) Cronq.; A. longifolius AA.; A. paniculatus Lar Lam. )
-~- Like a larger phase of the last. Main stem leaves
1-2 dm long, usually 1-2 cm wide. Involucre 4-6 mm high.
Ligules white; tubular flowers also white, rarely either
or both pinkish. Second half of summer. Wettish open
places. -- NF, NS, NB-cS, US.

In the estuary of the Saint Lawrence it gives way
to a smaller plant with shorter branches, known as var.
estuarinus Boivin.

The various species presented in this flora do not
always differ from one another in the same manner. Usual-
ly there is a definite morphological discontinuity between
closely related species and it is possible to assign to
each taxon definite morphological boundaries that are not
exceeded except in very unusual cases. Further, the spe-
cific criteria for most species exhibit constancy of as-
sociation. Such species are said to be monothetic; they
can be accurately defined by a minimum set of criteria
which are both essential and sufficient for the identifi-
cation of representative specimens. Taxa that do not fit
within the definition above are termed polythetic. See
Sneath, P.H.A., Symp. Soc. Gen. Microb. 12: 291-332. 1962,
and Morse, L.E., Taxon 20: 269-282, 1971.

Polythetic taxa grade into one another and cannot
be assigned precise morphological boundaries; they can
be defined only in terms of a series of criteria which
need not be all present; suffices to recognize such a
taxon that most or nearly all its criteria be present. Of-
ten, the various criteria have unequal value and one of
them may be much more important than the others. Field
experience, herbarium practice and a bit of flair are
helpful in dealing with such taxa and deciding which cha-
racter is to be given more importance. Known distribution
or ecological preferences can offer strong leads, but one
most not transform the place of origin or the habitat into
taxonomic criteria.

Taxa of all ranks, from form to family or higher,
can be either monothetic or polythetic.

Throughout this flora we have tried to restrict the
use the rank of species to monothetic taxa. Monothetic

119 ASTER

36 PH YT 0 b0*G ks Vol. 23, no. 1

species are presumably isolated genetically, while poly-
thetic ones usually stand in partial isolation only.
Whenever two or three taxa intergrade to form a monothe-
tic cluster of polythetic entities, we have usually trea-
ted them as so many varieties of a single species. Hence
our varieties are mostly polythetic taxa.

We have also tried to use each rank, such as spe-
cies, for taxa that are roughly comparable with one ano-
ther; comparable as to the kinds, manners, and degrees
of differences,

The species is also the basic concept by which one
initially apprehends the various elements of a flora or
particular group; other ranks are usually apprehended
later as collections or subdivisions of species. Hence
it is essential that the species concept should corres-—
pond to the level of abstraction most easily assessible to
our expected readers. In simpler words the species should
be and remain a practical concept as stated in our prefa-
Cee

Now, it is not possible at all times to equate spe-
cies and monothetic, and at the same time define only spe-
cies that are roughly equivalent and recognize nothing
but practical units easily comprehended by the informed
(but not necessarily specialist) reader.

Some of the richer clusters of polythetic taxa are
very complex and the range of their morphological varia-
tions is so wide that they cannot possibly be held as
roughly equivalent to other species in the same genus.

As clusters they are also very difficult to apprehend

and delimit from the non-member taxa. Practical expe=
rience has shown that such clusters are easiest to deal
with when each major element is rated as a species and
allowance is made for a certain amount of integrading.
Specimens with intermediate morphology may be frequent and
are best treated as genuine intermediates rather than
hybrids, since there is no sound basis to assume that they
are significantly more heterozygous than most other indi-
viduals referred to the cluster; they merely seem to
present less frequent recombinations of characters.

Aster simplex stands at the center of a very com-
plex cluster of polythetic taxa, comprising with us A.
hesperius, A. borealis, A. subspicatus and A, lateriflorus.
The cluster extends into eastern Canada with the follo-
wing additions A. dumosus L., A. vimineus Lam., A. Tra-
descantii L., A. novi-belgii L. and A. puniceus. More
elements, such as A. praealtus Poiret, also occur fur-
ther south.

If the cluster was much smaller, say if it compri-
sed only A. simplex and A. borealis, we would not have
hesitated to reduce it to a single species with a varie-

ASTER 120

1972 Boivin, Flora of Prairie Provinces 37

ty. But the cluster being as complex as it is, a diffe-
rent kind of solution is called for. Before one can a=
chieve a good general view of the A. simplex cluster, one
must first become acquainted with most of its major
units; hence in the present case the primary unit of
intellectual apprehension is not the monothetic cluster,
but the polythetic and intergrading species as we have
described them above and below. This is why we have re-
tained these intergrading taxa at the rank of species.

Rydberg's floras are good examples of texts based
primarily on the polythetic species. In such texts the
need to recognize subspecies and varieties is greatly
lessened, and most, if not all, taxa can be convenien-
tly termed species. At the other end of the range the
species as used by Gleason 1952 and Hitchcock 1969 is
mostly monothetic. Fernald's Manual is a halfway house.
Our approach is closer to that of Gleason and Hitchcock.

16. A; boseslis (T. & G.) Prov. (A. junceus AA.;
A. junciformis Rydb.) -- Like the last two, but the stem
thin and wiry, the leaves long and narrow, the heads
large and few. Leaves up to 2 dm long and usually less
than 5 mm wide. Heads mostly 1-8. Second half of summer.
Frequent in very marshy or boggy places. --Mack-Aka, NS-
BC, US.

Successively this taxon has been called A. laxifo-
lius, A. laxifolius var. borealis, A. borealis, A. jun-
ceus Aiton and more recently A. junciformis. But A.
borealis appears to be the earlier and correct name.

The synonymy is as follows:

A. borealis (T.& G.) Prov., Fl. Can. 1]: 308,

1863; A. laxifolius Nees var. borealis T. & G., Fl. N.
Am. 2: 138. 1841; A. Franklinianus Rydb., Bull. Torr.

Bot. Club 37: 141. 1910; A. junciformis Rydb., Bull.

Torr. Bot. Club 37: 142. 1910.

This entity was usually placed under A. laxifo-
lius until Gray 1884 pointed out that Nees' specimens
belonged to A. paniculatus (= A. simplex). As a correct
name Gray then took up A. junceus Aiton, the type of the
latter coming from Halifax. Now A. borealis does not
occur in mainland Nova Scotia; obviously Gray's choice
of name was unsound, but this escaped his attention,
probably because of the relatively limited number of spe-
cimens per species in the herbaria of the last century,
the distribution of any species being usually known only
in very general terms.

The matter rested there for another half century
until L.H. Shinners, Am. Midl. Nat. 26: 411-412. 1941,
pointed out that Aiton's specimen was unlikely to be i-
dentical with our plant since it came from outside the
range of our taxon. Hence A. junceus Aiton had to be

121 ASTER

38 P.Bye.2 O}L50),.4 Ech Vol. 23, no. 1

rejected as misapplied and another name was substituted,
namely A. junciformis. The latter name had been propo-=-
sed by Rydbem as a western segregate of what we call A.
borealis. We do not consider A. junciformis to be 4
tenable segregate of A. borealis and the latter is re-
tained as the earlier name.

A. longulus Sheldon is reputed to be the hybrid A.
borealis (or junciformis AA.) X A. hesperius (or caeru-
lescens AA.) and is the source of many obscure or baf-
fling reports for western Canada. Rydberg 1917 and 1932
reports it from Sask. and B.C.; Russell 1944 and 1954
lists collections from Pike Lake and Swift Current; Brei-
tung 1947 and 1957, repeated by Boivin 1967, mentions
McKague and Wallwort.

The type of A. longulus was the object of a detai-
led study by L.H. Shinners in Rhodora 44: 338-9. 1949.
It seems to be essentially similar to A. borealis except
for the pubescence being more abundant in the manner of
A. puniceus. Shinners estimated that Rydberg's report
was probably based on specimens of A. hesperius (or A.
caerulescens AA.) His guess is partly confirmed by two
Saskatchewan (CAN) collections identified A. longulus
by Rydberg but which seem typical enough of A. hespe-
rius. Russell's specimens have not been seen. Brei-
tung quoted his collections 865, 871 and 1503; these
(DAO) have been revised to A. hesperius and A. simplex.
No specimen was seen from our area that we could clear-
ly place with A. longulus as described by Shinners.

17. A- subspigatus Nees var. subspicatus (A. ci=
liomarginatus Rydb.; A. foliaceus Lindley, var. Parryi
(D.C. Baton) Gray; A. frondeus (Gray) Rydb.; A. hespe-
rius Gray var. laetivirens (Greene) Crong.; A. Tweedyii
AA.) -- Pitnagen, Tea-Flower -- A few outer tegules lon-
ger and larger than the middle and (usually) the inner
ones. Stem pilose along a longitudinal strip. Leaves
+ lanceolate, the middle and lower ones often oblanceo-
late, the middle ones most often narrowed below to 4
narrowly clasping base, the margin scabrous, the limb
glabrous to scabrous above. Tegules usually green to
the tip, sometimes squarrose, ciliate, the involucre
otherwise glabrous, but the peduncle heavily pilose.
Ligules blueish. Mid summer to late fall. Near moun-
tain streams. -- K, Aka, L-NF, NS, NB-eMan-BC, US --
Var. apricus (Gray) Boivin -- Smaller, only 2.5 dm high
or less. Leaves narrowly oblanceolate. Tegules usually
purple-tipped. The more common phase at higher altitu-
des. -= swAlta-BC, (wUS).

Var. apricus (Gray) stat n., A. foliaceus Lindley
var. apricus Gray, Syn. Fl. 1,2: 193. 1884,

ASTER 122

1972 Boivin, Flora of Prairie Provinces 39

subspicatus is a difficult species, difficult to
SR AGS difficult to grasp, and difficult to organize
into its component variations. It is clearly related to
A. hesperius into which it grades so thoroughly that the
two are separable only arbitrarily. In the east it gra-
des similarly into A. novi-belgii L. All three are howe-
ver only partly sympatric and, on that basis, their dis-
tinction is considered significant. There is also some
conflict as to what constitutes here the most satisfac-
tory taxonomic boundary.

We have placed into subspicatus all those specimens
with a few outer tegules green throughout and more or
less equaling or overtopping the inner ones. Such speci-
mens are also commonly separable from A. hesperius by a
number of other characters none of which is quite cons-
tant. In A. subspicatus the leaves are commonly narrowly
lanceolate and somewhat larger, usually around 1 dm long
and 1 cm wide, the sessile blade is less narrowed at
base and definitely clasping (not or barely clasping in
hesperius); the larger lower leaves will be slightly
contracted below into a broadly winged and ill-defined
petiole. The peduncle is densely pubescent (only pube-
rulent in lines in A. hesperius). Pappus at first whi-
te; often becoming purple-tinged. Further in A. subs-
picatus the outer and broader tegules are green throu-
ghout or nearly so, while the other tegules have a
rather broad and intense green patch, usually around
1 mm wide, the net effect is that the involucre is as
green or somewhat darker green than the rest of the fo-
liage. In A. hesperius the tegules are narrower by half
and the green patches still narrower and light green,
the involucre is obviously paler than the leaves. The
ligules are usually of a deeper blue in A. subspicatus.

As defined here A. subspicatus is very similar to
A. foliaceus sensu Fernald 1950, except that the latter
would distinguish as A. crenifolius (Fern.) Cronqg. one
broad-leaved collection, perhaps of hybrid origin. In
Cronquist 1952 the subdivision is on a different basis
and the bulk of the eastern material that we call A.
subspicatus is placed in A. johannensis along with much
of the eastern specimens of A. hesperius, while the broa-
der-leaved are placed in A. crenifolius. In Cronquist
1955 the western material of what we call A. subspicatus
is distributed between three species. If the leaves are
not contracted at base into a broadly winged petiole, it
is called A. hesperius var. laetivirens (vel sphalmate
var. laetevirens). If the middle and lower leaves are ‘
contracted below, they may be called A. foliaceus if the
leaves are entire or only obscurely denticulate, but A.
subspicatus if the leaves are obviously serrate.

123 ASTER

0 Pslye2 O2Li0+G DA Vol. 23, Nos.

As can be gathered by Cronquist's treatment, the
western phase of A. subspicatus is more variable; its
lower leaves are sometimes attenuate into a barely
winged petiole, more infrequently a basal rosette will
be produced. Thus we are not fully convinced that
the eastern and western populations should be treated
as a single taxon. But being unable to establish a
satisfactory morphological basis for a taxonomic dis-
tinction, we have retained them as a single taxon.

18. A; occidentalis Nutt. var. occidentalis --
Ligules blueish like the last, but the tegules imbri-
cated, the outer being shorter and often squarrose.
Mostly 2-4 dm high. Leaves narrowly lanceolate, usual-
ly less than 1 cm wide, the lower long petiolate. Heads
mostly less than 10, paniculate or more often corymbose,
borne on branches bearing few, if any, bracts. Second
half of summer. Near shores. --(Mack), wAlta-BC, US.

Grades into the more southern var. intermedius
Gray, taller, with more heads, these paniculate, and
borne on branches with numerous bracts.

19. A. Batonii (Gray) Howell (A. Mearnsii Rydb.)
-- Outer tegules oblanceolate with a dark green and
broad tip, glabrous dorsally and somewhat ciliate, from
much shorter to much longer than the inner. Resembling
the above two, but the stem leaves not dimegueth, rather
narrowly lanceolate to linear. Stem pubescence also si-
milarly fine and not in lines. Mid summer. Along creeks
and rivers: Cypress Hills and Rockies. --swS-sAlta-sBC,
wUs.

Reportedly the ligules vary from white to blue;
but the latter colour clearly prevailed in all the Cana-
dian specimens checked.

20. A, adscendens Lindley (A. chilensis Nees ssp.
adscendens (Lindley) Cronq.; A. oblongifolius AA.; A.
Richardsonii AA.; A. subgriseus Rydb.) -- Involucre simi-
lar to that of A. ericoides but the heads larger and
the ligules blue. Mostly 2-4 dm high. Herbage more or
less villous, especially the stem. Leaves linear, the
lower much longer, narrowly oblanceolate and tapering to
@ subpetiolar base. Tegules much imbricated, squarrose,
with a broad green tip, spinulose=-mucronate. Pappus
rather dark gray violet. Late summer. Steppes; umncom-
mon. --S-seBC, wUS.

21. A. Engelmannii (Eaton) Gray -- Tegules broad-
ly margined in purple in the upper third. Leaves soft
pubescent and finely glandular below, to nearly glabrous.
Heads largest, 3-5 cm across. Peduncles naked. Ligules

mauve. Mid to late summer. Mountain meadows. --swAlta-
sBC, wUS.

ASTER 124

1972 Boivin, Flora of Prairie Provinces 41

22. A. pauciflorys Nutt. -- Heads few on elongate
and bracteolate peduncles, in a densely glandular-~puberu-
lent corymb. Loosely tufted, the stems 2-3 dm high, de-
cumbent at base. Leaves narrow and long linear, much redu-
ced upwards and grading into the small inflorescence
bracts. Ligules white, sometimes fading mauve and drying
pale blue. Second half of summer. Alkali flats. --
sMack, Man-sAlta, cUS.

23. A. umbellatys Miller var. pubeyns Gray (A. pu-
bentior Cronq.; Doellingeria pubens (Gray) Rydb.) --
Heads white in a broad corymb. Leaves numerous, broad,
lanceolate. Involucre only slightly longer than the ma-
ture seeds. Second half of summer. Moist open ground.
-- (NB-Q)-O-Alta, (US).

In our variety the herbage is densely puberulent
including the tegules. Also the upper leaf surface is
less densely puberulent than the lower, and ligules are
only 4-7 mm long. In the more eastern var. umbellatus
the stem and tegules are glabrous or nearly so, the lea-
ves are glabrous below, or at least less puberulent
below than above, the involucre is sometimes shorter
than the seeds, and the ligules are more variable, being
commonly narrower and up to 8-(12) mm long. Budd 1957
and 1964 reports the typical variety to be common in Ma-
nitoba, but in 1969 all the specimens at SCS were cor-
rectly filed as var. pubens. The source of Budd's re-
port remains unclear, unless it was based on such ear-
lier reports as were discounted by Scoggan 1957.

Var. pubens is sometimes rated as a distinct spe-
cies, but except for the density of pubescence, none of
the diagnostic criteria amounts to much more than weak
tendencies with broad zones of overlap.

24. A. laurentianus Fern. (A. angustus (Lindley)
T. & G.; A. Brachyactis Blake) -= Discoid, annual. Lea-
ves long linear, the earlier ones + fleshy and longer,
but deciduous and usually lacking in herbarium speci-
mens. Glabrous except for the scabrous-ciliate leaves.
Tegules isomegueth or nearly so. Inflorescence narrow-
ly paniculate to much diffused, Pappus often becoming
much larger as the specimen dries. Late summer to frost.
Saline shores, sometimes weedy. --seK-Y, PEI-BC, US,
(eBur).

Young plants of A. Brachyactis Blake have larger
and fleshy leaves which soon wither. These are usually
lacking in herbarium specimens. The type material of A.
laurentianus, a name older than A. Brachyactis, is made
up of such juvenile and fleshy plants.

25. A, alpinus L. (var. Vierhapperi (Onno) Cronq.)
-- With a single large head. Tufted, 1-2-(3) dm high.

125 ASTER

2 PAB? OMLOvG IA Vol. 23, no. 1

Herbage densely pilose and inconspicuously glandular-
puberulent throughout. Upper leaves linear, much re-
duced. Heads 3.5-4.0 cm across. Tegules purple-mar-
gined and almost isomegueth, thus resembling an Erige-
ron. Ligules usually white, varying to pink or mauve.
First half of summer. Dry mountain slopes and Pine fo-
rests. -- Mack-Y, swAlta-BC, (US), nEur.

A. Tradescantii L. was credited to Saskatchewan
by Macoun 1886, repeating a report by Gray 1884, but
was ignored by later authors. The basis for this deba-
table report has not been investigated.

14. MACHAERANTHERA Nees

A minor segregate of Aster, without stolons but
with a well defined taproot.

a. Leaves at least pinnatifid...... 1. M. tanacetifolia
aa. Serrate to entire.
b. Leaves all or mostly entire..... 2. M. canescens
bb. All spinulose=serrate........ 3. M. grindelioides

1. M,. tanacetifolia (HBK.) Nees -- Leaves pinnati-
fid to tripinnatifid. Annual. Herbage abundantly glan-
dular-puberulent, sometimes also pilose. Leaf lobes whi-
te-spinulose at tip. Heads + corymbose. Tegules long
squarrose, green above the middle, whitish below. Ligu-
les blue. (Early summer?). Arroyos, rare. --swAlta,
cUS, (CA).

Still known only from the original collection by
Dawson in 1881 along the Belly River (CAN), presumably
near the 49th parallel. Its occurrence in Canada has yet
to be confirmed.

2. M. canescens (Pursh) Gray (M. pulverulenta

(Nutt.) Greene; Aster canescens Pursh, var. viscosus
(Nutt.) Gray) -- Herbage densely grayish-puberulent. An-

nual, diffusely branched. Leaves mainly entire, but the
earlier stem leaves often remotely dentate. Tegules
short-squarrose, the deflexed part deep green, densely
puberulent and densely glandular, the lower part whitish
and glandless. Ligules blue. Summer. Badlands and sali-
ne flats. --sS-sBC, wUS.

3. M: ggindelioides (Nutt.) Shinners var. grinde-
Jioides (Haplopappus Nuttallii T. € G.; Sideranthus grin-
delioides (Nutt.) Britton) -- Leaves thickish and quite

regularly spinulose-serrate. Perennial with tufted annual
stems arising from a thick and porous=-woody taproot. Leaf
teeth ending in long, white, spinulose setae. Heads dis-
coid, yellow. Tegules densely puberulent and densely
glandular. First half of summer. Dry hills and badlands.
--swS-sAlta, US.

ASTER 126

1972 Boivin, Flora of Prairie Provinces 43

The more southern var. depressa (Maguire) Crong. &
Keck is essentially a smaller plant, perhaps only a dwarf
form of more arid places.

15. ERIGERON L. FLEABANE
Closely resembling Aster and perhaps grading into
it. Tegules usually isomegueth and usually very narrow,
i.e. less than 1 mm wide; ligules mostly similarly nar-
row, Not stoloniferous but loosely tufted or taprooted
and frequently with few or only one large head, usually
borne on a long subnaked peduncle.

a. Heads discoid or with very short (1-3 mm) and in-
CONMBPLCUCUS! LIigullebissssciessclasceveuvcsccacess (Group .A
aa. Heads conspicuously radiate.
b. Less than 2 dm high and typically monocephalous.
c. Stem leaves numerous,

Ge) uGules) Wilt! isis 'es sss 's0 «se ee else oh CLOUD INE
dd fnigilies cOLOUred es csns ce se clinee CLOUD GC

cc. Foliage mainly basal, the stem leaves
MOSEL *h= SO) NOME felsielelaicicisisla aiclepisies GDOUp D

bb. Taller or many-headed, usually both.
See LOU Wille, veils cistssidisa sien a's elec ei sisia, LOLOUpD.
Se. uioGulLes COLOULEd Clicescerscstesirics sos 5 5 GLOUD C

Group A

Heads discoid or merely with short and inconspi-
cuous ligules projecting only 1-(3) mm beyond the tegu-
les.

a. Leaves deeply dissected .seccoccecss 15, Ez compositus
aa. Entire. ie
b. Monocephalous and 1 dm high or less.
c. Involucre usually + 7 mm high and
heavily lanate in dark blue,. 13. E. uniflorus
cc. Involucre smaller, 5-6 mm high, =
and merely pilose towards the
base with hyaline hairs ...... 21. E. Scotteri
bb. Normally taller and the heads many to

numerous; involucre not lanate.
d. Involucre only 2.5=5.0 mm high and
Glabrous ......seeceesee+-. 24, BE. canadensis

dd. Heads larger and variously pubescent
or glandular.
e. Involucre finely glandular, little
if at all pubescent; inflorescence
tending-~to corymbose........ 22. E. acris
ee. Involucre hirsute and not glandular.

127 ASTER

hh Poniet OPLiOwG IA Vol. 23, nowd

f. Leaves oblong-linear, shorter
than the peduncles of the lower
heads scesseccsse esses. 2oe) emenheeme
ff. Leaves very long linear and
overtopping the lower heads
of the racemose inflorescen-

CO scecvcsvcceces 40. EB. lonchommaliae

Group B
Ligules white; stem quite leafy.

a. Middle leaves longeSt ceoc--e+co 19. EB. hyssopifolius
aa. Leaves gradually reduced upwards, the upper
less than half as long as the lower.
b. Ligulate florets with a minute pappus;
plants usually over 4 dm high and with
many heads «....scecc.cecccsecsesssas 18. He annuus
bb. Ligulate florets with normal pappus, like
the inner florets.
c. Leaves 3 mm wide or less; stem leaves
all uniformly narrow, the upper merely
SHOETCT ssc 0 0 wiepwpois ain aicls oleae ey 9 (Om Eee | Meee
cc. Lower and basal leaves larger, the
upper leaves gradually narrower and
shorter.
d. Stem leaves 10-20; stems tufted
from a taproot .........9. H. caespitosus
dd. Stem leaves fewer, mostly, 5-7;
tufted but not forming a taproot,
the caudex merely covered by fi-
brous roots.
e. Leaves scabrous or pilose on
both faces; upper leaves
SSMS, OG OOGObOGE COOK -. 4. 5. Yaspex
ee. Leaves scabrous at margin only,
glabrous below and lightly pu-
bescent above .ecoeoee 3. Ee glabellus

Group C
Ligules coloured; stem quite leafy.

a. Middle leaves longest, 1-3 cm long..cccoccaccac
aonodaegosaoaandor seeecccoreeeee 19. E. hyssopifolius
aa. Leaves much longer and the lowest longest.
b. All or nearly all leaves auriculate-
clasping at base ....0....-. 16. E. philadelphicus
bb. None or only the upper leaves clasping.
c. Ligules (1.5)-2.0-(3.0) mm wide.
d. Stem leaves few, lightly hirsute
below, more densely so above ...

cece rece ccc seer eeeee§ Gf BE. grandiflorus

ERIGERON 128

1972 Boivin, Flora of Prairie Provinces 5

dd. Stem leaves numerous and at least the
middle and lower glabrous on both fa-
ces except for the midnerve and the
marginal ciliation ..... 1. E. peregrinus
cc. Ligules filiform, less than 1 mm wide.
e. Involucre pubescent; stem leaves
mostly 5-7, pubescent above .....
Cer r errr reve ceseccccecees So Be glabellus
ee. Involucre merely finely glandular
sometimes also pubescent; stem
leaves much more numerous and ci-

liate seccccevcsccececses 2s BE. speciosus

Group D
Small and monocephalous with the foliage mainly
basal. Less than 2 dm high. Stem leaves usually 1-3
and much reduced.

a. Leaves finely dissected ........... 15. E. compositus
aa. Leaves entire to 3-toothed at apex.
De hi gules yellow S.cee.scce-scncseees 145 EB. aureus
bb. White or pink to mauve or blueish, sometimes
yellowish in drying.
c. Involucre 9-13 mm high, heavily lanate.
d. Ligules white; some leaves 3-toothed
Mi cial=\ ol ee OSEO OO Cae see 12. E. lanatus
dd. Ligules coloured; all leaves entire ..
sere ceceesecesceoee 9. EB. grandiflorus
cc. ereatene only 4-8 mm high and not la-
nate, although sometimes heavily pilose.
e. Leaves mostly spatulate, varying
from narrowly obovate to oblanceo-
late.
f. Some leaves 3=5-toothed at
QPEX oe. -ecereseeeeeee 14. BE. pallens
fF. All entire.
ge Long stoloniferous; her-
bage Strigose ..scsccccecs
ecccescco+s-eeo 17. BE flagellaris
gg. Tufted and the stem pilose
with spreading hairs.
he Ligules + 3 mm long and very
narrow; plant usually 3-5
om high ..... 21. E. Scotteri
hh. Ligules longer and + 1 mm
wide; plant about 1 dm
high ....e.... 10. BE. Arthurii
ee. Leaves long linear; ligules broad.
i, Leaves 2 cm long or less; stem
usually scapose ..... 8. E. radicatus

129 ERIGERON

6 PBT OL, OG TA Vol. 23, no. 1

ii. Leaves much longer; stem bearing
a few leaves ..... 7. BE. ochroleucus

1. E. peregrinus (Pursh) Greene var. scaposus (T. &
G.) Crongq. (E. callianthemus Greene; E. salsuginosus AA.)
-- Ligules broad, 1.5-3.0 mm wide, like an Aster. Usual-
ly monocephalous and (1) -3-4(7) dm high, with 5-10 oblong
to oblanceolate leaves. MInvolucre densely and finely
glandular. Mid summer. Open woods in the mountains.
--Y-(Aka), swAlta-BC, wUS.

In the more western typical phase the involucre is
merely villous, not glandular, and the foliage tends to
be more ample.

2. E. speciosus (Lindley) DC. var. speciosus --
Resembling the last, but the blue ligules narrow and the
leaves more numerous. Heads solitary or more commonly
3-5. Herbage nearly glabrous except in the inflorescen-
ce, but the leaves ciliate. Ligules mauve, often drying

yellowish. Mid summer. Open places in the mountain. --
swAlta-sBC, US, (CA) -- Var. conspicuus (Rydb.) Boivin

(BE. subtrinervis Rydb. var. conspicuus (Rydb.) Cronq.)
-- More pubescent. Stem somewhat hirsute. Leaves some-
what pubescent on both faces, more densely so along the
midnerve.--swAlta-sBC, nwUS.

Stat. n., E. conspicuus Rydb., Mem. N.Y. Bot. Gard.
i: 400. 1900.

3. & glabellus Nutt. var. glabellus (var. pubescens
Hooker; E. anodontus Lunell; E. asper Nutt. var. pubescens
(Hooker) Breitung, f. roseata (Lunell) Breitung; E. Drum-
mondii Greene; E. oligodontus Lunell; E. speciosus AA.)
--A common and showy, tufted prairie species, mostly 3-4
dm high, with few heads and pink filiform ligules. Lea-
ves glabrous below, scabrous at margin, lightly pubes-
cent above. Upper leaves 1/3-1/2 as long as the lower.
Involucre hairy but not glandular. Mid summer, Frequent
on better prairie soils. -- Mack, Man-BC, US.

There is a fair amount of confusion in the herbaria
and in the botanical literature about this and the next
species. We have therefore based our distributions sole-
ly on the specimens examined for each entity.

At DAO all the Yukon and Alaskan specimens under
E. glabellus and its synonyms have been revised to E.
asper. We are assuming that the material in other herba-
ria should be similarly revised,

Further south our typical plant is replaced by a
var. yisgidus (Rydb.) stat. n., E. formosissimus Greene
var. viscidus Rydb., Bull. Torr. Bot. Club 28: 24, 1901,
inconspicuously glandular on the involucre, the gladulo-
sity being often somewhat hidden under the copious and
longer pubescence.

ERIGERON 130

1972 Boivin, Flora of Prairie Provinces 7

4. » gsper Nutt. (var. pubescens AA., f. roseata
AA; E. glabellus var. pubescens -; EB. oblonceolatus AA.)
-- Like the last, with which it is sometimes confused, but
more pubescent, the ligules white, the upper leaves more
reduced, and flowering earlier. Leaves scabrous-pubescent
to coarsely pilose on both faces. Upper leaves distant
and much reduced, down to about 1 cm. Ligules sometimes
fading pink. Early summer. Common on prairies. --Y-
Aka, Man-BC, US.

Because the ligules tend to fade and dry pink, the
latter colour is more common in the herbarium than in the
field.

5. E; gaespitosys Nutt. (E. condensatus AA.) --
Rather resembling E. asper, but smaller and in tufts from
a taproot, the stem leaves more numerous, the upper not
reduced so much. Taproot thickish, branched at top.
Stems 2 dm high or less, mostly monocephalous. Pubescen-
ce denser and shorter than in the next and the last two,
the hairs mostly 0.3-0.5 mm long. Ligules white. Mid
summer. Common on steppes and hillsides. --Y-Aka, Man-
BC, US.

E. Engelmannii Nelson was reported for southern
Saskatchewan by Budd 1957 and 1964. It proved to be ba-
sed on a Nashlyn (DAO, SCS) collection revised to E. caes-
pitosus by Breitung in 1955. We concur.

6. E. pumilus Nutt. var. pumilus -- Daisy -- For-
ming an obvious series with the last three species. Lea-
ves narrowest, long linear, and less than 3 mm wide. Tuf-
ted from a taproot. Mostly around 1 dm high. Herbage
villous throughout, especially so on the stem. Tegules
lightly hirsute. Ligules white. Late spring and early
summer. Sandy hills. --sS-sAlta, US.

The more western var. gracilior Cronq. is 4 bigger
plant, commonly about twice as tall and often with 5-8
heads; pappus of shorter and often of squammiform setae.

7. B- gcbreleucus Nutt. var. Goribneri (Canby)
Cronq. -- A small monocephalous type with narrow leaves
like the last, but the pubescence somewhat lanate at
least on the involucre, the ligules usually lavender and
the tegules with squarrose purplish tips. Basal leaves
about half as high as the stem, the latter with few lea-
ves, mostly 3, and much reduced. Heads showy. Involu-
cre about 7 mm high. Early summer. Alpine meadows and
summits. --swAlta, US.

Reports for the Cypress Hills, see Brittonia 6:
189. 1947, are to be discounted. They have not been
confirmed by modern collections and the sole justifying
sheet located, Macoun, Cypress. Hills, June 28, 1894 (MO),
has since been revised to E, radicatus.

Typical var. ochroleucus is 4 taller plant, 1-4 dm
high, with larger and more numerous stem leaves.

131 ERIGERON

48 Pon YT OCLO%G TA Vol. 23, nO st

8. E. radicatus Hooker (EB. peucephyllus AA.) --
Basal leaves narrow and rather short, less than 2 cm
long and not over 3 mm wide. Tufted, subscapose, less
than 1 dm high. Somewhat lanate towards the base of the
involucre, otherwise lightly strigose throughout. Invo-
lucre about 5 mm high. Ligules + 2 mm wide. Early sun-
mer. Rare on dry ridges and hilltops: Old Wives Creek,
Eastend, Wood Mountain, Cypress Hills, Jasper Lake and
Moose Mountain Creek in Alberta. --swS-swAlta, wUS.

9. E. grandiflorus Hooker -- Ligules wide, and
generally like a sma E. peregrinus, but the herbage
long pilose throughout, including the involucre. Only
1-2 dm high. Leaves not so wide, narrowly oblanceolate
to linear, densely long ciliate, the stem leaves about
5. Ligules often turn yellowish in drying. lEHarly sum-

mer. Alpine prairies. --wF, Mack-Aka, swAlta-eBC.
10. 5. Agthusii Boivin (E. acris var. asteroides
X aureus AA.; E. uncialis var. conjugans AA.) -- Similar

in habit and size to the next, but the ligules narrower
and not yellow. Also approching E. Scotteri but general-
ly larger, especially the ligules. Stem + 1 dm high.
Rosette leaves spathulate to oblanceolate, up to 1 cm
wide. Stem leaves (1)-2-(4), much smaller, the upper ob-
curely glandular. Pubescence and glandulosity as in the
next species, except that the involucral pilosity is not
so dense and the stem and the basal leaves are similarly
pilose. Head solitary or rarely with a smaller head ari-
sing from the upper axil. MInvolucre 6-7 mm high, appres-
sed, the tips purple. lLigules about 1 cm long and + 1 mm
wide, at first white, soon turning rose (or perhaps light
mauve), but the half-grown ligules often yellowish. Mid
summer. Alpine gravel slopes. -- swAlta-swBEC.

Sp. n.; EH. acris X aureus sensu Cronquist 1947 et
sensu Boivin 1967. Perennis, caespitosus, decimetralis et
monocephalus. Pilosus omnino nisi ad summas tegularum
ubi minute glandulosus. Folia rosettae 2-10 cm long, ad
1 cm lat,,a spathulatis oblanceolata. Caulis pilosa et
t glandulosa, dense et minute glandulosa in peduculo,
foliis (1)-2-(4). Involucrum appressum, 6-7 mm alt. Tegu-
lae ad basas pilosae et obscure glandulosae, ad summas pur-
pureae et minute glandulosae. Ligulae circa 50, in pri-
mis albae, deinde roseae (vel forsan pallide lilacinae),
+ 1 co long.,+imm lat. Flores disci jutei. Pappus 4-5
mm. Semen puberulum. Typus: Calder & Holm 24064, Twin
Cairn Peak, B.C., gravelly steep slope about 7950, July
29, 1959 (DAO; isotypi: ALTA,CAN). Paratypi: Calder &
Holm 24067, eodem (DAO); Calder & alii 19698 A, Quinis-
coe L., Keremeos, B.C., Aug. 3, 1963 (DAO); Taylor &
Ferguson, Lakit Mountain, B.C., July 15, 1958 (DAO);

E. Scamman 6710, Mt. Assiniboine, B.C., Aug. 7-17, 1952

ERIGERON 132

1972 Boivin, Flora of Prairie Provinces ho

(CAN); Macoun 70350, Mount Forget-me-not, Alta., July 16,
1897 (CAN); D. Be Liaar 274, Banff, Cascade Mt., Aug., 18,
1916 (CAN) ; D. Pelluet 210, Banff, Mt. Inglismaldie, 5 ar
ig LOLS (CAN).

A putative hybrid of acris X aureus was reported in
Brittonia 6: 230. 1947 on the basis of three collections
of which we have yet seen only one, the Forget-me-not spe-
cimen cited above.

So named after Dr. Arthur Cronquist, author of an
excellent monograph of the genus for north America and
apparently the first to have noticed the morphological
originality of this taxon.

ll. §&. aureus Greene -- Ligules yellow, fading
brown, 5-7-(10) mm long, (1.5)-2.0-(2.5) mm wide. A
small monocephalous type, the stem usually unifoliate,
obscurely glandular and spreading pilose. Rosette leaves
more densely pubescent than the stem and with shorter and
somewhat appressed hairs. Involucre (6)-7-(8) mm high,
appressed to irregularly long squarrose, lightly to hea-
vily long lanate with white to purple-black hairs. Sum-
mer. Alpine prairies and summits -=- swAlta-BC, nwUS.

12. E, lanatus Hooker -- Heavily long- Taeetae es-
pecially the involucre, with hairs up to 2-5 mm long.
Basal leaves narrowly oblanceolate, some of them 3-toothed
at apex. Ligules 1-3 mm wide, usually white. Mid summer.
High alpine on talus slopes and summits. --swAlta-seBC,
US.

A recent range extension to Lake Kluane in southwes-
tern Yukon in Can. Field-Nat. 82: 114-5. 1968 proved to
be based on a specimen of E. purpuratus var. dilatatus.

The monocephalous Aster alpinus is habitally simi-
lar to E. aureus and E. lanatus, but the ligules of A.
alpinus are white to pinkish and 1-2 mm wide, the tegu-
les coarser, 1-2 mm wide, tending to oblanceolate and
less than 9 mm long, the leaves entire, heavily pilose
and oblanceolate.

13. 5. uniflorus L. var. unalaschkensis (DC.) Boi-
vin (EB. humilis Graham; E. unalaschkensis (DC.) Vier.)

-- Involucre heavily lanate with dark bluish-tinted hairs,
the dark blue hue being due to the deep purple cross-
walls of the hairs. Less than 1 dm high and monocepha-=
lous. Ligules filiform, 3 mm long or less, white to
bluish. Second half of summer. Tundra and alpine slo-
pes. --G-Aka, L, nQ, nMan, swAlta-BC, US, Bur.

Stat. n., B. pulchellus var. Ee Ae DC.,
Prodr. 5: 287. 1836. In the var. eriocephalus (J. Vahl)
stat. n., E. eriocephalus J. Vahl ex Horn., Fl. Dan. 13:
2299. 1840, the involucre is lanate in white. The latter
is more strictly arctic and the more southern records
from B.C. are to be discounted; all B.C. specimens at

133 ERIGERON

50 PoboreT Ob!) 0vG sToA Vol. 23, snopal

DAO, QK and MTJB, including the mount Avalanche sheet ci-
ted by Macoun 1896, have been revised to var. unalaschken-
sis, while other herbaria visited, including UBC and V,
held no specimen at all.

There is also in B.C. a colour form that is likely
to turn up in Alberta and could create some confusion: E.
uniflorus var. unalaschkensis f. pallidus (Cronq.) stat,
n., EB. humilis f. pallidus Cronq., Brittonia 6: 239. 1947,
in which the involucral pubescence is white or essential-
ly so. Such plants have the color of var. eriocephalus
but the head size and appearance of var. unalaschkensis.
In the latter the involucre is purple black, (6)-7-(9) mm
high, and the tegules are all appressed or the tips some-
times lax. In var. eriocephalus the involucre is deep
purple, more densely lanate, 8-9-(10) mm high, and laxer,
the outer tegules tending to be long squarrose.

14. §£. pallens Cronq. -- Leaves spathulate, mostly
3-toothed at apex. Low and monocephalous. Herbage long
villous and inconspicuously glandular throughout, beco-
ming yellowish lanate on the involucre. Ligules usually
white. Mid summer. High alpine on shale slides. --swAl-
ta-seBc.

Known to us only from Mount Saskatchewan (DAO),
the Hanging Glacier (NY), Lake Louise (CAN), Shovel Pass
(CAN), Bee Mountain (CAN), and Mount Copperstain (UBC),
the latter two from B.C. We have not seen the Mount
MacDonald collection.

The range was recently extended northward into
western Mackenzie on the basis of a series of specimens
(DAO) originally identified as E. purpuratus Greene, la-
ter revised to E. pallens and cited as such by Cody 1969.
The justifying specimens have entire leaves, their invo-
lucre is heavily pilose with + purplish hairs, their pap-
pus turn purplish at anthesis, etc.; they do not differ
substantially from E. purpuratus except for their con-
sistently broader leaves which give them a superficial
similarity to the closely related E. pallens. They ap-
parently represent a hitherto unrecognized geographical
variant and may be known as follows:

E. purpuratus var. dilatatus var. n., foliis la-
tioribus et modo brevoribus, a spathulatis oblanceolatis,
praecipue 2-3 mm lat. Typus: Kvale & Haggard 131, Mac-
kenzie Mtns, Redstone River, dry soil and talus, 4 July
1963 (DAO). Paratypes from the Mackenzie Mountains as
cited by Cody and also from the Quill Creek area (DAO)
in Yukon. The latter was reported as E. humilis in Bot.
Not. 199: 204. 1956. ote ee

15. &; compositus Pursh (var. glabratus Macoun,

var. discoideus AA.: E. trifidus Hooker) -- Leaves dee-

ply dissected into narrow segments, tripartite to tri-
ternatifid. 2 dm high or less and monocephalous. Stem

ERIGERON 134

1972 Boivin, Flora of Prairie Provinces 51

leaves few, linear, entire, reduced, Ligules usually
white, variable in length. Late spring and early summer.
Infrequent in open places, including steppes and alpine
prairies. --G-F-(K)-Mack-Y-(Aka), NF, seQ, sS-BC, US
--F. discoi (Gray) Vict. & Rouss. --Heads discoid.
--Y, NF, seQ, swS-BC, US.

Many varieties have been described. We are in-
clined to regard them as ecologically conditioned va-
riants: plants from lower altitudes or from along wa-
tercourses tend to be taller, their leaves are more di-
vided, the lobes longer, etc.

16. &, ww L. var. philadelphicus
(B. purpureus Aiton) -- Most stem leaves clasping at
base. Shallow-rooted and + biennial. Herbage long vil-
lous or hirsute. Inflorescence corymbose. Ligules pink
or mauve, filiform, numerous. Early summer. Wettish
ground, --swMack-Y, NF, NS-(PEI)-NB-BC, US.

Hultén 1950 extended the range to southern Yukon,
but his only cited specimen was from Liard Hot Springs
in northern B.C. We have however checked the following
Yukon collection: E. Schoff, W. Dawson, August 1904
(TRT). A collection from Old Crow is also cited by Hul-
tén 1967.

The more eastern var. Provancheri (Vict. & Rouss.)
stat. n., BE. Provancheri Vict. & Rouss., Contr. Inst.
Bot. Un. Mtr. 36: 58. 1940, from the estuaries of the
Saint Lawrence and of the Hudson, is smaller and essen-
tially glabrous.

17. B& €lagellaris Gray -- With long and conspi-
cuous superficial stolons. Mostly 1-2 dm high and the
stolons about as long as the stem. Rosette leaves more
numerous, spatulate-oblanceolate. Stem leaves only 1-2
-(3) and much smaller, oblanceolate, similar to the sto-
lon leaves, but the latter more numerous. Mid summer.
Mountain meadows in Waterton. --swAlta-sBC, US.

18. 5B, annuus (L.) Pers. (BE. ramosus (Walter)
BSP.; 5. strigosus Muhl. var. septentrionalis (Fern. &
Wieg.) Fern.) -- Sweet Scabious, White Top -- Stem rid-
ged with 3 lines of decurrence from each leaf, one for
the mid-nerve and one for each margin. Shallow-rooted
and + biennial. Herbage rough=-pubescent. Leaves entire
or serrate, attenuate to a sessile base, the lower petio-
late. Inflorescence becoming a lax and broad corymb.
Central head larger, flowering first, eventually overtop-
ped by the newer heads. Ligules white, filiform, nume-
rous. Mid summer. Open woods. -=-NF, NS-BC, US, Eur.

Usually subdivided in two species on what seems to
us to be an essentially arbitrary basis.

19. 5, hyssopifolius Mx. var. hyssopifolius- ~
Wild Daisy -- Heads few or only one, borne on a very

135 ERIGERON

52 RAAT O1i,0-G LA Vol. 23, no. 1

long and subnaked peduncle. No basal leaves, lower lea-
ves smaller, middle leaves linear, mostly 2-3 cm long,
about 3 mm wide and subtending short sterile shoots,

thus the plant is much more heavily leafy towards the
middle. Stem glabrous to sparsely pubescent. Mid sum-
mer. River banks and wet openings in coniferous forests.
--K-Mack, NF, NS, NB-Alta-(neBC), US.

A highly localized sinolaurentian variant, var.
villicaulis Fern., is a smaller plant, abundantly pubes-
cent on the stem, the bracteolate peduncle about as long
as, or longer than, the leafy part of the stem.

20. 5. lonchophyllus Hooker (E. minor (Hooker)
Rydb.) -- Inflorescence becoming racemose. Shallowly
rooted biennial. Stem hirsute. Leaves commonly about
3 mm wide. glabrous on both faces, long ciliate towards
the base, linear, the lower very long linear, but some
of the basal ones oblanceolate and petiolate. Ligules
white, + 1 mm long. Mid to late summer. Pioneer on wet,
disturbed soils. --K-Aka, Q-BC, US, SA.

21. E. Scotteri Boivin (E. acris var. asteroides

FE RRR = —_— ———_—
X uniflorus var. unalaschkensis AA.; E. acris var. debi-
lis X humilis AA.; E. Evermannii AA.; EB. vagus AA.) --
Small and monocephalous, similar to E. pallens and E.
uniflorus, but much less pubescent than either, at least
the larger leaves glabrous above. Perennial, 2-6-(12) cm
high, the stems few or commonly solitary. Leaves entire,
ciliate, + pubescent dorsally. Stem heavily pubescent
with hairs + 0.5 mm long, also finely glandular. Involu-
cre rather small, only 5-6 mm high; tegules ciliate, den-
sely and finely glandular on back, densely pilose at
base, sometimes sparsely pilosealeng the midnerve. Ligu-
les pink, + 3 mm long, + O.3 mm wide, rare alpine in
Banff and Jasper parks. --swAlta-seBC.

Sp. n. sectionis Trimorphaeae. Perennis. Radix
brevis, caudicibus nullis. Caulis saepius solitarius,
2-6-(12) cm alt., dense et minute glandulosus, valde pilo-
sus, pilis + 0.5 mm, lucidis. Folia rosettae petiolata,
a spathulatis anguste oblanceolata, ciliata et inferne
+ pubescentia, superne glabrescentia. Involucrum 5-6 mm
alt., viride. Tegulae circa 0.5 mm lat., sensim acumina-
tae. Ligulae roseae, + 3 mm long., (0.2)-0.3-(0.5) mm ,
lat., 40-100 in capite. Flores filiformes perpaucaée.
Flores disci 4-5 mm long., tubo + 1 mm long., luteae, lo-
bis brunnescentibus vel purpurascentibus. Pappus 3-4 mm
long. Semen + 2 mm long., anguste lanceolatum, compres=
sum, puberulens.

Typus: G.W. Scotter 9796A, Alberta, Jasper Nat.
Park, Maligne Lake, alpine, Aug. 8, 1968 (DAO). Paraty-
pi : G.W. Scotter 9776:-& 9801, eodem (DAO); J.A. Calder
24031, Lake Agnes, 1959 (DAO); L. Jenkins 7809, Jasper

ERIGERON 136

1972 Boivin, Flora of Prairie Provinces 53

Park, from Maligne Lake to Lorraine Lake, el. about 6000
feet, common among boulders on open slope, Aug. 6, 1957
(DAO); F.J. Hermann 12865, mossy north shore of Lake Loui-
se, alt. 5,700 ft., July 18, 1956 (ALTA, CAN); N.B. San-
son 1030, Lerch Valley, Ad tes3 1923 (NY); J. Macoun 11002,
78762, B.C., Kicking Horse Lake, 1886 (CAN, NY) ; Bostock,
Yoho, B.C., 1927 (DAO); Calder & Holm 24066B, 24067, Twin
Cairn Peak, BC., July 29, 1959 (DAO).

From its relatives our new species may be distin-
guished by a number of characters, such as the very small
involucre. Its very narrow and short ligules will readi-
ly separate it from most other small monocephalous spe-
cies in our area, Its light pubescence also readily sets
it apart from most other small alpine species. The length
of the pappus will separate it from small and monocephea-
lous specimens of E. acris. In the latter ligules are
usually white and the disk florets are conspicuously
overtopped by the pappus while in E. Scotteri the pappus
is shorter and slightly overtopped by the disk florets.

Despite its assignment to section Trimorphaea, it
seems that E. Scotteri is most closely related and quite
similar to E. uniflorus. However the latter is heavily
tomentose on the involucre with much longer hairs, its
tegules are longer and purplish, and its pappus is some-
what longer than the disk florets.

Though rarely collected, this new entity has been
known for quite some time and has gone through a4 surpri-
singly elaborate series of avatars.

The first collection may have been that of Macoun
at Kicking Horse Lake in 1886; it was identified as E.
acris (CAN) or E. uniflorus (NY) and reported as the lat-
ter the following year. But Macoun in 1897 mentions it
again as his only specimens of E. alpinus. In 1923 a
Sanson collection from Larch Valley was also identified,
probably at NY, as E. uniflorus. On page 239 of his mono-
graph Cronquist refers casually to hybrids of E. acris
var. debilis X humilis. This report was investigated in
1965 and turned out to be based on the above two collec-
tions. Borrowed and examined in 1966, they proved to be
rather intermediate morphologically to the postulated
parents and were incorporated in our Enumeration of 1966
as BE. acris var. asteroides X uniflorus var. unalaschken-
sis. Hermann collected it at Lake Louise in 1957, iden-

tified it to the partly glabrous E. Evermannii, aad dis-
tributed duplicates with a note that it was new to Canada.
His collection was the basis for a last minute inclusion
of the latter name in the Flora of Alberta of Moss 1959,
and in our Enumération of 1966. A Jenkins collection sub-
mitted for identification in 1958 was estimated to be an
unusually depauperate specimen of E. acris and was so
identified. ig

137 ERIGERON

Sh Payee? OL-OG TA Vol. 23, mow a

There is also in one of the Botanical Congress
guide books a mention by Porsild 1959 of E. vagus from
Mt. Temple. In our 1962 survey of the genus at CAN
there were no specimens under this names and a curso-
ry check in 1969 and 1971 gave similarly negative re-
sults. In the absence of justifying specimens, noti-
cing the absence of the putative hybrid and of E. Ever-
mannii from Porsild's list, considering that E. vagus
would be far out of range in the Canadian Rockies, and
considering that it is generally similar to E. Scotteri
in the same manner as the latter is similar to E. Ever-
mannii, we are tentatively referring the Canadian report
of E. vagus to E. Scotteri.

Thus, this attractive little species has already
accumulated a complex history and a rather large and
unwieldy series of conflicting identifications.

22. 5E. acris L. var. asteroides (Andrz.) DC.
(var. debilis Gray; E. angulosus Gaudin var. kamtscha-
ticus (DC.) Hara; E. droebachensis O. Mueller) --
Farewell-to-Summer (Vergerette) -- Ligules short and
inconspicuous like the last two and the next, but the
involucre minutely glandular. Herbage otherwise + pu-
bescent or hirsute. Leaves oblanceolate to linear, up
to 1 cm wide. Heads rarely solitary, usually few ina
variable inflorescence, commonly corymbose, sometimes
paniculate or thyrsoid. Peduncles widely spreading.
Mid summer. Wettish open spots in coniferous forests.
-K-Aka, L, NB-BC, (US, Bur).

Many other varieties also occur in Eurasia.

23. E. elatus (Hooker) Greene (E. acris L. var.
elatus (Hooker) Cronq.) -- Obviously related to the
last. Involucre hirsute, not glandular. Heads only
one or few on nearly erect peduncles in a racemose in=
florescence. Usually smaller and with fewer stem lea-
ves, First half of summer. Open wet ground on light
soils. --K-Aka, L-NF, Q-Man-(S)-Alta-Bc.

Smaller specimens may be monocephalous and should
not be confused with E. uniflorus. In the latter the
somewhat larger head is very heavily lanate with flexuous
hairs mostly 2-3 mm long. But in E. elatus the involu-
cral pubescence is much less dense and the shorter hairs
are all or mostly under 1 mn.

24, E. GANADBNSIS L. var. CANADENSIS (Conyza ca-
nadensis (L.) Cronq.; Leptilon canadense (L.) Britton)
-- Fireweed, Horseweed -- (Fausse Camomille, Herbe des
Frangais) -- Heads small and usually very numerous in
@ + cylindric inflorescence. Annual. Leaves numerous,
linear. Herbage villous. Mid to late summer. Frequent
on disturbed soil, especially in sandy or gravelly. --
Mack, (NF)-SPM, NS-BC, US, SA, Bur, (Oc).

ERIGERON 138

1972 Boivin, Flora of Prairie Provinces SS

Reputedly native in Canada but we are unconvinced.
From coast to coast we have often come across it; eve-
ry time it had the usual weedy behavior of an alien in-
vading disturbed soils. Nowhere did it recur regularly
as a normal component of a natural habitat.

A coastal plain variant, var. pusillus (Nutt.)
stat. n., E. pusillus Nutt., Gen. g: 148. 1818, is
glabrous or nearly so and its tegules are purple-tipped.

16. PSILOCARPHUS Nutt.

With the general presentation of an Antennaria, or
better a Gnaphalium, but the leaves opposite. Pappus lac-
king. Heads without involucre but subtended by a few fo-
liage leaves. Each floret enclosed by a wooly bract.

1. 2B. elatior Gray -- Woolly Heads -- Woolly
annual resembling a Gnaphalium with opposite leaves. Less
than 1 dm high, simple to dichotomously branched. Heads
small, rounded, sessile, overtopped by a number of sub-
tending foliage leaves. Mid summer. Dried slough bot-
toms, rare: Redcliff. --swAlta-swBC, nwUS.

17. ANTENNARIA Gaertner
PUSSY-TOES

White-woolly herbs with dioecious flowers. Heads
discoid, but rather showy because the tegules are peta-
loid in the upper half, white-woolly in the lower half.
Pappus of bristles, these somewhat clavate in the stami-
nate plants.

This genus has been much studied by various taxono-
mists for some three quarters of a century now and we
have not yet had a chance to evaluate some of the many
described entities. At least the following have been re-
ported for Alberta and should eventually be added to our
text either as additional taxa or as additional synonyms.

Antennaria acuta Rydb. (= Rydberg 1917).

Antennaria albescens E. Nelson (= Rydberg 1917).

Antennaria alborosea Pors. (= Porsild 1950).

Antennaria Sansonii Greene (= Rydberg 1917).

a. Inflorescence an open raceme ........e- 5. A. racemosa
aa. Inflorescence a corymb, or sometimes a solitary
head.

b. Stem very short or vestigial, not overtopping
the basal leaves .......00.+-+.-e- 17. A. dimorpha
bb. Much taller.
c. Basal and lower leaves 3-15 cm long,
lanceolate to linear, acute ........ Group A
cc. Shorter and relatively wider, usually
rounded at tip.

139 PSILOCARPHUS

56 PHY T O'L'OG Th Vol. 23, no. 1

d. Tegules coloured in the upper half,
greenish, straw, brownish or pink eo.
aiasnyale iiaveleloie ale etelele iw slwletels cle clleietelete) atmmGdaGhtty mes
dd. As Acieaditiee to light sulphur-yellow.
e. Leaves rather large, the larger
ones over 5 mm wide and usually
glabrous or glabrescent above ..
TOO OA OOUGOOOoOon Oooo COOcooecdc. (Garey (C
ee. Rather narrow, rarely over 5 mm
wide, grayish or whitish tomen-
EOSE), (ADOVEN so os cleiclslocte «1c + + ee OLOUDEE

Group A
Tufted or stoloniferous but the stolons burried.
Rather large species with the leaves all or mainly cau-
line. Rosette leaves, if present, erect.
a. Tegules glabrous to the base or the outer ones some-=
what tomentose near the base........ 3. A. luzuloides
aa. Tegules heavily tomentose in the lower 1/3;
heads larger.
b. Tufted; upper stem leaves at least half
as long as the lower ......+0+---+-. 4. A. lanata
bb. Stoloniferous; upper stem leaves much redu-
ced and many times shorter than the lower.
c. Tegules not white at tip, or the white
tip less than half the length of the
tegulle S..csic siteleiete se Jeeves ILS AS pulchernme
cc. White tips longer, more than half the
length of the tegule ..... 2. A. anaphaloides

Group B
Tegule tips variously coloured, Rosette-forming
species,

Ble SE GULSS Pa ere) <icjeiniolejeil= clelele)clejelcls ciel el-ielol nee enEGOSee
aa. Straw-coloured to brownish or greenish. .
b. Tegules pale to dark brown above the middle.
c. Involucre 6-7 mm high.
d. Basal leaves 5-10 mm wide ....

enews rene eccccccccvcescee 15. A. Russellii
dd. Narrower ....e-cevcveses-eee 18. A. alpina

cc. Involucre only 4-5 mm high; plants
smaller ....sscccccccccceee 16. A. umbrinella

bb. Tegules with dark=coloured and greenish tips.
e. Involucre 8-10 mm high; leaves very
narrowly oblanceolate.
f. Herbage grayish-tomentose ...

ease vice cesssecceceie se: GOe, Ay Angustate
ff. Green and nearly glabrous ...
Cer escecrsecrccsccsecs Che Ae QGiebueiee

ANTENNARIA 140

1972 Boivin, Flora of Prairie Provinces 57

ee. Heads smaller, the involucre only
4-6-(7) mm high; leaves oblanceolate,
mostly 3-5 mm wide.
ge Typically monocephalous; tegules
strongly SquarrOSe ..-ees-eee
seecscccccsscacee 19. A, monocephala
gg. MoseLy with 3-5 heads; tegules
appressed ....-ccecceee 18. A. alpina

Group C
Rosette forming species. Tegule tips all, or at
least the inner, white, or in one species light sulphur-
yellow. Basal leaves rather broad, the larger ones
mostly over 1 cm wide and mostly glabrous or glabrescent
above.

a. Involucre 4-7 mm high.
b. Basal leaves ovate or obovate, 1-2 cm wide when

Full Grown o.cccacccccacsesescsecs/s Aa Denikeana
bb. Spathulate or oblanceolate and all or “mostly
O.5-1.0 cm wide s..ccaccccecscece 9. A. neodioica

aa. Involucre 7-11 mm high.
c. Rosette leaves 1.5-5.0 cm wide ... 6. A. Parlinii
cc. All or mostly narrower.
d. Basal leaves cuneate-oblanceolate,
gradually narrowed at base, not dis-
tinctly petiolate; new rosettes not
developed until fruiting time .cccoo.
siaca ola sake oye nies olay spice misersiatiaiaie ats wh O otrtin CIOME Lb.
dd. Blade + obovate and abruptly narrowed
to a winged petiole; new rosettes pre-
sent at flowering ...-sseccee Ge A. neodioica

Group D
Like the last, but the leaves more tomentose. Whi-
tish or grayish above, and narrower, all or mostly less
than 5 mm wide.
a. Heads rather large; involucre 7-10 mm high ...-.

PR a wainnc niet SAR On sense dirien tae. Os Le SDELCE
aa. Heads smaller; involucre 5-7 mm high.
b. Reduced plants, less than 5 cm high .......

mina WAKE aime aw ainbbiie 6's ga. « sala’ sh ate etek RADI Lee.
bb. Geer well over 1 dm high.
c. Basal leaves obovate-spathulate, the
blade less than 3 times as long as
WELOG. «cine os skin o Kin Samara ke BLO sos see Ueto LILA
cc. Narrowly oblanceolate.
d. Middle and upper stem-leaves ending
in a scarious flat and glabrous ap-
pendage; tegule tips dirty white...

Soe ¢ Geecaevws AGis aee GG GE 2 2 a 2 8 a8 14. A. isolepis
141 ANTENNARIA

58 Pony? O°L'O'G Ik Vol. 23, HO.

dd. All or nearly all stem-leaves subu-
late at tip; tegules tips white ...

LBs inat'al ein! a MRMaNG tel otal eRe Vober alars 12. A. corymbosa

te es pulcherring (Hooker) Greene -- Our largest
species and with the longest foliage leaves. Stoloni-
ferous, but the stolons much elongate and underground,
the plant thus not carpet-forming. Tomentose throughout.
Stem 3-6 dm high. Lower and basal leaves petiolate, nar-
rowly lanceolate, the blade 5-15 cm long. Involucre 7-9
mm high. Tegules dirty green or dirty brown to white at
tip, the basal part green and white-tomentose, the middle
part dark brown. Just before mid summer. Wet and open
clay soils in the coniferous forest regions. --K-Aka,
Q-BC, wUS, (Bur).

2. A. gnaphaloides Rydb. --Very much like the
last but somewhat smaller and the tegules broader and
more conspicuous, the white tips being more than half
the length of the tegule. Involucre smaller, 5-7 mm
high. Early summer. Mountane prairies and open Lodge-
pole woods: Cypress and Rockies. =--Aka, swS-BC, nwUS.

3. A. luguloides T. & G. -- Resembles the last
two, but the heads smaller and less pubescent. Stem 2-4
dm high. Short stoloniferous. Leaves less than 1 cm
wide, mostly linear. Heads numerous. Involucre + 4 mm
high, lanate at the very base only. First half of sum-
mer. Rocky alpine slopes in Waterton. --swAlta-sBC,
nwUS.

4, A. lanata (Hooker) Greene -- With the habit of
the last 3 and with elongate lanceolate leaves, but loo-
sely tufted and loosely lanate throughout, especially
so in the inflorescence, 1-3 dm high. Ligule tips
squarrose, brown to greenish black. Mid summer. Alpine
meadows. --SwAlta-BC, nwUS.

5. A, racemosa Hooker -- Inflorescence a loose ra-
ceme, the lowest peduncle 2-3 cm long. With long super-
ficial stolons. Basal leaves ovate, green above and
mostly 2-3 cm wide. Stem leaves oblong-lanceolate. Har-
ly summer. Montane Pine woods. -- swAlta-BC, nwUS.

6. A. Paglinii Fern. var. Parlinii (A. munda Fern.)
-- Rosette leaves rather large, obovate, mostly 2-3 cm
wide. 2-4 dm high. Heads 4-8 in a rounded corymb. Long
stoloniferous and forming dense carpets, the new rosettes
not fully grown till fruiting time. Dry open places;
rare: Indian Bay. --NS, Q-seMan, US.

In the magnilacustrine var. Farwellii (Greene) Boi-
vin the leaves are cuneate obovate, + truncate, and the
pappus is somewhat shorter, about 6 mm long.

7. A. Denikeana Boivin (A. plantaginifolia AA.)

-- Similar to A. Parlinii, but the heads smaller and the

ANTENNARTIA 142

1972 Boivin, Flora of Prairie Provinces 59

leaves permanently grayish tomentose above. Involucre
only 4.0-4.5 mm high. Late spring. Dry fields. -- sMan.

A. plantaginifolia has often been used sensu amplo,
especially be the older authors, and the justifying sheets
must be examined in each case when attempting to dispose
of old records. Its Canadian distribution appears to be
restricted to southwestern Quebec and southern Ontario.
Of the old collections by Dawson's party we have examined
sheets from Emerson (TRT) and Duffering (TRT). They were
revised partly to A. neodioica, partly to A. Howellii var.
campestris. Old collections by Bell (QK) have been revi-
sed partly to A. neodioica, partly to A. Howellii var.
athabascensis. The more recent report of Scoggan 1957
was partly based on A. Denikeana.

8. A. Howellii Greene var. Yowellii (A. neglecta
Greene var. Howellii (Greene) Cronq.) --One of the 3 com-
mon species, the one which forms loose carpets with the
leaves green and glabrous above. 2.0-4.5 dm high, the
basal leaves 2.0-4.5 long and (0.8)-1.0-1.5 cm wide. Mid-
dle and upper stem leaves commonly ending in 4 scarious
appendage. Spring. Dry Pine woods and mixed forests,
--sY, S-BC, US -- var. athabascensis (Greene) Boivin --
Less than 2 dm high at Flowering, but elongating to 3.5
dm at maturity. Rosette leaves narrower, mostly 2-3 cm
long, but only 0.5-1.0-(1.2) cm wide. Middle and upper
stem leaves ending in a scarious appendage. Prairies and
Aspen bluffs. More or less transitional to the next spe-
cies. --sMack, Man-BC -- Var. campestris (Rydb.) Boivin
(A. campestris Rydb.; A. canadensis AA.; A. neglecta AA.;
A. racemosa AA.) -- Smaller and the stem leaves with sca-
rious or subulate tips. Less than 2 dm high. Rosette
leaves shorter, only 1-2 cm long and 0.5-1.0 cm wide.
Steppes and dry hills. --sMack, wO-BC, US.

In A. Howellii Greene f. concolor (Piper) stat.
ne, A. concolor Piper, Contr. US, Nat. Herb. Jl: 604.
1906, the leaves remain somewhat tomentose above. We
know of no specimen from our area and we suppose that an
Alberta report by Moss 1957, querried by Boivin 1967,was
merely a speculative entry.

A. Howellii Greene var. athabascensis (Greene) stat.
n., A. athabascensis Greene, Ott. Nat. 19: 197. 1906.

A. Howellii Greene (May 19, 1897} “Var. campestris
(Rydb.) stat. n., A. campestris Rydb., Bull. Torr. Bot.
Club 24: 304, (June 29, 1897}.

9. A. neodioica Greene var. negdioica (A. neglecta
Greene var. attenuata (Fern.) Cronq.); A. obovata E.
Nelson; A. oxyphylla Greene; A. petaloidea AA.) -- Like
the last, but the leaf blades shorter, more abruptly con-
tracted into a winged petiole, more permanently tomentose
above. Long stoloniferous and forming loose carpets with

143 ANTENNARIA

60 PAY? 0-L'(O°G Ts Vol. 23, no. l

the new rosettes already full grown(but not fully spread
out) at flowering time. Leaves up to 1.5 cm wide, but
more commonly less than 1 cm. Winged petiole at least
half as long as the blade. Involucrum 7-9 mm high. Late
spring. Dry, open woods, --NF-SPM, NS-BC, US -- Var.
Randii (Fern.) Boivin (A. canadensis Greene; A. neglecta
Greene var. Randii (Fern.) Cronq.) -- Leaves glabrous
above, even when young -- (NF, NS-PEI)-NB-Q-(nMan, neUS).

Var. Randii (Fern.) stat. n., A. canadensis Greene
var. Randii Fern., Proc. Bost. Soc. Nat. Hist. 28: 246.
1898.

Further west there is also a local variant, var.
chlorantha (Greene) stat. ne, A. chlorantha Greene, Ott.
Nat. 18: 38. 1904, with a more deeply coloured involu-
cre, the tegules being of a rather dark green towards the
tip. Still known only from Chilliwack, B.C.

All Manitoba specimens (DAO) reported as A. peta-
loidea Fern. have since been revised to A. neodioica.

An Alberta report of A. petaloidea by Raup 1935
was based on a sheet from Pine Lake (CAN) since revised
to A. Howellii.

An Alberta report of A. canadensis by Raup 1935
has not been investigated but is held to be improbable.

10. A. parvifolia Nutt. var. parvifolia (A. arida
Nelson; A. microphylla Rydb.; A. nitida Greene; A. rosea
Greene var. nitida (Greene) Breitung) -- Just before an-
thesis the stem is recurved downwards and the inflorescen-
ce is drooping, soon to become erect. Tegules tips often
tinted in sulphur yellow. One of the 3 common species,
the one with the smaller leaves and the denser carpet,
the stolons being very short. Stems 1-3 dm high, the her-
bege whitish or grayish tomentose throughout, including
the upper leaf surfaces, somewhat glandular in the inflo-
rescence. Basal leaves (0.5)-0.8-1.0-(1.5) cm long, 6 mm
wide or less, spatulate, the stem leaves 1.0-1.8 cm long.
Involucre 5-7 mm high. Staminate plant similar, somewhat
smaller and about as common as the pistillate plant. (Sta-
minate plants are rare or unknown far most of our species.)
Early summer. Common in prairies and steppes. --K-Aka,
O-BC, US -- Var. bracteosa (Rydb.) Boivin -- Larger, 3.0-
4.5 dm high, the stem leaves + 2 cm long and the involu-
cre + 7 mm high. Rare, Cypress Hills, --swS-seAlta, (US).

On the application of the names A. aprica, A. micro-

hyllea, A. parvifolia and A. rosea, see Boivin 1951 and
1953's

ll. A. posea Greene var. rosea -- Conspicuous by
its tegules variously tinted in light pink to cherry red.
Stems 1-4 dm high, the inflorescence nodding before anthe-
Sis like the last. Basal leaves (0.8)-1.5-2.0-(2.5) cm
long, including the ill-defined petiole, oblanceolate,

ANTENNARIA 144

1972 Boivin, Flora of Prairie Provinces 61

mostly 2-3 mm wide arid nearly erect; stem leaves linear,

+ 2 mm wide. Otherwise similar to A. parvifolia. Early
summer. Open woods and prairies. -- K-Aka, wcQ-nO-KC,

US -- F, decipiens Boivin -- Tegules white. A rare local
form not to be confused with the previous species. Note
the narrower and more erect leaves of A. rosea. --¥%Alta-
BC -- Var. jmbricata (Nelson) Boivin -- Leaves larger, the
basal ones obovate-spatulate, 4-6 mm wide; stem leaves
oblong-lanceolate, 3-5 mm wide. Cypress Hills and Rockies.
--swS-Alta, US.

12. A. cor E. Nelson -- Rather similar to
the last, but the tegules not tinted and the basal lea-
ves narrower and more elongate, linear-oblanceolate and
commonly 2,5-4.0 cm long. Barly summer. Open montane
forest. --swS-Alta, wUS.

Perhaps only a minor variant to be consolidated
with A. rosea f. decipiens.

13. A. pprice Greene var. aprica (A. parvifolia
sensu Cronq., etc.; A. parviflora sphalm.) -- The lowest
of the 3 common species, short, with large heads, and lea-
ves equally tomentose on both faces. Stem commonly + 1 dm.
high, stiffly erect. Basal leaves (0.6)-1.0-(1.5) cm long,
cuneate-oblanceolate. Involucre 7-10 mm high. Late
spring. Common in prairies and steppes. --w0-BC, US, (CA)
-- F. roseoides Boivin (A. parviflora Nutt. f. roseoides
(Boivin) Breitung) -- Tegules pinkish at tip. -- S-BC,

US -- F. byynnea Boivin (A. parvifolia Nutt. f. brunnea
(Boivin) Breitung) -- Tegules strawbrown at tip. Boisé
Coteau. -- swS -- Var. minuscula Boivin -- Smaller, only
1-3 cm high, and the involucre only 5.5-7.0 mm high. Ra-
re: Touchwood Hills. --sS.

Var. minuscula (Boivin) stat. n., A. minuscula
Boivin, Nat. Can. 80: 122-123, 1953.

14. A. isolepis Greene -- Differenciated from the
last 4 by the scarious appendages of its middle and up-
per stem leaves. Said appendages largest and most noti-
ceable of all our species. Stem 1-2 dm high; herbage
grayish-tomentose. Basal leaves + 1 cm long, oblanceo-
late. Perhaps nodding before anthesis. Tegule tips
transitional to the subsequent species: the outer squar-
rose and brownish, the inner white but somewhat finely
speckled in brown. Mid summer. Dry, sandy or gravelly
arctic tundra. -- K-Mack-(Y)-Aka, L. nQ, (nMan, nBC).

15. A. Rassellii Boivin (A. oxyphylla AA.) -- Te-
gule tips golden brown to straw=-coloured. Otherwise
Similar to A. neodioica, the leaf blades obovate and
abruptly narrowed to a winged petiole, the upper surface
permanently tomentose. Heads slightly smaller, the in-
volucrum only 6-7 mm high. SHarly summer. Dry hills and
open woods: Cypress Hills. --swS, wUS.

145 ANTENNARIA

62 PubsYoT .O.L.0.G: TA Vol. 23, nos 1

The name A. oxyphylla has been used in a rather
wide variety of meanings. Greene's original description
seems to correspond to a phase of A. neodioica in which
the leaves are more heavily tomentose above. Russell
1954 and Breitung 1954 were dealing with A. Russellii.
Porsild 1950 seems to refer to a minor segregate of A.
rosea, at least as far as his northern-most specimens are
concerned. And our own 1960 report for Cranbrook in
British Columbia was based on specimens we now place in
A. Howellii var. athabascensis. Raup's 1936 and 1947
reports have not been investigated.

Sp. n., A. oxyphylla sensu Russell 1954, sensu
Breitung 1954. Superficialis et coloniam laxam evolvans.
Caulis 2-3 dm alt. Stolones procumbentes, radicantes,
2-6 cm long., rosettam novam gaudentes aetate florendi.
Folia inferne albo-tomentosa, superne griseo-tomentosa 3
rosularia 1.5-2.5 cm long., late radiantia, lamina obova-
ta 5-10 mm lat., ad summas rotundata, mucronulata, ad
basas in petiolum alatum angustata; caulinaria ad summas
subulata, nec appendiculata nisi interdum superiora 1-2,
Inflorescentia rotundata-corymbosa ex 5-6 capitulis. In-
volucrum 6-7 mm alt., tegulis ad summas brunneo-stramineis.
Planta mascula mihi ignota. Type: A.J. Breitung 4414,
Cypress Hills Park, open Pine and Aspen woods, July 7,
1947 (DAO).

16. A. umbrinella Rydb. (A. aizoides Greene) --
Heads rather small and the tegule tips brownish. About
1 dm high, the leaves equally grayish=-tomentose on both
faces. Stolons short, forming small dense carpets of
rather short and broad leaves, mostly obovate. Lower
stem leaves similar, but narrower. Involucrum 4-5 mm
high. Late spring. Gravel slopes and shale slides in
the mountains: Cypress Hills and Rockies. --swS-seBC,
US.

17. A. dimorpha (Nutt.) T. & G. -- Scapose or near-
ly so and forming dense and exclusive patches. Perennial
from a taproot. Basal leaves 1.5-2.5 cm high, oblanceo-
late, overtopping the single head. Tegule tips brownish.
First half of spring. Eroded steppes, often pioneering.
Rare. --swS-sAlta-BC, wUS.

18. A. glpine (L.) Gaertner var. alpina -- Tegule
tips greenish to dirty green or dark green. Short sto-
loniferous and forming small and dense carpets. Stem
(0.5)-1.0-(1.5) dm high. Herbage grayish tomentose,
the rosette leaves discolour, very densely tomentose be=
low, but glabrescent above, the older ones greenish and
glabrous or nearly so on the upper surface. Middle and
upper stem leaves conspicuously appendiculate. Involucre
usually 5-6 mm high. Mid summer. Alpine meadows and
summits, rare. -- G-(F-K, L), nQ, swAlta, Bur -- Var.

ANTENNARIA 146

1972 Boivin, Flora of Prairie Provinces 63

canescens Lange (var. media (Greene) Jepson; A. canes-
cens (Lange) Malte; A. media Greene; A. mucronata E,
Nelson; A. subcanescens Ost.) -- Leaves permanently
whitish tomentose on both faces. Tegules tips brown to
greenish, as in var. alpina. Much more common. -- G-
(F) -K-Mack-(Y)-Aka, L, Q, swAlta-BC, wUS.

9. ff. monocephale DC. var. monocephala -- Tegu-
les tips also greenish and otherwise much as in A. alpi-
na, but typically monocephalous, sometimes with 4 se-
cond and smaller head. Stolons short, tending to form
small tufts, or the plants sometimes solitary. Rosette
leaves green and glabrous or nearly so above. Tegules
strongly squarrose and very dark green. Leaves whitish-
lanate below, glabrous or nearly so above. Sarly sum-
mer. High alpine. -- wMack-Aka, swAlta-eBc.

In the Alaskan var. exilis (Greene) Hultén the
tufts are laxer, the stolons being up to 5 cm long, and
the leaves are white tomentose on both faces, with 4
more elongate petiole.

20. A; angustata Greene -- Tegule tips greenish
like the last two, But the heads larger and the narro-
wer leaves longer. Not stoloniferous, but tufted, the
new shoots being short and ascending. Leaves linear-
oblanceolate, tomentose, commonly glabrescent above.
Heads usually solitary. Involucrum 8-10 mm high,
lightly to heavily lanate towards the base. Early sum-
mer. Alpine slopes. --(G)-F-Mack-(Y-Aka), L, nQ, swAltae-
(eBC).

21. §- glabrata (J. Vahl) Greene -- Similar to
the last, of which it is perhaps only a rare phenotype.
Herbage, and especially the rosette, much less pubescent,
green and glabrous to merely lightly tomentose. (Early
summer?). Wettish alpine slopes. -- G-(F-K)-Mack,
swAlta.

18. ANAPHALIS DC.

EVERLASTING
White-woolly and the tegules petaloid like the
last and the next, but the flowers unisexual; the pis-

tillate and staminate present together in each head.

1. A. margaritacea (L.) B.& H. (var. subalpina
Gray) -- Straw-Flower, White Daisy (Mortelle, Immortelle)
-- Like a large Antennaria without basal leaves but with
long and numerous stem leaves. Stoloniferous, 3-8 dm high,
virgate. Leaves 5-15 cm long, linear-lanceolate, green
and often floccose above, revolute. Heads numerous and
showy in a corymbiform inflorescence. Tegules milky-
white, strongly contrasting with the darker center. Mid
summer. Light soils in semi-open Coniferous forests,
--wMack, Aka, L-SPM, NS=-BC, US, Bur.

147 ANAPHALIS

6h Pt TO, 0-G A Vol. 23, no. 1

Quite local across our area: southeastern Manitoba,
Cypress Hills and Rocky Mountains. We were unable to
substantiate a report from Cutknife, Sask., by Fraser
1944, Russell 1954, and Breitung 1957.

Many varieties have been proposed, based mainly on
size of plant, number, size and width of leaves, and den-
sity of pubescence. We are not yet convinced that these
characters are sufficiently correlated inter se and with
@ well defined and individualized distribution to justify
taxonomic recognition. Some of the variation could be
ecologically conditioned.

19. GNAPHALIUM L.

CUDWEED
Lanate and with petaloid tegules like the last two.

Not dioecious. All flowers either perfect or pistillate
and both types present in each head. Our species all
annual.

a. Stem simple below the terminal corymb or panicle.
b. Stem glandular-viscid ........c0.. 3. G. viscosum
bb. White-tomentose, not glandular...cecee
Soca ee siiereteviedecescscevens He GoM tGl Oommen
aa. Stem much branched and leafy under the head
clusters.
c. Tegules acutish and dirty green to
brownish in the upper half ..... 1. G. uliginosum
cc. Rather rounded at tip, the inner hyaline
in the upper half. ...+.wecccosionss Cn Ge apAluebee

1. G. ULIGINOSUM L. -- Wartwort -- Branchy annual
with numerous small terminal clusters much overtopped by
the surrounding foliage leaves. 2 dm high or less. To-
mentum on the stem thinner than the thickness of the stem
itself. Leaves oblanceolate to long linear. Summer.
Infrequent in exundated places. -- (G), Mack-(Y-Aka), L-
SPM, NS-BC, US, Eur.

Specimens have been checked from Angusville, Rose-
town Paradise Hill, Saskatoon, Medicine Hat and Fort
Saskatchewan. Such sporadism almost surely denotes an
introduced entity.

2. g& palustre Nutt. -- Similar, more woolly and
the leaves broader. Tomentum looser, especially upwards,
becoming thicker than the width of the stem and branches.
Leaves ovate to oblanceolate, becoming shorter and broa-
der in the inflorescence and not overtopping the heads
so much. Mid summer, Marshy depressions. -- S=-BC, US.

3. G. yiscosum HBK. (G. Macounii Greene) -- (Po-
verty-Weed) -- Leaves oblinear, discolour, decurrent for
3-10 mm. Tufted biennial. Stem and upper leaf surfaces

densely glandular-pubescent, lower leaf surfaces
GNAPHALIUM 148

1972 Boivin, Flora of Prairie Provinces 65

and inflorescence white tomentose. Heads numerous in 4
dense and lightly coloured inflorescence. Tegules light
yellow to nearly hyaline. Rare in forest openings: Car-
bondale. -- NS-O, swAlta-BC, US.

G. Macounii was reported by Budd 1957 and 1964
from southern Manitoba, but this may have been a lapsus
calami as the only sheet found at SCS was the Carbondale
specimen described above.

4. G. microcephalus Nutt. (var. thermele (Nelson)
Cronqg.) -- Tufted perennial with numerous, small, whitish
heads. Stems 3-4 dm high, + decumbent at base. Stem
leaves gradually shorter upwards and grading into the
short ultimate bracts subtending the heads. Just after
mid summer. Dry foothill gravels in Waterton. --swAltae-
sBC, wUS.

Re G. obtusifolium L. reported from Manitoba by
Gleason 1952, see comment under Buchlo# dactyloides. A
further report by Budd 1957, 1964 is presumably based on
Gleason's as no corresponding specimen could be located
at SCS in 1967.

20. ADENOCAULON Hooker

Involucral bracts few, only 4 or 5. Heads without
ligules, without chaff and without pappus.

1. A. bicolor Hooker -- Pathfinder, Silver-Green
-- Leaves large, deltoid-cordate and lanate below, rather
suggesting those of Petasites vitifolius. Stem leafy.
Petioles winged. Inflorescence a diffuse panicle, almost
bractless. Achenes few and not enclosed by the very small
and reflexed involucrum. Mid summer. Moist montane woods
in Waterton. --(0), swAlta-sBC, US.

21. IVA L.
MARSH- ELDER
Similar to the next two, but the involucre not be-
coming indurated nor spinescent at maturity. Heads small
and discoid with a chaffy receptacle. Pappus lacking.
Main leaves opposite, the upper alternate.

a. Heads solitary in the axils......... 1. 1. axillaris
aa. Heads numerous in a panicle of racemes .....

@eeceeeecesreceereseceeceseer arses eose es Geece Ze i. xanthiifolia
l. J. axillaris Pursh var. axillaris -- Poverty-

Weed -- Heads solitary and drooping on recurved pedicels.
Branchy herb with lanceolate to linear leaves, the main
ones opposite, the upper alternate. Herbage inconspi-
cuously glandular, not punctate. Leaves nearly glabrous
on both faces, becoming pubescent towards the edges. Mid
summer. Alkaline soils, sometimes aggressive in cultiva-
ted or disturbed ground. == sMan-Alta, cUS -- Var. robus-

149 IVA

66 Pon YT O/OG IA Vol. 23, mos it

tior Hooker -- Herbage glandular-dotted in yellow or
brown and abundantly pubescent. Leaves lanceolate to
elliptic. Wood Mountain. -- scS, scBC, wUS.

2. I. xanthiifolia Nutt. (Cyclachaena xanthii-
folia (Nutt. ) Fres.) -- Coarse annual with large, ovate
and irregularly serrate leaves. Herbage + scabrous,
but the stem smooth below. Leaves paler beneath, the
main ones opposite, the upper alternate. Late summer.
Exundated shores of saline waters, invading disturbed
soils and waste places. -- NS-BC, US, Bur.

Apparently native around sloughs from southwes-
tern Saskatchewan westward to the Rockies, a casual
adventive elsewhere.

22. AMBROSIA L.
RAGWEED

Heads unisexual, the staminate ones in long termi-
nal racemes, the pistillate ones axillary and strongly
modified, containing a single flower without corolla, the
tegules fused together into a pod=like bur which is aci-
cular in the upper part, and becomes semi-woody. Heads
rayless and with filiform chaff on the receptacle. Pappus
none.

a. Leaves both trifid and serrate or merely

SCSRNESD Aadocsoeeae sh ooducnccaboncode 1. A. txatida
aa. Leaves pinnatifid to bipinnatipartite.
b. Perennial; leaves opposite .......sceeec

cece ec ee ee ee cece veseeee 3. A. pSilostachya

bb. Annual; leaves all or mostly alternate.
ce. Involucre of the staminate flowers

QNELTE ssccevscvcccnece, & As) arteninianalae
cc. Deeply lobed; stem acicular-hispid ..
Cette cece cece cer eceeecees 4. A. acanthicarpa

1. A. toifiida L. var. trifida (f. integrifolia
(Muhl.) Fern.) -- Great Ragweed, Buffalo-Weed (Grande her-

be a poux) -- Main leaves large, opposite and trifid.

Tall and coarse annual, usually little branched. Leaves
sometimes ovate and merely serrate. Petiole winged. Cor-
ners of the achene prolonged into short points. Second
half of summer. Riverward edge of galerie-forests and
casually as a weed indisturbed places. -- NS-BC, US, Bur.

Seems native from southeastern Saskatchewan east=
ward to southwestern Quebec; an uncommon adventive else=
where.

Leaf shape is rather variable. Stem leaves are ty-=-
pically trilobed to tripartite. Upper leaves, lower lea-
ves, and leaves from depauperate individuals may be unlo-
bed (= f. integrifolia). South of us there is another va-
riant, var. texana Scheele, in which the petioles are

AMBROSIA 150

1972 Boivin, Flora of Prairie Provinces 67

wingless and the achenes tuberculate rather than acicu-
lar on the angles.

2. A. grtemisiifolia L. (var. elatior (L.) Des-
courtils, f. villosa Fern. & Grisa; A. elatior L.) --
Ragweed, Hogweed (Herbe A poux, Roupie) -- An annual
inconspicuous except for the long terminal racemes of
staminate heads. Leaves + bipinnatipartite, narrowly
decurrent on the petiole, the lobes mostly 2-3 mm wide,
the pubescence short and nearly appressed. Stem pubescen-
ce very long, spreading hirsute. Fruit with a row of
spreading spines near the top, these not very sharp. Af-
ter mid summer. Common as a weed, but also native around
dessicating sloughs west of the Missouri Coteau. -=- NF,
NS-BC, US (CA), SA, Bur.

3. A. psilostachya DC. var. coronopifolia (T. &
G.) Farw. (A. coronopifolia T. & G.) -= Much like the
last, but perennial by deeply buried rhizomes and the
leaves not so deeply divided. Stem leaves all opposite,
merely pinnatipartite, the lobes mostly around 5 mm wide,
decurrent on the winged petiole. Stem pubescence like
that of the leaves, only a little longer. Fruit someti-
mes spineless, but mostly with a crown of not very long
and not very sharp projections. Mid to late summer. Oc-=
casional in somewhat alkaline prairies and shores. NS-
PEL, Q-BC, (US, CA), Bur.

Apparently present in Alberta only as a railway weed
at Craigmyle where it was collected by Brinkman in 1922
(ALTA). This collection is the justification of the re-
port by Moss 1959. An earlier report by Groh 1944 was
based on Macoun 949, Red Deer Lakes, July 21, 1879 (DAO).
But the specimen belongs to A. artemisiifolia and the Red
Deer Lakes (or Coteau Lakes) are in Saskatchewan, 10-15
miles southwest of Outlook.

The typical phase is Mexican and is reported to be
more finely pubescent on the staminate involucrum.

4. £- acanthicar Hooker (Franseria acanthicarpa
(Hooker) Cov.) -- Sandbur =-- Fruit a bur with many and
very sharp spines. Annual and mostly similar to A. arte-
misiifolia, but very rough pubescent, the stem almost
acicular=pubescent. Bur usually with a terminal spine and
two rings of lateral ones. Second half of summer. Pio-
neer onWind eroded sandhills; also adventive at Saskatoon.
--swMan-sAlta, US.

23. XANTHIUM L. CLOT BUR
Fruit a bur formed of fused tegules and covered
with numerous acicules hooked at tip. Heads unisexual,
the staminate ones few and not obvious. Pistillate ones
reduced to 2 flowers, maturing into a bilocular woody bur.

152 XANTHIUM

68 PsBiYeT GEOG: Tok Vol. 23, no. 1

a. Ferociously armed with axillary spines....ceo

Corer ccc ccc ccccccceccrcccnccsceossececoe Le X. Spinosum
aa. Not spiny except the burs ...ec..-. 2. X. strumarium

1, X. SPINOSUM L. -=- Cocklebur, Bathurst-Bur (Peti-
te bardane) -- Very spiny herb with numerous, yellow,
branched, very sharp and very long spines, Leaves lobed,
whitish=-tomentose below. Bur smaller than in the next.
Late summer. Rare weed: Steelman. --O, S, BC, US, SA,
Bur.

2. X. strumarium L. (var. canadense (Miller) T.

& G., var. glabratum (DC.) Cronq.; X. canadense Miller;
X. commune Britton; X. echinatum Murray; X. glandulife-
rum Greene; X. italicum Moretti) -- Cocklebur (Gratia,
Glouteron) -- Fruit a bur about 2 cm long and covered
with catchy acicules, the top two acicules stronger. Ve-
ry scabrous annual. Leaves deltoid-ovate, irregularly
serrate. Mid summer. Shores; weedy on occasion. --NS-
BC, US, (CA, SA), Bur, (Oc).

This shore plant is at the origin of the name of
the Riviére aux Gratias. The latter is the French topo-
nym of the lower half of the Boyne River, the upper half
being the Riviére des Ilets de Bois.

24, HELIOPSIS Persoon OX-EYE

Resembles Helianthus both habitally and technical-
ly, but the receptacle conical, hence the flower center
is raised. Also the peripheral florets are fertile (ste-
rile in Helianthus). Rays marcescent, like Zinnia to
which it is related.

1. H. helianthoides (L.) Sweet var. sgabra (Dunal)
Fern. (ssp. occidentalis Fischer) -- Ox-Bye--Monocephalous
and showy perennial, resembling Helianthus, but the ligu-
les sulphur (rather than orange) yellow. Very scabrous
virgate herb with ovate, opposite, serrate leaves, Pedun-
cle elongate, thickened below the head. Tegules rounded
at tip. Mid summer. Open woods and outer edge of gale-
rie-forests. --PEIl-ecS, (BC), US.

Our taxonomy differs from that of the latest mono-
graphic study by T.C. Fischer, in Ohio Journ. Sc. $8:
97-107, 1958, and is justified as follows.

The typical veriety is common in the eastern half
of the U.S.A., barely entering Canada in southern Ontario.
Its leaves are thin, triangular-lanceolate, rounded or
cuneate at base, glabrous on both faces, varying to light-
ly scabrous above and somewhat short pilose below.

The more northern and more widespread phase has
thicker and coriaceous leaves, deltoid-ovate, truncate
at base, scabrous on both faces, more strongly so above.
We are calling it var. scabra because its short and sca-

HELIOPSIS 152

1972 Boivin, Flora of Prairie Provinces 69

brous leaves seem to fit Dunal's original description
(foliis scabris, ovate oblongo ...) better than the next
variety does.

In a yet undescribed variety ranging from Illinois
to Texas the leaves are narrower but scabrous. It was
called ssp. scabra by Fischer but this interpretation is
questionable as pointed out above. This southwestern
phase may then properly be known as var. Fischeri var.
n. (= ssp. scabra sensu Fischer, nec Dunal), folis sca-
bris, ab ovato-lanceolatis anguste lanceolatis, saepius
ter longioribus quem latis. Type: D. Demaree 66484, near
Avoca, Arkansas, May 17, 1929 (DAO).

25. RUDBECKIA L. CONE-FLOWER
This, the last and the next two genera are easily
spotted by the very protuberant center of the head, be-
cause of a conical to cylindric receptacle. Disk flowers,
but not the ligulate flowers, subtended by bracts. Ray
flowers sterile.

a. Leaves pinnatifid to serrate ........ 1. R. laciniata
@a. Entire to shallowly serrate ...-.-cccece 2% Re hirta

1. RR. laciniata L. var. iniata -- Coneflower,
Golden Glow -- Very tall herb with large heads, the disk
ovoid and the ligules yellow. Often 1-2 m high. Nearly
smooth except for the scabrous leaf margins. Leaves lar-
ge, trifid to pinnatifid with serrate lobes, the upper
leaves often not lobed but ovate and merely serrate.

Heads few, corymbose on long peduncles. Second half of
summer. Galerie-forests, often along the inner (or river)
edge. --NS-sMan, US, Eur.

In our plant the leaves are glabrous above and +
strigose below, while the disk scales are only 3-4 mm
long. It is seemingly native from southern Manitoba to
southwestern Quebec. Its occurrence still further east
is probably related to its cultivation as an ornamental.
A variant from the central U.S., var. ampla (Nelson)
Cronq., has somewhat larger heads, its scales larger, +
7 mm long, and its leaves glabrous below but usually
strigose-muricate above.

2. B- hirtea L. (R. serotina Nutt.) -- Brown-eyed
Susan, Nigger-Heads (Marguerite jaune, Obéliscaire) --
Showy herb with bicolour heads, the ligules yellow, the
semi-hemispheric center purple-black. Stem abundantly
punctate in purple-brown, coarsely hirsute and 4-7 dm
high, usually virgate and monocephalous. Leaves lanceo-
late and commonly entire. Tegules nearly as long as the
ligules. Mid summer. Open places, mostly on chernozems.
--NF, NS-BC, US.

Native with us, but only an introduction in B.C.,
mainly an introduction in Bastern Canada.

153 RUDBECKIA

70 PHT On 0G TA Vol. 23, no. 1

26. ECHINACEA Moench

Like the last but the receptacle bracts spinescent
and overtopping the disk florets.

1. E, angustifolia DC. var. angustifolia -- Very
showy and very conspicuous; rather similar to the more
common Rudbeckia hirta in herbage and habit, but the lon-
ger and drooping ligules are pink, fading purple. Disk
purple brown. Just before mid summer. Bluffs of cou-
lées and sandy deltas, locally abundant. --swMan-seS,
cUS.

Sometimes treated as a variety of the more southern
E. pallida Nutt. but the material at hand shown no inter-
mediates.

Our var. angustifolia is usually monocephalous and
the stem leaves and peduncle are hirsute to hispid. More
southern plants, especially those from Oklahoma, may bear
a few heads and be more or less strigose on the leaves and
peduncles, these have been named var. strigosa McGregor.

27. RATIBIDA Raf.

Like Rudbeckia, but all florets subtended by bracts.

This and the last are perhaps not generically dis-=
tinct from Rudbeckia.

1. B- columpifera (Nutt.) Woot. & Standl. (Lepachys
columnaris (Sims) T. & G.) -- Very showy composite with
the center of the head cylindric (!) and subtended by ue
sually 4-(6) large, drooping, yellow ligules. Tufted pe-
rennial with pinnatipartite alternate leaves, Terminal
lobe largest, the lower ones sucessively smaller. Head
with brownish to purple disk. Mid summer. Frequent weed
along railroads, roads, etc., also apparently indigenous
at least in southern Saskatchewan. --O-seBC, US (CA) --
F. pulcherrime (DC.) Fern. -- Ligules purple. --Man-Alta,
US -- F. denudata Boivin -- Ligules lacking. Val-Marie
and Bowmantown. --S-Alta.

F. genudata (Boivin) stat. n., R. columnaris Sims
£. denudata Boivin, Nat. Can. 87: 46. 1960.

The area of native occurrence is not easy to defi-
ne. Generally found along roadsides, railway embank-
ments, etc. But in the extreme south of Saskatchewan it
seems to recur in some places as a normal element of
steppic vegetation. We hold no opinion about Manitoba
and Alberta, but in Ontario it is certainly an introduc-
tion.

28. BALSAMORHIZA Nutt.

Related to Helianthus, with the peripheral florets
fertile and the foliage mostly basal.

ECHINACEA 154

1972 Boivin, Flora of Prairie Provinces 71

1. B. sagittata (Pursh) Nutt. -- Balsam-Root --
Forming rather conspicuous rosettes of large leaves
similar to those of Petasites or Arctium. Stem monocepha-
lous and with smaller and narrower leaves. Herbage soft
tomentose, the tomentum especially dense near the head.
Basal leaves with blades at least 1 dm long and triangu-
lar-sagittate; stem leaves few, + lanceolate. Head 6-8
cm wide. Late spring and early summer. Foothill prai-
ries, --swAlta-BC, US.

Reports for Saskatchewan by Rydberg and later au-
thors do not appear to be substantiated by any actual
collection from the province.

29. HELIANTHUS L. SUNFLOWER

A basie type with large heads radiate in yellow and
pappus reduced to 2 caducous awn=scales. Receptacle chaf-
fy. Ligulate flowers sterile.

a. Annual; leaves alternate.
b. Tegules 5 mm wide or more, long caudate ..
Pier ct clale vicls’ ajo eisisiola'elele ain win slatsietelersicic wt ip, He ATIMUUS
bb. Narrower and barely caudate .... 2. H. Couplandii
aa. Perennial; leaves all or mostly opposite.
ce Tegules strongly imbricate, broadly
acute to rounded at tip .....3. H. subrhomboideus
cc. Outer tegules narrowly acute to acumi-
nate and mostly about as long as the
involucre.
d. Leaves ovate, rounded to a petiole
2-5 cm long .--scecceesseeve 6. Hs tuberosus
dd. Leaf blade oblong=ovate to linear;
petiole 1 cm long or less.
e. Leaves conduplicate and falcate
in the smaller plants; heads ra-
cemose in the larger plants ....
eee ccccceccesccoccces§ 4. H. Maximilianii
ee. Leaves flat and the heads corym-
bose; petioles long ciliate ...e.
Cor ccesccccraccsvescccses Ys He Nuttallii

1. H. ANNUUS L. cv. GIGANTEUS -- (var. macrocar-
pus (DC.) Cockerell) -- Sunflower (Soleil, Tourne-soleil)
-- Typically with a single gigantic head nodding towards
the sun. Disk 5 cm wide or over. Cultivated and casual-
ly reseeding itself, but labile. --NS-PEI, Q-Alta, US,
(Bur) -- F. lenticularis (Douglas) Boivin (H. lenticula-
ris Douglas; H. petiolaris Nutt.) -- Wild form of the
above. Herbage very rough. Leaves serrate, ovate or
deltoid-ovate, 3-10 cm wide. Disk 3-5 cm wide. Tegules
5-8 mm wide, 1.2-2.0 cm long, ovate and abruptly long
caudate. Disk purple. Mainly in late summer. Rare

155 BALSAMORHIZA

(2 Relieyer OF O1G ie Vol. 23, no. 1

native along eroded coulées; mostly a common weed, espe-
cially of roadsides. --PEI-BC, US -- F. FALLAX Boivin --
Disk florets orange. Forget. --sS.

A Manitoba report of f. fallax by Boivin 1960 is to
be discounted as it was based on a variant of cv. Gigan-
teus in which the disk florets are orange instead of the
usual deep purple.

F. lenticularis seems to be present as a native
pionneer species along a few eroded river banks of south-
western Saskatchewan and extreme southern Alberta.

2. H. Couplandii Boivin (H. aridus AA.; H. petio~
laris AA.) -- Habitally resembling the last but more deli-
cate and somewhat shiny when alive. Very branchy or with
suppressed shoots in all leaf axils. Leaves 0.5-3.0 cm
wide, triangular-ovate to triangular-lanceolate, usually
entire. Tegules 0.7=1.5 cm long, 3.0-4.5 mm wide, lan-
ceolate, slightly acuminate. Ligules shorter, only 1.5-
2.0 cm long. Mid summer. Showy on eroded sandhills;
also on disturbed sands. --O-seBC, US.

Sp. n., H. aridus auctorum, nec Rydb.; H. petiola-
ris auct. nec Nutt. Annuus scaber, (2)-4-(7) dm alt.,
ramosissimus vel fasciculiferus in axillis foliorum
praecipuis. Folia persaepius integra, ab ovatis lanceo-
lata, ad basas late cuneata vel rotundata, 2-5 cm long.,
0.5-3.0 cm lat. Petiolus 1-4 cm. Capita saepius nume=-
rosa et paniculata, interdum unicum, centro atropurpureo.
Tegulae ab ovato-lanceolatis lanceolatae, (0.7)-1.0-(1.5)
cm long., 3.0-4.5 cm lat. Ligulae 1.5-2.2 cm. Type: B.
Boivin 6682, entre Big Stick Lake et Crane Lake, espéce
pionniére sur les dunes éventrées, 28 juillet 1949 (DAO).
We have had an opportunity to examine the types of H.
aridus Rydb. (NY) and of H. petiolaris Nutt. (K). Both
belong to H. annuus f. lenticularis.

Dr. R.T. Coupland of the University of Saskatche=-
wan is the author of many important papers on our area,
including Ecology of Mixed Prairie, Ecol. Mon. 20: 271-
315), L950).

3. HH. subrhombeideys, Rydb. (H. laetiflorus Pers.
var. subrhomboideus (Rydb.) Fern.) -- Main leaves rhom-
boid-ovate to rhomboid-lanceolate. Long stoloniferous,
hardly tuberous. Heads on very long peduncles and so-
litary, or the peduncles incurved into a candelabriform
inflorescence. Tegules ciliate, glabrous dorsally, the
inner acutish, the outer often rounded, and less than
half as long as the involucre. Mid summer. Dry, open
places, usually on chernozems. =--Y, NB-BC, US, Bur.

The related H. laetiflorus was reported for Sas-=-
katchewan by Fernald 1950, but we found no correspon-
ding specimen at GH in 1965.

HBLIANTHUS 156

1972 Boivin, Flora of Prairie Provinces 73

4. HH. Maximilianii Schrader -- Smaller plants
monocephalous, grayish, the leaves conduplicate and
strongly falcate. Larger plants with the foliage not
quite so characteristic but the inflorescence elongated,
somewhat racemose and often secund. Roots tuberous in
the manner of the next. Leaves alternate in the upper
third, long attenuate at base, the middle, and upper ones
sessile, or sometimes the middle ones tapered into 4 short,
ill-defined and winged petiole. Leaf pubescence abundant
on both faces, the hairs less than 0.5 mm long and nearly
strigose; stem pubescence similar, but sometimes less
dense; leaf base not conspicuously ciliate. Involucre
1-2 cm high. Ligules about 2 cm long. Second half of
summer. Chernozems, especially around depressions, --
PEI, Q-BC, US.

Native on chernozems between Lake of the Woods
and Regina, frequently introduced between Regina and
Saskatoon, a sporadic introduction elsewhere; Swift
Current, Redcliffe, Calgary, etc.

A dot map in Brittonia 18: 74, 1966 credits this
species with a substantially more northern reach (north
to Churchill River and James Bay), than the outline gi-
ven just above. The discrepancy is apparently related
to a difference in taxonomic treatment; what we are here
calling H. Nuttallii var. subtuberosus being reassigned
by Long partly to ssp. canadensis, partly to H. Maximi-
lianii. Hence the similarity in northern limits for the
maps on pages 74 and 76 of said paper.

4X. H. Alexidis, Boivin -- Hybrid with the next
and combining various characteristics of both, such as
the inflorescence broad, but the involucre high and the
leaves conduplicate. Thornhill and probably elsewhere
also. -- sMan.

Hybr. n. Verosimiliter H. Maximilianii X Nuttal-
lii. Variabilis et exhibens notas varias parentiun,
vege: inflorescentia lata, corymbosa atque folia late lan-
ceolata, sed involucrum majus, 15-20 mm alt., folia con-
duplicata, ligulae + 2 cm long, etc. Type: J.F. Alex
121, Manitoba, Lisgar District, Thornhill, 1 mile south,
native grassland along dry waterway adjacent to highway,
Sept. 4, 1957 (DAO).

S. H. Nyuttallii T. & G. var. Nuttallii -- Often
closely resembling the last, but the leaves flat and
the heads corymbose if more than one. Stolons mostly
5-10 cm long, tuberous at tip, producing rootlets which
are tuberous towards their attachment. Leaves + 1 cm
wide, linear to linear-lanceolate, opposite except
perhaps the upper 1-3, cuneate at base, petiolate. Pe-
tioles successively shorter, the middle ones around 1 cm
long, coarsely ciliate, the cilia over 1 mm long. Stem

LST HELIANTHUS

7h PHY? OL ove WA Vol. 23, no. 1

lightly pubescent to nearly glabrous, the coarse hairs si-
milar to the cilia, but somewhat shorter. Midnerve pu-=
bescent like the stem, but the hairs still shorter. Lea-
ves heavily scabrous on both faces with very short hairs
inflated at base. Heads monochrome. Tegules 1.5 cm long
or less. Ligules 2-5 cm long. Second half of summer,
Wettish prairies and along watercourses. --wO-BC, US --
Var. subtuberosus (Britton) Boivin (ssp. canadensis Long;
H. fascicularis Greene; H. giganteus AA.; var. subtubero-
sus Britton; H. subtuberosus Britton) -- Leaves broader,
+ lanceolate, mostly 2-3 cm wide. Commoner. -- (Mack),
NF, NS, NB-BC, US -- F. yerticillatus Boivin -- Leaves
verticillate in 3's or more. Local: Candle Lake. -- S,
US -- Var. Rydbergii (Britton) Boivin -- Leaves broad

and short, ovate to narrowly oblong., less than 1 dm
long. The common phase along creeks in the steppe re-
gions. --sMan-sAlta, US.

Var. subtuberosus (Britton) stat. n., H. giganteus
L. var. subtuberosus Britton ex Britt. & Brown, Ill. Fl.
3: 425. 1898.

F. verticillatyus nom. ne, H, giganteus L. var. sub-
tuberosus Britton f. verticillatus Lakela, nom. ill.,
Rhodora 49: 21. 1947, nec H. giganteus L. var. verticilla-
tus Farwell 1927.

Var. Rydbergii (Britton) stat. n., H. Rydbergii Brit-
ton, Man. Fl. N. Stat. & Can. 993-994, 1901.

The trio H. Nuttallii, subtuberosus and Rydbergii
constitute €@ series of strongly overlapping and complete-=
ly intergradient phases; their recognition is undoubtedly
mechanical. Yet each taxon presents a certain ecological
specialization and some degree of geographical individua=
lity; we have felt justified to retain them at the varie-
tal level.

We are not happy yet about the degree and quality
of distinctiveness of var. subtuberosus from the eastern
H. giganteus L. However we are for the present retaining
them as specifically distinct as per the more common cur-
rent practice. The accepted criteria are as follows. Var.
subtuberosus: leaves all opposite, or the upper 1-(3) al-
ternate; tegules ciliate with hairs under 1 mm long and
only half as long as the petiolar cilia; leaf pubescence
similarly dense and short on both faces, although the
hairs are more strongly bulbous=based on the upper face.
H. giganteus: the upper (3)-5-(7) leaves alternate; tegu-
lar and petiolar cilia similar in size and over 1 mm long;
leaf pubescence of dense, short, very stiff and strongly
bulbous-based hairs on the upper face, but on the lower
face the hairs are not bulbous=-based,much less dense, and
obviously longer, commonly 0.5=1.0 mm long. Unfortunately
these criteria seem far from constant and discrete; it

HELIANTHUS 158

1972 Boivin, Flora of Prairie Provinces 75

might be preferable to subordinate H. Nuttallii and its
varieties to the earlier H. giganteus.

If the latter solution proves to be preferable,
the correct names of our three varieties would be as
follows. The common and widespread phase with leaves of
middling width returns to H. giganteus var. subtuberosus
Britton. The narrow-leaved phase of steppic regions be-
comes H. giganteus var. utahensis D.C. Eaton. The broad-
leaved phase of the steppic regions would require 4 new
transfer.

6. HH, BAR SEO REE L. var. subcanescens Gray -- (Es-
quebois) -- Leaves largest, 5-10 cm wide, ovate, conspi-
cuously 3-nerved, serrate, acuminate, rounded to 4 winged
petiole, opposite. Very stoloniferous, the stolons very
long and ending in a purplish potato. Mostly 1-2 m high.
Leaves somewhat velvety below, the herbage otherwise
scabrous. Heads few in a corymbose inflorescence. Late
summer. Galerie-forests. --O-seS, (US).

In the more eastern typical phase the stem leaves
are mostly alternate and scabrous on both faces.

Reports of H. divaricatus L. from Saskatchewan ha-
ve yet to be tied down to specimens actually collected
within our area,

30. COREOPSIS L. TICKSEED

Intermediate to Bidens, the pappus being of two mi-
nute teeth. Tegules dimegueth and in two rings; the in-
ner adnate at base and petaloid at tip, the outer much
smaller and free. Disk not chaffy.

1. ¢. ginctoria Nutt. (C. Atkinsoniana Douglas) --
Bye-Flower, Tickseed -- Leaves opposite and pectinati-
partite to bipectinatipartite. Biennial with rather
scanty foliage, branchy and the branches opposite. Heads
many, bicolour, the disk purple-brown, the ligules golden
yellow with a purple brown patch near the base, cuneate
with a ragged tip. Mid summer. Exundated places in
drier parts of southwestern Saskatchewan and southern
Alberta, elsewhere a casual escape from cultivation. --
swQ-BC, US.

Appears to be native in southern Alberta and in
southwest Saskatchewan, casually adventive or escaped
from cultivation elsewhere.

C. Atkinsoniana is merely a form with narrowly
winged seeds, sporadic in the range of the species, of no
particular significance, not forming a distinct population.

Coreopsis lancolata L. and C. verticillata L. were
reported by Macoun 1884 for Western Canada but this may
have been a lapsus calami for Canada West. The latter was
an alternate name for Upper Canada, now the southern part

159 COREOPSIS

76 PobY oT OL OC PA Vol. 23, nosed
of the province of Ontario.

31. THELESPERMA Less.

Obviously similar to the last but the disk chaffy.
Pappus also of 2 reduced bristles. Inner involucral
bracts adnate for at least one third of their length.

1. 2. marginatum Rydb. -- Heads discoid, its in-
volucre campanulate, of fused bracts, its lobes broadly
margined in white. Perennial herb with narrowly dissec-=
ted leaves and 1 or more heads on very long and subnaked
peduncles. (Mid summer?). Eroded hills, rare: Medicine
Hat. --seAlta, ncUS.

We know of no Canadian collections other than the
ones from The Hat and there is none other at the New
York Botanical Garden. The various reports for Saskat-=
chewan must therefore be the result of Medicine Hat
being assigned to the wrong province.

32. BIDENS L.

Achenes catchy by 2 or 4 barbellate terminal aci-
cules termed "horns" or "teeth''. Tegules in two rings
and dimorphic, the inner + petaloid.

a. Submerged aquatic with leaves divided into fi-
liform SegmentS .....sseccccesceeeceeoe 4. Be. Beckii
aa. Terrestrial and the leaves with a well defi-

ned flat limb.

b. Head radiate; achene with 4 horns .. 1. B. cernua

bb. Eradiate; achene with only 2 horns.
c. Leaves compound......e..+-+e2 3. B. frondosa
cc. Simple, merely serrate to trifid ..

ayneeNe le ssicus Onieeseacscsesececons 4e Be uteri pamtane

1. B. gcernua L. (B. glaucescens Greene) -- Stick-
tight, Pitchfork (Fourchettes) -- Heads radiate in yel=

low; however the ligule-like appendages are not deri-=
ved from the outer florets, but from the inner petaloid
tegules of the involucre. Leaves + lanceolate, sessile,
serrate. Achenes with 4 horns. Mid to late summer.
Common in wet places and shores. -- sMack, Aka, NS-BC,
US, Eur.

1X. B-. amplissimg Greene (B. Stevensonis Boivin,
nomen) -- Hybrid with B. frondosa var. puberula. Luxu-
riant annual with irregularly lobed to trifid or pinna=
tifid leaves. Rachis and petiole broadly winged. Heads
irregularly radiate. Rare: Brandon. --sMan, sSwBC.

2. 3B. TRIPARTITA L. var. TRIPARTITA (B. comosa
(Gray) Wieg.; B. connata Muhl.) -- Beggar-Ticks, Stick-
tight (Fourchettes, Cornes) -- Leaves typically petio-
late and trilobed, but very variable and ranging from

BIDENS 160

1972 Boivin, Flora of Prairie Provinces 77

merely serrate to trifid, the upper leaves sometimes mere-
ly attenuate to a sessile base. Leaves and outer tegules
scarious at margin, the latter sometimes scabrous, the
herbage otherwise glabrous. Other characters pretty much
as in the next species. Late summer. A rare adventive

of wet places: Ronalane. -- NF, NS-O, seAlta-(BC), US,
Bur.

Other reports from our area are probably to be
discounted. The Manitoba records were discounted by
Scoggan 1957. Reports of B. connata Muhl. from the Sas-
katchewan are based on a Drummond collection from Cumber-
land House. Its specimen basis has not been investiga-
ted yet, but since it has never been confirmed, it is
expected to be based on B. frondosa var. puberula.

In our typical variety the achenes are over 2 mm
wide, as contrasted with a sinolaurentian var. heterodoxa
Fern. in which the disk achenes are only 1-2 mm wide.

3. B. frondosa L. var. frondosa -- Beggar~-Ticks,
Boot-Jacks (Fourchettes) -- Main leaves compound, mostly
trifoliate, the petiole not winged. Glabrous to hirsute.
Bracts of the outer series mostly 5-8, green and folia-
ceous, longer than both the inner series and the disk.
Achenes with 2 horns, 2.5=5.0 mm long. Mid to late sum-
mer. Shores. -- NF, NS-Man, US, BC, Bur -=- Var. puberu-
la Wieg. (B. vulgata Greene, var. puberula (Wieg.) Y Greene)
-- Coarser and the heads with 10-15-(20) tegules in the
outer and longer series. Central achenes with horns 4-8
mm long. -- NS, (NB)-Q-BC, US.

A doubtful Alberta report of B. frondosa by Scoggan
1957 could not be substantiated and is herewith discoun-
ted.

4. B, Beckii Torrey (Megalodonta Beckii (Torrey)
Greene) -- Water-Marigold == Submerged and usually steri-
le herb with opposite leaves dissected into filiform seg-
ments. Emersed leaves, when present, entire to pinnatifid.
Head solitary, radiate. Achene with 3-6 horns longer than
the body of the fruit. Late summer. Quiet waters, rare:
Wildnest River and Cumberland Lake eastward. -- NS, NB-
ecS, sBC, US.

33. GALINSOGA R. & P.

Leaves opposite, the heads radiate, the pappus chef-
fy.
a. Villous with hairs 0.5-1.0 mm long.... 1. G. ciliata
aa. Glabrous or finely pubescent, the hairs
a fetrigose ies cinen dc eels. Cot ees. “SenG. parviflers
1. G. CILIATA (Raf.) Blake -- Quickweed -- A. city

weed with small heads briefly radiate in white. Villous
annual, Leaves ovate, serrate. Heads about 5 mm high.

161 GALINSOGA

78 PHY? O'L0G TA Vol. 23, no. 1

All seeds bear a pappus of scales about as long as the
body of the seed. Late summer and fall. Neglected
lawns and back lanes, rare. --NS-Man, Alta-BC, US,
(CA, SA), Bur. (Afr, Oc).

We have checked only two collections from our area,
Winnipeg and Calgary, both at DAO. The Boissevain re-
port was based on a sheet of Potentilla norvegica. The
Grand Beach and Edwin report have not been checked.

The inclusion of Saskatchewan in its range by
Frankton 1970 was apparently based on a@ specimen culti-
vated at Saskatoon (DAO), the inference being that the
initial seed supply originated somewhere within the pro-
vince.

2. G. PARVIFLORA Cav. -- Joey Hooker, Yellow
Weed -- Sufficiently similar to the first to be gene-
rally confused with it. Leaves entire or weakly crena=
te. Peripheral seedswithout pappus. A rare town weed:
Hamiota. --Q-Man, US, (CA, SA), Eur.

A rare weed in Canada; we have checked only three
Canadian collections: Sherbrooke, Bridgeport and Hamio-
ta, all DAO.

34. MADIA Molina TARWEED

Most of the tegules half-wrapped around the outer
achenes. Pappus none or much reduced,

1. M. glomerata Hooker -- Tarweed -- Heads narrow
and few-flowered, about half as wide as high. Heavily
glandular-pubescent annual, usually virgate. Heads not
very conspicuous, about 1 cm high, few, discoid or brie=
fly radiate in yellow, drying pink, the rays only 1-3,
the disk flowers also very few. Second half of summer.
Arroyos, sometimes weedy. --(Y-Aka), Q-BC, US.

Appears to be native from Saskatchewan westward,
but also occurring as an uncommon adventive. Known in
Manitoba from Souris (WIN) and Portage (SASK).

There has been a fair amount of confusion of this
species with M. sativa; all specimens named M. sativa
from eastern Canada that we have studied turned out to be

M. glomerata.
35. HYMENOPAPPUS L'Hér.

Pappus of small hyaline scales. Heads discoid. Te-
gules in one series, scarious margined, not imbricate.
Receptacle not chaffy.

l. BH, gilifolius Hooker var. polycephalus (Oster-
hout) B.L. Turner -- Leaves + bipectinatipartite. Tufted
perennial from a taproot, (1)-2-(4) dm high, more or less
white tomentose, especially the petioles, the base of the
stem and the margin of the tegules. Leaf segments + 0.5
mm wide, finely pitted and punctate in deep green. Head

MADIA 162

1972 Boivin, Flora of Prairie Provinces 79

yellow. Early summer. Local on badlands. -- swS-sAlta,
ncUS,

A highly variable species. In the latest mono-
graph it is subdivided into a very complex series of 13
intergrading varieties. Only var. polycephalus is recor-
ded as entering Canada. It is a fairly tomentose va-
riant, of medium height, average leafiness and smallish
flowers.

36. BAHIA Leg.

Not unlike the last, with a pappus of short hyali-
ne scales. But the leaves opposite and the heads redia-
te. Not chaffy. Tegules in only one series.

1. B. OPPOSITIFOLIA (Nutt.) DC. (Picradeniopsis
oppositifolia (Nutt.) Rydb.) -- A Composite with opposite
and narrowly dissected leaves. Deeply stoloniferous peren-
nial, 1-2 dm high. Densely puberulent, finely pitted and
glandular-punctate. Heads few, yellow, radiate, but the
ligules only 1-3 mm long and paler to nearly white. First
half of summer. Eroded and saline clays along arroyos
and ditches, rare. --swS-swAlta, US.

We have seen specimens from Nashlyn (DAO), Divide
(DAO), Pambrun (CAN, DAO) and Coaldale (CAN, DAO). It
has also been reported from Lethbridge.

37. HYMENOXIS Cass. RUBBER-WEED

Like the last two with a pappus of short hyaline
scales. Leaves alternate or basal. Heads radiate. Te-
gules more numerous, two layers thick, but isomegueth.

a. Leaves entire, all basal............+6. 1. H. acaulis
aa. Stem with deeply dissected leaves ......
SIRNA ee news eee es cee vccus sees ee He Richerdsont

1. BH, agaulis (Pursh) Parker var. acaulis (Tetra-
neuris simplex Nelson) -- Leaves densely soft sericeous
on both sides, entire, oblanceolate, all basal. Forming
a small dense cushion from a taproot. Scape about 1 dm
high, monocephalous. Head yellow, radiate, the ligules
fading white. Leaves finely glandular-punctate as in the
last two genera. Early summer. Upper part of eroded
hills; Mortlach and Cypress Hills westward. --swS-sAlta,
US,

In the magnilacustrine var. glabra (Gray) Parker,
the leaves are green, lightly villous or glabrescent,
the heads often somewhat larger.

2. He Bishardsonii, (Hooker) Cock. var. Richardso-

-- Leaves pectinately divided into 3-5 remote and
filiform segments. Herbage green. In small tufts, about
1 dm high. Also glandular-punctate. Ligules yellow,
fading white. Barly summer. Wind eroded hills and bad-

163 HYMENOXIS

80 BybreT Orb O.G Tok Vol. 23, noe 2

lands, --sS-sAlta, US.

In a more southern var. floribunda (Gray) Parker
the somewhat smaller heads are more numerous and the
whole plant tends to be larger.

Macoun 1884 mentions a collection of Chaenactis
Douglasii (Hooker) H. & A. by Dawson at Wood Mountain,
but we have found nothing at CAN or MTIMG under that na-
me or under the neighbouring genera. Unless this be 4
lapsus calami for Actinella Richardsonii (Hooker) Nutt.
(=Hymenoxys), the latter being represented at CAN by
an old Wood Mountain collection which Macoun does not
mention under the last two names.

38. HELENIUM L. SNEEZEWEED

A basic type, similar to Helianthus, the heads ra-
diate in yellow, but pappus present, of two or more se=
ries of hyaline scales. Receptacle not chaffy.

1. H, autumnale L. var. montanum (Nutt.) Fern. --
Sneezeweed -- Leaf blade decurrent down to the next node,
the stem thus narrowly winged. Leaves lanceolate, usual-
ly entire, finely glandular-punctate in yellowish to pale
brown. Heads yellow, more than hemispheric, with paler
and drooping ligules. The latter (0.8)-1.0-(1.5) cm long,
obtriangular, 3-lobed at apex. Mid to late summer. Wet-
tish meadows and edge of woods. --wO-BC, (US) -- Var.
grandiflorum (Nutt.) T. & G. (H. macranthum Rydb.) --
Heads larger, the ligules 1.5-2.5 cm long. --sMack, swAl-
ta-sEC.

The collections from Saskatchewan distributed by
Breitung as H. macranthum and reported by him as H. autum-
nale have been revised to var. montanum.

39. GAILLARDIA Foug.

Receptacle chaffy. Otherwise similar to Helenium.
Ligules also conspicuously 3-lobed at apex. Receptacle
convex to subglobose.

1. G. aristata Pursh -- Very showy bicolour head
with a purple center and orange-yellow ligules purple at
base. Short-lived perennial, hirsute, commonly monocepha-
lous. Leaves entire to pinnatifid. Peduncle elongate.
Head 4-8 cm across, the disk hemispheric. Tegules elon-
gate and very unequal. Early summer. Occasional in prai-
ries. -- sMack, swQ-BC, US-- F. monochroma Boivin -- Li-
gules and disk florests of a single colour, yellow throu-
ghout. Local: Waldheim, Milk River, Porcupine Hills. --
S-BC.

Specimens with smaller heads are found throughout,
but the range of size variation increases gradually west-
ward and the largest heads are found in the Rockies. This

HELENI UM 164

1972 Boivin, Flora of Prairie Provinces 81

was noted by Macoun long ago, but it seems difficult to
define this situation in taxonomic terms, although it is
not much unlike the situation in Helenium,

40. ANTHEMIS L. CHAMOMILE

This and the next 5 genera similar to Helianthus,
Helenium, etc., but the tegules scarious or hyaline along
the margin. Receptacle conical, chaffy. Pappus none or
vestigial.

a. Heads radiate in white .....eeeeeeeeeee 1. A. Cotula
ea. In yellow .....cccrcvcscccrveccvessse Se A. tinctoria

1. A. COTULA L. -- Mayweed, Dogfennel (Petite Mar-
guerite, Maroute) -- Peduncle pubescent. Tegules acutish
at tip. Receptacle chaffy in the central half. Otherwi-
se very similar to, and not readily distinguished from,
the more common Matricaria Chamomilla. Summer. Rare
railway weed: Morris, Killarney, Wetaskiwin, Troy. --
(Y-Aka, NF), NS-(PEI)-NB-Man=-(S)-Alta-BC, (US, Bur.).

We have checked specimens from Morris (DAO) and We-
taskiwin (SASK) while Dr. C. Frankton has also checked
specimens from Killarney (CAN) and Troy (CAN). Other-
wise all reports from our area are held as questionable
because of the frequent confusion with Matricaria. Mani-
toba and Saskatchewan reports by Groh 1948 were based on
specimens of Matricaria Chamomilla, and for Alberta on
M. maritima.

2. A. TINCTORIA L. (Cota tinctoria (L.) Gay) --
Yellow Chamomile (Oeil de boeuf, Camomille jaune) -- Heads
ligulate and resembling a Daisy, but bright yellow. Lea-
ves pinnatipartite, the segments dimegueth, the larger ones
pinnatifid, more or less alternating with much smaller and
entire segments. Summer. Infrequent escape, mainly along
roadsides. -- Aka, NF, NS, NB-BC, US, Eur.

41. ACHILLEFA L. YARROW

Like the last, but the receptacle flattish and the
heads quite small.

BeTnGNVeB SSGLTEtG .sccsaccecsse se cieeceee Tle A. Ee tAaALMICa
aa. Much more deeply dissected.
b. Leaves pinnatifid, the lobes dentate ...

MU ee Sa CRE STARR eee EE eee oe Hee Le hice
bb. ich more deeply and finely dissected ..
Screws ccccesecrcccsseveceeccce§ Se A. Millefolium

1. A. PTARMICA L. f. MULTIPLEX (Reynier) Heimerl
-- Sneezeweed, White Tansy (Herbe & éternuer) -- Double-
flowered heads small and white in a corymb. Stolonife-
rous. More or less virgate, 3-10 dm high. Leaves li-

165 ANTHEMIS

82 PHYTOLOGIA Vol, 23, no. 1

near-lanceolate. Summer. Cultivated and sometimes sprea-
ding to roadsides and waste areas. -- Aka, NS, Q-Man,
Alta-BC.

2. A. sibinica Led. (A. multiflora Hooker) -= With
small heads and obviously resembling the more common A.
Millefolium, but taller and the leaves less divided. Vir-
gate and tufted, 10-15 dm high. Leaves elongate, pinna-
tifid, the lobes oblong and serrate. Mid summer. Moist
spots in forested regions, not frequent. -- Mack-Aka,
Q=nBC, cnUS, (Eur).

3. A. MILLEFOLIUM L. f. PURPUREA (Gouan) Schinz
& Thellung -- Yarrow, Fern-Tansy (Herbe & dindes) -- Li-
gules velvet-purplish above, pink below. Stamens lac-
king, hence sterile and spreading only be rhizomes. O-
therwise similar to the common var. occidentalis, but
somewhat taller and less densely pubescent. Late summer.
Sometimes cultivated and rarely escaping to railway em=
bankments, etc. --Aka, NS-Man, US, (Bur) -- Var. occiden-
talis DC. (var. lanulosa (Nutt.) Piper; A. lanulosa Nutt.)
-- An almost ubiquitous herb with very finely dissected
leaves and a corymb of small, white heads. Long stoloni-
ferous. Leaves bipinnatipartite to tripinnatipartite
into numerous small segments less than 1 mm wide. Invo-
lucre usually 4-5 mm high. Tegules pale brown to hyali-=-
ne at margin. Ligules 1-4 mm long, white. Summer. Ve-=
ry common in open places, mainly steppes and prairies,
sometimes weedy. --Mack, Aka, L-SPM, NS-(PEIL)-NB-BC, US,
(CA) -- F. pegsea Rand & Redfield (f. roseoides Breitung)
-- Ligules pink above, nearly white below. Local -- (NF-
(SPM), NB-BC, (US) -- Var. megacephala (Raup) Boivin (A.
megacephala Raup) -- Heads larger, the involucre mostly
6-7 mm high. Mostly sand dunes. -- sMack, nwS-Alta --
Var. nigrescens E. Meyer (var. alpicola (Rydb.) Garrett,
var. borealis (Bongard) Farw.; A. borealis Bongard) --
Tegules with a darker margin, brown to blackish. The
more common or even exclusive phase northward. -- G,
K-Aka, L-SPM, NS-BC, US, Bur-- F. soseiflora Boivin --
Tegules darker as in var. nigrescens and the ligules pink
as inf. rosea. -- K-Aka, L-NF, Q-0,S—BC.

The first described is an uncommonly escaped orne=
mental. All specimens examined lacked anthers. That
they are of european origin seems hardly questionable.
The rest of the north American material is apparently
native.

If we except the highly local and larger-headed var.
megacephala, our specimens are fairly readily referable
to the two varieties above. We have not however been able
to detect a clear morphological gap between our american
types and the legion of minor eurasian variants. We have
been equally unable to relate our plants clearly to the

ACHILLEA 166

1972 Boivin, Flora of Prairie Provinces 83

many eurasian variants. Hence we could not be sure that
the varietal epiteths used are actually the earliest
available.

The european plants are hexaploid and seem closest
cytologically to var. nigrescens. But by their morpholo-
gy it is var. occidentalis and var. Millefolium that are
nearest to one another and hence often confused. This is
also part of the uncertain nomenclatural situation.

Of our two main types, var. occidentalis is common-
ly tetraploid while var. nigrescens is hexaploid(2n=54).
The level of morphological differenciation is low and its
quality is poor. However it is possible to state that,
grosso modo, the tetreploid var. occidentalis is the com-
com and wide-spread type in North America, while in the
mountains, on the Pacific slope and in subarctic habitats
it generally gives way to the hexaploid var. nigrescens,

Our two varieties are further recognizable with the
help of a good microscope as there is a slight average
difference in the outer diameter of the pollen grains,

a difference apparently related to the chromosome num-
bers. The following figures were obtained by Mulligan
and Bassett in 1950:
var. occidentalis: 2n=36; diam (26)-27-30-(31)n.
var. nigrescens: 2n=54; diam (31)-32-33-(34)yp.

Specimens of var. occidentalis from the interior of
the continent are usually readily distinguished from var.
Millefolium, but eastward the morphological distinction
becomes gradually less convincing. Further, some of the
eastern specimens with the apparent morphology of var.
occidentalis have the pollen size of var. borealis. The
opinion has been expressed that these could represent
& european introduction, but the evidence in favor of
the latter is rather negative and we are more inclined
to treat this material as an intergrading series bet-
ween our two main phenotypes. A similar situation pre-
vails in the western U.S.A. where one meets with an he-
xaploid, var. californica (Pollard) Jepson,which approa-
ches var. occidentalis in its morphology. In short, the
correlation morphology-cytology is incomplete.

The tetraploid seems absent from Europe; obviously
var. occidentalis should be regarded as native. Since
var. nigrescens is primarily a plant of native habitats,
it too is expected to be a native variant, even if also
found in northern Scandinavia.

Our 1951 classification was rather elaborate and
has not proved to be a good and practical scheme.

42, MATRICARIA L. WILD CHAMOMILE

Quite similar to Anthemis, yet the conical recepta-
cle not chaffy.

167 ACHILLEA

8h PoHP Yel OF LsOvG= iA Vol. 23, no. 1

a. Heads discoid...-..cccsccceseree Se Me matricarioides
aa. Ligulate.
b. Receptacle hemispheric, somewhat broader
than high,sometimes becoming conical in fruit;
herb odorleSS ..cecceceecccaccece le Mo maritima
bb. Receptacle conical and much higher than
wide; herb pineapple-scented ... 2. M. Chamomilla

1. M. MARITIMA L. var. MARITIMA (var. agrestis
(Knaf) Weiss; M. inodora L.; Chamomilla inodora (L.) Gi-
lib.) -- Bachelor's Button, Barnyard-Daisy -- Much like
the next species and not readily distinguished from it.
Odorless. Corolla lobes yellow with a brown spot towards
the tip. Achene with 3 very strong ribs and 2 large
brown glands near the top on the outer face. Annual or
biennial, 3-10 dm high. Tegules light-coloured along
the margin, hyaline to pale brown. Mid to late summer.
Casual weed, mostly of roadsides and railways. -- G, Mack,
Aka, L-SPM, NS-BC, US, Bur -- Var. nana (Hooker) Boivin
(M. ambigua (Led.) Krylov) -- Tegules dark-margined in
brown to blackish. Often perennial and usually shorter,
1-4 dm high. Sandy arctic coasts. -- G-Aka, L, nQ-n0-
nMan, (nEur).

Var. nana (Hooker) stat. n., Pyrethrum inodorum
(L.) Sm. var. ni nanum Hooker, Fl. Bor. Am. 1: 320. 1834.

2. M. CHAMOMILLA L. (Chamomilla Chamomilla (L.)
Rydb.) -- Wild Chamomile (Chamomille) -- Large-Headed
and suggesting a Daisy by its bicolour heads, but the
leaves bi-to tripectinatipartite into numerous segments
less than 1 mm wide. Ligules white, marcescent and even-=
tually drooping. Disk yellow, hemispheric, tending to
conicél in fruit. Achene without glands. Closely resem-
bling both M. maritima and Anthemis Cotula. From M. ma=
ritima it differs by being pinapple-scented when freshly
crushed; corolla lobes pure yellow; achene rugose with
5-7 nerves; pappus reduced to a short crown=-like ridge.
From Anthemis Cotula it differs by its herbage glabrous or
nearly so; tegules rounded at tip; receptacle not chaffy.
Barly summer. Infrequent weed of farmyards and roadsi-
des. =-- G, NF, NS, NB-(Q)-O-BC, US, Eur, (Oc).

3. M. MATRICARIOIDES (Less.) Porter (M. suaveolens
(Pursh) Buch.; Chamomilla suaveolens (Pursh) Rydb.) --
Pineapple-Weed, Wild Marigold (Herbe 4 crapaud) -- Dis-
coid and strongly pineapple-scented when freshly crushed.
Annual, up to 5 dm high. Leaves finely dissected like
the last two. Tegules broader, + oblong and more stron-=
gly cucullate at tip. Early summer to frost. Disturbed
or bare soils, a common weed, very tolerant of tramping.
--(G)-seF, Mack-Aka, L-SPM, NS-BC, US, (CA), Bur.

Reputedly native in ane western U.S. A. it has al-
ways seemed to us introduced wherever we met with ite an

MATRICARIA 168

1972 Boivin, Flora of Prairie Provinces 85

Canada. In 1884 Macoun knew it only from the Pacific
coast and from the upper Kootenay. This gives an idea
of its path of entry or, conversely, of its original
area as a native plant, assuming that it ever was na-
tive in Canada.

The range is extended to Franklin District on the
basis of the following collection: A. Dutilly, Terre de
Baffin, Cap Dorset, 25 aofit 1936 (QPA).

43. CHRYSANTHEMUM L. OXEYE-DAISY

Like Anthemis and Matricaria, but the receptable
flattish and not chaffy. Head typically ligulate.

@. Heads discoid ....-.ceseccecececesee Se C. Balsamita
aa. Ligulate.
b. Leaves long cuneate,grading into the
ISN OL Gm vine «cele eleie > ela sets aoetmuclens 2. C. arcticum
bb. Leaves dimorphic, the lower and basal
petiolate, the middle sessile and not
narrowed At DASE eecccccccce 1. C. Leucanthemum

1. C. LBEUCANTHEMUM L. var. LEUCANTHEMUM (var. pin-
natifidum Lec. & Lam.; Leucanthemum vulgére Lam.) -- Daisy,
Bull's Bye (Marguerite, Marguerite blanche) -- The typical
Daisy, a loosely tufted herb with virgate monocephalous
stems, the head with a yellow centre and long white ligu-
les. Stems 3-8 dm high. Leaves lyrate-pinnatifid, more
deeply so towards the base. Head 3-5 cm across. Mainly
the first half of summer and then sporadically till fall.
Formerly cultivated and now frequently spreading to wet-=
ter spots in pastures and along roadsides. --(NS-NB)-Q-
(O)-Man-Alta-(BC, US, Bur) -- Var. BOEBCHERI Boivin (C.
ircutianum Turcz.) -- Not so deeply dissected, the stem
leaves merely serrate or may be some of them subpinnati-
fid towards the base. Similar habitats, but tetraploid.
--K, Y, L-(NP) -SPM,NS-PEI -( NB) -Q-Alta-(BC), US, SA, Bur.

Var. Boecheri, nom. ne, C. ircutianum Turcz., Bull.
Soc. Nat. Moscou 39: 177. 1846. See also T.W. B&cher &
K,. Larsen, Cytotaxonomical Studies in the Chrysanthemum
Leucanthemum Complex, Watsonia 4: 11-16. 1957.

2. {. arcticum L. var. polaris (Hultén) Boivin --
(Chrysanthéme du Kamtchatka). Heads much as in the last,
but the foliage fleshy. Stem 1-2-(3) dm high, subscapose
or the foliage nearly all basal. Leaves long cuneate
into a winged petiole, the blade coarsely toothed to lo-
bed, and obovate to cuneate in shape. Tegules conspi-
cuously blackish=bordered. Early to mid summer. Arctic
coasts. --K-Mack-(Y-Aka), nQ-nMan, nBur.

Stat. n., ssp. polaris Hultén, Svensk Bot. Tidskr.
43: 776. 1949. The typical phase occurs west of us, in
the Queen Charlotte Islands westward to eastern Asia; it

169 CHRYSANTHEMUM

86 Pelpyol OFLFOyG TyA Vol. 23, no. 1

is commonly a taller plant with the leaves more deeply
cut, mostly trifid to pinnatifid.

3. C. BALSAMITA L. f£. TANACETOIDES (Boiss.) Boi-
vin =-- Costmary, Mint Geranium (Herbe au coq, Grand Bau-
me) -- Numerous discoid and yellow heads in 4 terminal
corymb. Leaves thickish, serrate, elliptic to lanceola-
te, the lower long petiolate and much larger, the upper
somewhat glaucous. Fall. Rarely spreading from cultiva-=
tion: Lloydminster. --swQ-O, S, US, (Eur).

Stat.n., Pyrethrum Balsamita (L.) W. var. tanace-
toides Boiss., Fl. Or. 3: 346. 1875. The heads are rayed
in the typical form, the latter apparently not known as
an escape in Canada.

44, TANACETUM L. TANSY

Ligules very short or lacking; otherwise hardly dif-
ferent from Chrysanthemun,
a. Leaf segments 3-10 mm wide ..-c...-+-. 1. T. vulgare
aa. Much more finely divided, the segments about
PSMA WLOS “sc cle vcs e ese sloe «cee cisisic 6 ces = (ie lis) siete

1. T. VULGARE L. -- Tansy, English Fern (Tanaisie,
Tenzé) -- Numerous yellowish=-green discoid heads ina
terminal corymb. Leaves pinnatipartite, the primary seg-
ments pinnatifid, the ultimate lobes entire to serrate.
Heads mostly less than 1 cm across, from slightly depres-
sed to somewhat convex at center. Mid summer. Often
cultivated and readily spreading to roadsides. -- Mack,
(Aka), sL-SPM, NS-BC, US, Eur -- Cv. CRISPUM -- More dee-
ply dissected, + bipinnatifid, the lobes overlapping,
crisp and upwardly curled at the tips. Less common. --
(NS)-PEI-O, (S)-Alta-(BC).

2. T. hurgnense Nutt. (var. bifarium Fern., var.
floccosum Raup, var. monocephalum Boivin, var. terrae-
novae Fern.; T. bipinnatum AA., ssp. huronense (Nutt.)
Breitung) -- Leaves feathery, very finely dissected,
tripectinatipartite, the ultimate segments about 1 mm
wide. Lightly to heavily tomentose stoloniferous peren=
nial, Heads few, mostly 2-5, and usually 1-2 cm across,
nearly discoid, the yellow ligules only 1-2 mm long.

Mid summer. Sandy shores, infrequent. -- K-Y-(Aka), NF,
NB-BC, US.

Highly variable and many phenotypes have received
names, We have been unable to bring them into a satis-=
factory classification, although the total range is con=
veniently broken up in a series of discrete areas. In
each area 4 particular type tends to dominate, such as
a single large head around Hudson Bay (var. monocephalum) ,
or more heavily lanate around Lake Athabaska (var. flocco-
sum), or the leaves somewhat fleshy along the Pacific

TANACETUM 170

1972 Boivin, Flora of Prairie Provinces 87

Coast (T. Douglasii DC.), etc., yet each local population
is highly variable, so variable indeed that its morpholo-
gical originality can be acurately expressed only in terms
of higher frequency of a particular phenotype in a4 parti-
cular area.

45. ARTEMISIA L. WORMWOOD

Heads small and resembling Achillea. However the
heads are discoid and the receptacle is not cheffy.

a. Leaves entire to coarsely lobed ......+.-..... GroupA
aa. Pinnatipartite to tripinnatipartite ......... Group B

Group A

Main stem leaves varying from entire to coarsely
and deeply lobed.

a. Shrubby; leaves grayish to whitish tomentose on both
faces.
b. Leaves oblanceolate to linear, entire ...
Cece e cers cesses cereressececcseesecesetde Ay CANA
bb. Cuneate, the apex truncate and three-
toothed .....ecccccccscccscccee 12, A. tridentata
aa. Herbaceous.
c. Main leaves entire, green and usually

GIADTOUS oe cecscesecsssccasess 2e A. Dracunculus
cc. White-arachnoid below, commonly serrate
to lobed.

d. Leaves heavily grayish or whitish-
arachnoid on both faces ...11. A. ludoviciana
dd. Less pubescent and much darker above.
e. Leaves entire and strongly revo-
Lute ce. ceceserceceeesee Ge A. longifolia
ee. Flat and at least the lower
ones coarsely lobed ...... 8. A. Tilesii

Group B

Leaves deeply and narrowly dissected; pinnatipartite
to tripinnatipartite.

a. Leaf segments narrow, all or mostly less than 1 mm
wide, usually entire.
b. Sterile; many ultimate segments with 1-(2)
teeth wccscescrccresecceccceccececs Ie Ae pontica
bb. Heads normally present in season; ultima-
te segments entire.
c. Whitish tomentose throughout, inclu-
ding the involucre ........... 15. A. frigida
cc. At least the involucre greenish.
d. Branchy shrub, woody below ...
Cece cccccccescccescccses§ 4. A. Abrotanum

a fy gal ARTEMISIA

88 Pre TOL iO1G Tuk Vol... 23, mite

dd. Herb, the stem not branched....
dala leteis\ sitio slelaicl'e sleleve ctele «, © Let AeMCAlperiaaers
aa. Segments broader and mostly toothed or lobed.
e. Leaves discolour, grayish to white-to-
mentose below, less densely so to gla-
brous above.
f. Leaves petiolate, without stipu-
LES ois. oe sin aisle + wvicleniceietele cle) LA. (ALA bea ei sm
ff. Leaves sessile, the lower pair of
segments often stipule-like.
g. Herb 2-4-(5) dm high; leaf seg-
ments 1-3 mm wide .....10. A. Michauxiana
gg. Much taller plant with broader
leaf segments ..e-.e..--4 7- Aw vulgaris
ee. Leaves green and similar on both faces.
h. Heads 5-10 mm across «.-ecee- 6. A. norvegica
hh. Much smaller and very numerous .eco
PO OC Ce eee eu

de A: campestris L. var. Wgrmsk joldii ( Besser)
Cronq. (var. Spithamea (Pursh) Peck; A. borealis Pallas)

-- (Aurone sauvage, Armoise rouge) -- Virgate peren-
nial with finely dissected leaves and a narrow panicle
of few heads. Tufted, with heavy rosettes, the foliage
primarily basal. Leaves pinnatipartite to bipinnatipar-
tite, green or grayish and more or less pubescent. Stems
1-3-(4) dm high. Stem leaves few. Inflorescence of uni=
form width, its lower branches long overtopped by the sub=
tending leaves. Corollas usually purplish in the upper
part. Mid summer. Alpine slopes and subarctic shores.
--(G)-F-Aka, L-NF, Q-nO, nwS-(Alta)-BC, nwUS, (Bur) --
Var. Scouleriana (Besser) Cronq. (var. caudata (Mx.)
Palm. & Stey.; A. canadensis Mx.; A. caudata Mx.) -- Tal-
ler, mostly 3-8 dm high, the stem more leafy, the inflo-
rescence & narrow panicle with the lower branches mostly
overtopping the leaves. More often biennial. Herbage
mainly cauline, green and more or less pubescent. Corol=
las yellow, the lobes often purple-margined. Mid summer.
Mostly sandy shores and open, sandy woods. --seK-Y-(Aka,
L)-NF, NS, NB-BC, US -- Var. Douglagiana (Besser) Boivin
(A. Bourgeauiana Rydb.; A. camporum Rydb.; A. caudata Mx.
var. calvens Lunell; A. Forwoodii Watson)-- Like the last
variety, but more pubescent, the herbage grayish-tomento-
se. Steppes and prairies, common. ==Mack-Y, Q-BC, (US).

Var. Douglasiana (Besser) stat. n., A. desertorum
Sprengel var. Douglasiana Besser ex Hooker, Fl. Bor. Am.
i: 325. 1833.

A rather variable type, both in the Old World and
in the New. Our three varieties are somewhat arbitrary.

ARTEMISTIA 172

1972 Boivin, Flora of Prairie Provinces 89

By increasing the level of arbitrariness one could reco-
gnize still more segregates as we did in 1955 (two spe-
cies, seven varieties). To-day we regard this earlier
classification as too arbitrary, too eleborate, and
hardly worth retaining.

2. A> REALS, © (A. dracunculoides Pursh; A.
glauca Pallas) -- Tarragon (Estragon, Herbe au dragon) --
With numerous small heads and numerous linear-ligulate,

entire leaves. Stem mostly 5-10 dm high. Leaves 1.5-
10.0 cm long, 1.5-4.0 mm wide, the lower ones often tri-

fid. Mid to late summer. Steppes and hillsides. -- (Y)-
Aka, wO-BC, US, (CA), Eur.
3. A. biennis W. -- (Herbe Saint-Jean) -- Biennial.

Glabrous and branchy. Upper leaves linear and entire, the
middle and lower pinnatipartite to bipinnatipartite, the
segments mostly 2-3 mm wide, sharply and irregularly ser-
rate to lobed. Inflorescence a panicle of numerous spi-
ciform groups of small heads. Late summer and fall.
Common on shores where apparently native; a frequent weed
of disturbed soils. -- Mack, NS-BC, US, (eEur, Oc).

4. A. ABROTANUM L. -- Southernwood, Sweet Benjamin
(Aurone, Citronelle) -- Perennial, woody below, and the
leaf segments mostly 0.2-0.3 mm wide. Herbage puberu-
lent, very densely so on growing parts. Otherwise much
resembling A. biennis. Second half of summer and early
fall. Cultivated and spreading to roadsides and waste
places. --Q-Alta, (US, Eur).

S. A. PONTICA L. -= Roman Wormwood (Petite Absin-
the, Plante de beauté) -- Normally sterile with us. A
Simple, virgate, gray-blue perennial herb growing in den-
se colonies. Herbage densely puberulent, the leaves whi-
tish below. Leaves bipinnatipartite, the segments 0.5-
1.0 mm wide. Panicles are rarely produced in late sum-
mer. Sometimes cultivated, long persistent and sprea-
ding vegetatively to waste places: Dauphin. -- NS, Q-
Man, US, Bur.

6. A-norvegica Fries var. saxatilis (Besser) Jep-
son -- Heads few, largest and commonly racemose. Tufted
perennial 2-5 dm high. Leaves mostly basal, bipinnati-
partite, the ultimate segments entire or nearly so. Te-
gules broadly margined in purple black. Heads 5-10 mm
wide, drooping on erect peduncles. Mid summer. Alpine
Slopes. --(wF), Mack-Aka, (neO), swAlta-BC, wUS, (eBur).

The typical phase of western Eurasia is a usually
smaller plant, its heads tend to be larger, and the cen-=-
tral rachis of the leaf is shorter so the limb seems al-
most palmately cut.

7. 4A. VULGARIS L. -= Mugwort (Herbe Saint-Jean,
Herbe & cent gofits) -- Leaf seemingly stipulate, the lo-
wer 1-2 pairs of lobes or leaflets being borne at the base

173 ARTEMISIA

90 PreHyee? OjL OG TA Vol. 23, nosis

of the petiole-like rachis. Branchy perennial. Leaves
dark green and glabrous or nearly so above, white-tomen-
tose below, pinnatifid or pinnatipartite to compound to-
wards the base. Tegules with a deep green midnerve and
white=-tomentose limb. Mid summer to frost. Rare weed of
waste places. --(G), NF, NS-S, BC, US, (CA), Bur.

8. A. Tilesii Led. (var. unalaschkensis Besser; A.
Herriottii Rydb.) -- Very variable type, but with the
stem leaves,;or at least the lower ones,long-lanceolate
with a few lanceolate lobes. About 1m high. Leaves
0.5-1.5 dm long, 5-20 mm wide, white-arachnoid below,
grayish tomentose above when young, becoming glabrous.
Heads few to many, small to large. Sometimes resembling
the last species, but lacking the stipule-like lobes.
Sometimes close to the next, but the leaves thinner and
often larger and at least the lower ones lobed. Mid sum-
mer. Open woods and river flats. -=-(wF)-K-Aka, wQ=BC,
(US), nEur.

A rather polymorphic type, perhaps divisible in two
or three geographical variants. We have not yet been able
to establish or recognize a sound morphological basis for
the distinction of such variants.

9. £- Jongifalia Nutt. -- Linear leaves strongly
revolute and white-arachnoid below. Densely tufted from
a@ woody base and taproot. Leaves (2)-3-5-(8) mm wide,
thickish, lightly arachnoid above, entire. Stem simple,
3-8 dm high. Involucre arachnoid. Mid summer. Wind ero-
ded steppes and badlands or lightly alkaline soils. --
sMan=-sbC, US.

Has been recently detected west of us at Osoyoos
(DAO), Kelowna (DAO) and Summerland (UBC). The Ontario
report by Fernald 1950, repeated by Scoggan 1957, quer-
ried by Boivin 1967, is to be discounted as it could not
be substantiated at GH or elsewhere.

10. A. Michauxiang Besser -- Segments of the lower
pair stipule-like as in A. vulgaris, but the stem shorter
and simple and the leaf segments narrower. With a ta-
proot and somewhat stoloniferous, forming loose colonies.
Leaves green above, white-arachnoid below, pectinatipar-
tite to bipectinatipartite, the segments 1-3 mm wide.
Inflorescence very narrow, sometimes subspiciform. Early
summer. Gravels and rocky exposures at mid altitude. ==
swAlta-EC, ae

als, ludovicigna Nutt. var. ludoviciana (var.

gnaphalodes / or ¥ var. latifolia (Besser) T. & G.; A. di-
versifolia Rydb.; A. gnaphalodes AA.; A. Purshiana Besser)
-- Sage, White Sage -- Long stoloniferous. Leaves mostly
1 cm wide, lanceolate, grayish to white-tomentose on both
faces, entire to paucidentate towards the apex. Heads
arachnoid with a purplish or brownish disk. Late summer.

ARTEMISIA 174

1972 Boivin, Flora of Prairie Provinces 91

Prairies and open places, common. --sMack, PBI-BC, US--
Var. gnaphalodes (Nutt.) T. & G. (A. pabularis (Nelson)
Rydb.) -- Smaller and often yellowish=pubescent. Leaves
t linear, 3-5 mm wide, most often conduplicate. Steppes.
--swQ-Alta, US.

With us this is essentially a native plant, but its
occurence in Eastern Canada is mainly in the form of a4
not particularly agressive weed invading open places.

The type of Artemisia gnaphalodes Nutt. (PH) is the
narrow-leaved phase commonly treated as A. pabularis (Nel-
son) Rydb., while the type of A. ludoviciana (also PH)
is merely a sterile shoot of the broader leaved phase col-
lected late in season and somewhat glabrescent above as
often happens in what is usually called A. gnaphalodes.
Hence the shift in names and the usage adopted here, in
which var. gnaphalodes becomes the correct name for the
narrow=leaved phase.

12. A. tridentata Nutt. var. tridentata -- Sage-
brush (Absinthe) -- Leaves narrowly cuneate, three-too-
thed at apex. Shrubby. Herbage grayish-tomentose throug-
hout. Heads numerous and small. Late summer and fall.
Steppes along the South Castle Creek in the Crowsnest
Area. swAlta-sBC, US, (CA).

Involucre 3-4 mm high. At Hedley B.C. (DAO) and
south there occurs a var. Vaseyana (Rydb.) stat. n., A.
Vaseyana Rydb., N. Am. Fl. 34: 283, 1916, with somewhat
larger heads, the involucrum + 5 mm high.

13. A. gana Pursh -- Wild Sage -- Shrubby with nar-
row and entire leaves equally whitish-tomentose on both
faces. Panicule leafy, the leaves mostly overtopping the
flowering branches. Late summer and early fall. Dry hills
and steppes. --swMan-sAlta, US.

14. A. ABSINTHIUM L. var. INSIPIDA Stechmann --
Wormwood, Absinth (Absinthe) -- Leaves grayish-
tomentose and pinnatifid to nearly tripinnatifid into li-
gulate and subentire segments about 2-4 mm wide. Panicle
ample with numerous drooping hemispheric heads about 4 mm
wide. Mid summer to fall. Cultivated and casually sprea-
ding to roadsides, etc. -- (NF-SPM), NS-(PEIL)-NB-BC, US,
Bur.

15. A; frigida W. -- French Sage, Prairie-Sage-
wort -- Foliage whitish and finely divided, the leaves
rather short. Tufted perennial, white-silky throughout.
Leaves less than 2 cm long, pinnatipartite to bipinnati-
partite, the segments entire and less than 1 mm wide.
Second half of summer. Vary common in steppes and prai-
ries, --Mack-Aka, NB-BC, US, Bur.

Essentially a prairie species, it is known in EKas-
tern Canada as a sproradic introduction, but is perhaps
also native at a few spots on Lake Superior.

46. PETASITES Miller SWEET COLTSFOOT

Resembles Senecio. Flowering stems shedding their

175 ARTEMISIA

92 P, HvXeT O/L10uG Tod Vol. 23, no. 1

seeds and evanescent by the time the basal leaves are
fully grown. Plants subdioecious. Stem leaves very much
reduced.

Species of this genus present some unusual and inhe-
rent difficulties of identification, partly because of the
alternance of biological phases. Most specimens will re-
present only one phase and the correlation of characters
is difficult to establish. Then herbarium specimens show=
ing both phases are a small minority and in most cases the
two phases are not root-connected, leaving open the possi-
bility that they may have come from different clones,
perhaps different species.

Many specimens have turned up identified or revised
to various hybrid combinations. We have studied a fair
number of such specimens at DAO, UBC and V, and we are
not fully satisfied that their morphology could justify
the postulate of hybridity. Most such specimens have
been revised or returned to P. vitifolius, others to P.
sagittatus or P. palmatus. However we have not yet seen
the many Yukon and Alaska intermediates discussed by
Hultén 1950.

a. Leaves suborbicular, palmatifid .......3. P. palmatus
aa. Deltoid to sagittate, undulate to lobed.
b. Deltoid and deeply lobed ..... 2. P. vitifolius
bb. Sagittate, the margin undulate to coar-
SelyGenitertc aisle elelsiclelstetelslclelsiatc 1. P. sagittatus

1. PB. sagittatus (Banks) Gray -- Like the next two,
but the leaves triangular-sagittate, 1-2 dm long, deeply
cordate at base, the margin sinuate to dentate. Stems
arachnoid=pubescent, not glandular. Pappus 15-22 mm long.
First half of spring. Wet places. -- (seF)-K-Aka, L, Q-
Be, US.

The more northern P. frigidus (L.). Fries was map=
ped by Porsild 1957, 1964, showing an unlikely dot near
Jasper, but was not listed by Porsild 1959. The specimen
basis of the dot could not be determined positively.

2. RB. virifolius Greene (P. frigidus (L.) Fries
var. nivalis (Greene) Cronq.) -- Like the next. Leaves
deltoid, deeply dordate, irregularly lobed and dentate,
about as wide as long, sometimes up to 2 dm across, but
mostly under 1 dm wide. Stem both arachnoid and glandu-
lar. Pappus 12-15 mm long. Spring. Wet or boggy pla-
ces. =-=(sK)-Mack-Aka, (L), Q-BC, US.

3. PB. palmatus (Aiton) Gray var. palmatus (P. fri-
gidus (L.) Fries var. palmatus (Aiton) Crong.) -- Herb at
first producing a simple flowering stem with leaves redu-
ced to large dilated stipules and a dense raceme elonga-=-
ting in fruit. Leaves appearing later in early summer
only. Stoloniferous. Stem 2-6 dm high, glandular-pubes-

PETASITES 176

1972

cent, rarely slightly arachnoid.

Boivin, Flora of Prairie Provinces 93

Leaves up to 2 dm across

suborbicCular, palmatifid. Heads short ligulate; involu-
cre up to 1 cm high, ligules yellowish, pappus 8-12 mm

long.

Spring. Frequent in low places. --(sK) -Mack-Y,

L-NF, NS-neBC, neUS.

A report of the related genus Erechtites hieracii-
folia (L.) Raf. by Macoun 1884 was based on a mention by
Hooker 1834 of Senecio hieraciifolius L. from Saskatche-
wan. This has never been confirmed and was not accepted
by later authors.

47. ARNICA L. ARNICA

Leaves opposite, otherwise as in Senecio. Tegules
isomegueth.

a. Leaves cordate to narrowly oblong.

b.

bb.

Larger stem leaves broadly cordate and long
petiolate, the petiole mostly about as long
as the blade; achene abundantly short hir-
EAUGO: dercucesvausratetaiticta ie covccceseccese 8. A. cordifolia
Stem leaves oblong to ovate-oblong, subcor-
date to cuneate at base; petiole less than
half as long as the blade.
c. Pappus pale brown; leaves closely and
sharply dentate ......... ll. A. diversifolia
cc. White; leaves somewhat remotely serra-
te or denticulate; achenes usually gla-
DYOUSS. J sicise sews Geaeess disease fe A. latifolia

aa. Narrower, oblong-lanceolate to linear.

d.

dd.

Stem leaves in (1)-2-3-(rarely 4) pairs ..
Bistetatia® ctaldleieleis us sicleWiaclelcle sls we clsisiseas ceeese GLOUD A
Stem leaves in (4)-5-6-(8) pairs.
e. Leaves closely and sharply dentate ..
Cece eres cccccscccsccccsccell. A. diversifolia
ee. Entire to remotely denticulate.
f. Tegules lanate-ciliate at tip..
Crees eesecsccccccerese Gs A. Chamissonis
f£. Tegules much more sharply acute
and the ciliation not unusually
dense at tip.
g. Leaves entire or nearly so;
tegule pubescence entirely
or essentially of short glan-
dular hairs........ 10. A. longifolia
gg. Leaves remotely denticulate;
tegule pubescence primarily
WIAACUN vasecstpascect ss. bis He MOLILe

Group A

Stem leaves rather narrow and few; mostly in (1)-2-
3 pairs and broadly lanceolate to linear. Pappus mostly

Lite ARNICA

9h POH tT. Orb ONG, TA Vol. 23, now 1

white.

a. Heads discoid; pappus pale brown ...... 13. A. Parryi
aa. Heads radiate.
b. Heads at first nodding; achene glabrous
or finely glandular below the middle,
sparsely pilose above ...ee.-+e-. 2. A. louiseana
bb. Brect; achene uniformly pilose; mostly
taller plants.
c. Leaves remotely and regularly den-=-
ticulate with distant and subopposi-
te teeth.
d. Pappus white ..«ee..e- 3. A. lonchophylla
dd. Pale brown; tegules longer ....
sjeveperateie ate a ayniow de ciee so 0a wale, « Loe Cee ee
cc, Leaves entire or irregularly and remotely
denticulate.
e. Tegules 9-11 mm long; leaves
broadly lanceolate ..... 4. A. Rydbergii
ee. Tegules 11-15 mm long.
f. Leaves 1-4 cm wide and near-
ly uniform in length but the
middle ones somewhat longer ..
ole ale ae melee el ele SRicl cee kes Aq melee
ff. Much reduced upward, the lo-
west at least twice longer
than the upper, also mostly
narrower and lanceolate to
linear.
ge Rhizome with tufts of long,
brown hairs; leaves + oblan-
ceolate, the lower at least
li em wide sscis.ssso0. Ag Eulgens
gg. Rhizome without tufts; leaves
lanceolate to linear and not
over 1 cm wide.

h. Ligules light yellow; te-
gules somewhat acuminate,
often purplish and squar-
rose at tip .... 1. A. alpina

hh. Ligules orange-yellow; te-
gules broadly acute at tip
and green ..... 6. A. sororia

1. A. alpina (L.) Olin var. ungavensis Boivin (ssp.
attenuata (Greene) Maguire; A. attenuata Greene) --A
middling type, 1-3 dm high with 2-3 pairs of stem leaves
and usually only 1 head. Leaves usually less than 1 cm
wide, linear-lanceolate, acuminate, entire. Involucre
obviously glandular-puberulent and lightly villous, more
densely so towards the base. Pappus clean white. Mid

ARNICA 178

1972 Boivin, Flora of Prairie Provinces 95

summer. Tundra and rocky alpine slopes and summits. --
K-Aka, L-(NF), nQ-nMan-(nS)-swAlta-nBC -- Var. vestita
Hultén (var. tomentosa (J.M. Macoun) Cronqg.) -- Densely
soft lanate, especially on the tegules and the base of
the involucre. Tegules also abundantly, but not obvious-
ly, short glandular-pubescent under the heavy lanosity.
Ligules rather short, usually of about 1 cm. Less com-
mon. --(Mack-Y)Aka, (wNF), swAlta-(BC, nwUS).

Var. ungavensig (Boivin) stat. n., A. Sornborgeri
Fern. var. ungavensis Boivin, Nat. Can. 75: 211. 1948.

2. A. loviseana Farr var. louiseana -- Smaller
than the last, usually around 1 dm high. Leaves broa-
der, mostly 1-2 cm wide, + lanceolate and not acuminate,
mostly basal, the stem usually bearing only 1 reduced
pair. Head nodding at anthesis and usually solitary.

Mid summer. Shale slides at high altitudes in the Banff
area. --Y, swAlta-neBC.

Vicariant of the eastern var. Griscomii (Fern.)
stat. n., A. Griscomii Fern., Rhodora 26: 105. 1924, from
Gaspé and Newfoundland. The latter has erect heads at
anthesis. Also it is a somewhat taller plant and its ache-
nes are slightly shorter.

3. A. lonchophylla Greene var. lonchophylla (A.
arnoglossa AA.) -- Resembles A. alpina, but somewhat lar-
ger and the leaves are remotely denticulate with a few
pairs of subopposite teeth. Mostly (2)-3-4-(6) dm high.
Upper leaves much reduced and usually entire, the basal
ones with 4 petiole at least half as long as the blade.
Involucre often shorter. Early summer. Near shores in
limestone regions. --(swK)-Mack-Y, NF, NS, NB-Alta,
ncUS.

In ours the herbage is both glandular and long pi-
lose. In the more southern var. arnoglossa (Greene) Boi-
vin the glandulosity is more abundant and obviously domi-
nant while the pilosity is much shorter and scanty, the
contrast is especially strong on the tegules where the
longer hairs are quite lacking or nearly so.

4. A. Rydbergii Greene -- Somewhat halfway between
A. alpina and is latifolia. Loosely tufted and 1.5-2.5 dm
high. Stem leaves mostly 2 pairs of which the upper are
broadly lanceolate, the lower broadly oblanceolate, enti-
re to irregularly denticulate. Head usually solitary, the
involucre rather short. Pappus clean white. Summer. Low
alpine, mostly on shale slides, -- swAlta-(eBC), nwUS.

5S. A. fulgens Pursh -=- Rhizome with many tufts of
long, brown hairs. Pubescence dense, primarily glandu-
lar and lightly tinted. Stem leaves 2-3 pairs, the upper
much reduced, the lower 2-4 times longer, oblanceolate,
entire to irregularly denticulate. Head usually solitary
and on a peduncle usually longer than any of the interno-

179 ARNICA

96 PART O LAG IA Vol. 23, no. 1

des. Outer tegules up to 2,5-3.5 mm wide. Early sun-
mer. Shallow depressions in the steppe. Infrequent but
showy. --SswMan-eBC, US.

6. A. roria Greene --Closely resembling the last,
but lacking the tufts of brown hairsat the base of the
stem and on the rhizome. Lower leaves not so clearly
oblanceolate and tending to narrower, usually less than
1 cm wide. Peduncles less elongate, often shorter than
any of the internodes. Tegules somewhat narrower, (1.0)-
2.0-(2.5) mm wide. Early to mid summer. Foothill prai-
ries and open montane woods. --sAlta-sBC, (wUS).

7. A. latifolia Bongard (A. gracilis Rydb.) --A
rather large-leaved species with typically two pairs of
stem leaves of which the lower are serrate, + oblong and
rounded at base to a winged petiole. Basal leaves some-
times cordate. Stem 2-6 dm high. Involucre rather nar-
row and high, the tegules 12-18 mm long. Pappus pure
white. Achaine commonly glabrous, sometimes sparsely pu-
berulent in the upper half,more rarely minutely glandular.
Early to mid summer. Wetter montane forests. --swMack-
Aka, swAlta-BC, wUS.

Plants from higher altitudes tend to be generally
smaller, including smaller heads. A number of names have
been proposed for this ecological extreme, the latest
being var. gracilis (Rydb.) Cronq. An essentially pa-
rallel variation occurs under the next species: var. pu-
mila (Rydb.) Maguire is available to single out such
smaller variants of higher altitudes.

8. A. cordifolia Hooker -- A rather showy forest
species resembling the last, yet the basal and lower lea-
ves deeply and broadly cordate. Head mostly 5-6 cm
across. The petioles elongate and not winged. Tegules
12-20 mm long, 3-5 mm wide, oblanceolate. Pappus white.
Late spring to mid summer. lLodgepole forests; common
at low altitudes in the Rockies, highly disjunct east-
ward. --swMack-Y, Man-BC, US.

Recently discovered in the Riding Mountain area
(Rassburn Tower Cabin), this cordillerian species also
occurs east of the Rochies in a rather unusual and highly
disjunct manner: Wintering Hills, Cypress Hills, Pasquia
Mountain and Riding Mountain, Also south of the border
in the Sweetgrass Buttes of Montana, the Black Hills of
South Dakota and the Keweenaw peninsula of Michigan. Ma-
ny of these isolated localities are in recently glacia-
ted territory, which would indicate a species enjoying a
range expansion in earlier post-glacial times followed
with a regression to the present highly sporadic condi-
tion.

Reports of A. cordifolia for southeastern Alaska
by Maguire 1943, Hultén 1950, Anderson 1952, Gleason 1952,
Cronquist 1955, querried by Boivin 1967, are apparently

ARNICA 180

1972 Boivin, Flora of Prairie Provinces 97

in need of confirmation. The original report was based
on @ collection by Cushing from Muir Glacier and two Krau-
se collections from Tlehini and Klokwan. The Muir Gla-
cier collection (CU) has the short petioles, the triangu-
lar-oblong leaves with broadly cordate base and the small
size typical of A. latifolia var. gracilis, and has been
revised accordingly. It is not possible to state if the
Krause collections should be similarly revised as these
were preserved at the Berlin Botanical Garden and were
presumably lost in the fire that destroyed their herba-
rium.

The Keweenaw plant has been described as a separa-
te species, A. Withneyi Fern., which differs in no subs-
tantial way from the cordilleran plant, yet the limited
Keweenaw populations exhibit, as would be expected, a nar-
rower range of variation than the multitude from the
Rockies and westward. Such a restricted type has no taxo-
nomic value by the mere fact of its restricted range and
variation. The other 6 isolated localities also support
populations of similarly restricted range and variation
and would also rate taxonomic rank if either limited va-
riability or localized occurrence were taxonomic charac-=
ters per se, a situation where the place of collecting
would actually become the primary taxonomic criterion.

9. A. Chamissonig Lessing (var. angustifolia Her-
der, var. incana (Gray) Hultén, ssp. foliosa (Nutt.) Ma-
guire; A. foliosa Nutt.)-- Stem leaves more numerous,
mostly in 4-5 pairs. Long stoloniferous and (3)-4-6-(8)
dm high. Leaves + lanceolate, 1-5 cm wide, remotely
denticulate or entire, commonly about as long as the in-
ternodes. Lower part of stem often purple. Herbage
abundantly long villous and glandular-puberulent. Heads
mostly 3-5, corymbose. Tegules broadly acute and lanate-
ciliate at tip. Pappus pale brown to nearly white.

Mid summer. Low lying patches in black soil regions.
--sMack-sAka, wcQ-BC, wUS.

Some average differences are fairly obvious when
specimens from opposite ends of the range are contrasted.
Thus Alaska specimens (var. Chamissonis) tend to larger
stem leaves, (1.5)-2.0-3.5-(5.0) cm wide, more obviously
toothed, the pappus usually tawny, varying to nearly
white, and Ontario specimens (var. angustifolia Herder)
have narrower leaves 1.0-2.0-(3.5) cm wide, entire to
weakly toothed, the pappus mostly nearly white. The Ca-
lifornia specimens (var. incana (Gray) Hultén) are often
quite heavily tomentose. However these morphological
types are merely statistical variants, they occur with
greater frequency in one area without being completely
absent from the rest of the range. Most specimens have
leaves about 2 cm wide, pappus light tawny, and average

181 ARNICA

98 PABVY LT OFL40%G Deh: Vol. 23, 205%

tomentum; it is difficult to sort them out into geographi-
cal variants without undue emphasis on the place of col-
lecting. Certainly, a realistic sorting of the material
at hand would not achieve the strong geographical res-
trictions illustrated by a dot map in Brittonia 4: 462,

1943,
The extreme with dense and felty tomentum could be

regarded as an ecological form (= f. incana (Gray) Boivin
of wetter years, often found standing in shallow water.
The evidence at hand is still too scanty to be conclu-=
sive.

10. A. longi ia D.C. Eaton -- Involucre not vil-
lous at base, or only slightly so. With 5-8 pairs of
stem leaves, lanceolate to narrowly lanceolate, thus re-
sembling the last, but greener, the dense and short glan-
dulosity not being mixed with any villosity. Leaves
entire to remotely denticulate, at least as long as the
internodes and commonly twice longer, the upper often
overtopping the heads, the basal ones absent at flowe-=-
ring. Late summer. Forming large patches along subal-
pine creeks in Waterton. --swAlta, wUS.

ll. A. diversifolia Greene -- Leaves closely and
very sharply dentate, the sinuses rounded, Stem leaves
mostly in 5 pairs, the larger ones £ ovate, the others
gradually shorter and much narrower, mostly petiolate,
the petioles winged. Herbage lightly villous and dense-
ly glandular-puberulent throughout. Pappus pale brown.
Mid summer. On wet cliffs and along subalpine creeks.
--(Y)-seAka, swAlta-BC, wuUS.

12, A. mollis Hooker var. mollis (A. lanceolata
Nutt.) -- A middling and non-descript type with 3-4 pairs
of stem leaves, nearly entire to serrate, + lanceolate,
1-4 cm wide. Pubescence mixed, partly glandular-puberu-
lent, partly villous or glandular-villous. Pappus pale
brown. Second half of summer. Wet or boggy places in
the mountains. -- (swMack-seY), nNB-seQ, swAlta-sBC,
wUS -- Var. aspera (Greene) Boivin (A. amplexicaulis
Nutt.) -- More leafy, mostly with 5-6 pairs of stem lea-
ves. -- (wMack, seAka), swAlta-BC, wUS.

Var. aspe (Greene) stat. n., A. aspera Greene,
Ott. Nat. 15: 281. 1902,

Another variant from the U.S. Northwest is var.
Piperi (St. John & Warren) stat. n., A. amplexicaulis
Nutt. var. Piperi St. John & Warren, Proc. Biol. Soc.
Wash. 44: 36. 1931, distinguished mainly by its more
ample foliage, the leaves up to 4-6-(8) cm wide.

One of the more remarkable cases of range dis-
junction in North American, widely distributed in the
Rockies and again around the Gulf of St. Lawrence. A
Great Lakes report by Macoun 1903 was based on a sheet

ARNICA 182

1972 Boivin, Flora of Prairie Provinces 99

labelled, R. Bell, Gros Cap, July 25, 1860 (QK); it is
to be discounted as the specimen belongs to Coreopsis
lanceolata var. lanceolata.

13. A. Pagryi Gray var. Parryi -- Discoid. Simi-
lar in habit to A. fulgens. Lower and basal leaves lan-
ceolate, broadest near the base, long petiolate. Middle
and upper stem leaves strongly contrasting, less than
half as long and sessile. Heads commonly 3-5 and fairly
large, but rayless. Mid summer. Mountain meadows towards
timberline. --(Y), swAlta-sBC, wUS.

In the more southern var. Sonnei (Greene) Cronq.
the heads are radiate.

48. SENECIO L. GROUNDSEL

Mostly a conspicuous herb with yellow heads ina
terminal corymb, often in an umbelliform corymb. A ba-
sic type, resembling Solidago with its yellow flowers and
ligules and its pappwof bristles, but the tegules iso-
megueth and more or less in a single row, or sometimes
dimegueth, the outer ones few in number and many times
shorter than the inner.

@. Monocephalous or annual ...-.ccae-eeeeeeeees Group A
aa. Perennial and polycephalous.
b. Leaves subentire to dentate ............ Group B
bb. Some or all leaves coarsely lobed to pin-
Ptr GLC dads isi dieleles o side daiam aisles see CLOUD &

Group A

Annual herbs, usually polycephalous. Or perennial
but monocephalous or sometimes with a second smaller
head.

a. Annual.
b. Annual from a bulbous base with fasciculate
BOOLLOCE & ners, ciwielatsalele satate sereeee 4. S. congestus
bb. Annual with a taproot.
c. Herbage nearly glabrous; outer tegules

black-tipped .....se+ee0--+-- 1. S. vulgaris
cc. Herbage heavily glandular; all tegu-
les green to the tip ......... 2 S. viscosus

aa. Perennial.
d. Leaves subentire; head largesSt..ccccoee

eee cece cece ccescccssesesecess De Se megacephalus
dd. Lower stem leaf deeply lobed .........

Ce ecoesessessesecesescsasses 17. S. resedifolius
Group B

Leaves subentire to dentate or serrate.
a. Head largest, solitary or nearly SO .oecee

Re CIS 6k ow) vei elec Sw Sw 8 Semi S. S. megacephalus
183 SENECIO

100

Pe hol ore TA Vol. 23, no. 1

aa. Polycephalous, with more than 2 heads.

b. Leaves + isomegueth, + truncate at base ..

os ne.e'e a'e6 G0 8 00/0:50018 Waele @ w(Sleiehe ete? phe o\ fekzks Aen RRa eae
bb. Lower leaves long cuneate at base.
c. Upper stem leaves sessile but other-
wise not much smaller than the lower
ones.
d. Stem leaves much shorter than
the internodes and much narrower
than the rosette leA€veS ..cecee
acogec Jseleracds (Lb S. streptanthitoliug
dd. Stem leaves rather longer than
the internodes.
e. No rosette leaves; main stem
leaves + obovate..... 6. S. Fremontii
ee. Rosette present; stem leaves
lanceolate or oblanceolate ..
old.ciccilaaicca nc . 260 (Strident eee
cc. Middle and upper stem leaves less than
half as long as the lower ones.
f. Heads in a corymbiform raceme, ra-
rely somewhat compound; herbage +
villous or tomentose, at least in
the inflorescence..... 9. S. integerrimus
ff, Outer branches bearing short ra-
cemes of 2-5 heads; herbage gla-
brous or lightly pubescent in
the inflorescence ......-. 8. S. foetidus

Group C

Polycephalous perennials with leaves more deeply

dissected. At least the stem leaves coarsely lobed to-
wards the base, more commonly pinnatifid to pinnatipar-
tite.

ae

a4.

Leaves all alike.
b. Herb 3 dm high or less; leaves slightly
fleshy, dentate to lobed ..16. S. tridenticulatus
bb. Much taller herb; leaves thin and more
deeply dissected....a+c2e22eccee Se Se Cremophilus
Leaves polymorphic: rosette leaves larger and
usually merely dentate; stem leaves more dee-
ply dissected, the upper ones smaller.
Basal leaves cuneate at base, usually less
than 2 cm wide.
d. Herbage grayish to whitish-tomentose, but
the leaves sometimes green above.... 10. S. canus
dd. Herbage less pubescent and green throughout.
e. Rosette leaves uniformly crenate or
serrate to base .......... 12. S. pauperculus

SENECIO 184

1972 Boivin, Flora of Prairie Provinces 101

ee. Mostly with only 3-5 teeth towards
the apex ......+-.+-+eell. S. streptanthifolius
cc. At least the larger leaves subtruncate to
cordate or broadly rounded at base.
f. Larger leaves cordate or ovate to reni-
form; heads radiate; tegules green ...
Co eee c eee cece ccccessesseseceseee 13. S. aureus
ff. Larger leaves more or less truncate at ba-
se; heads radiate or eradiate; tegules
+ purplish.
g. Herb 2-4 dm high; basal leaves 1-3
cm long o-.-.sescceseecece 15. S. pauciflorus
gg. Taller and the leaves larger ....c.
Ce eee cerecccccseseccesesoe 14. S. indecorus

1. S. VULGARIS L. -- Groundsel (Grand mouron, Toute
venue) -- Main tegules green, but the outer short tegu-
les black in the upper third. Leaves + oblanceolate, ir-
regularly lobed to pinnatifid, the lobes irregularly den-
tate. Discoid. All summer. Casual weed, rarely abundant,
-- (G), Mack-(Y)-Aka, L-SPM, NS-BC, US, Bur.

2. S. VISCOSUS L. --Stinking Groundsel -- Closely
resembling the first, but densely glandular puberulent.
Inner tegules with a small brown spot at tip; outer ones
green. Ligules very short, the heads almost discoid. Se-
cond half of summer and fall. Rare weed of disturbed
soils: Winnipeg. --(NF), NS-Man, BC, (US), Bur.

3. S. eremophilug Rich. var. eremophilus -- No ro-
settes, but the stem leaves numerous and pinnatifid to
pinnatipartite. Fairly showy tufted perennial, (0.6)-1.0-
(1.5) m high. Leaf lobes narrower than the sinuses.
Heads fairly large and long ligulate. Tegules finely
tipped in black. Mid summer. Wetter spots at edge of
forests. -- sMack, O-BC, US.

In the southern Rockies our plant gives way to
var. Kingii (Rydb.) Greemm.with smaller heads, the invo-
lucre only 5-7 mm high, the tegules more conspicuously
black-tipped.

4. S. cgongestys (Br.) DC. (var. palustris (L.)
Fern., var. tonsus Fern.; S. palustris (L.) Hooker) --
Marsh-Fleabane -- Annual from a bulbous base. Long-lana-
te throughout. Stem thick, hollow, up to 1m high, but
usually much smaller. Bulb also hollow. Stem leaves nu-
merous, undulate to pinnatifid. Heads in clusters. Li-
gules short. Late spring to mid summer. Exundated pla-
ces. -- F-Aka, L, Q-Alta-(BC), ncUS, Bur.

5. S. megacephalus Nutt. -- Heads largest, 2.0 -
2.5 cm high, and usually solitary or sometimes with 4
Second smaller one borne on a longer peduncle. Herbage
tomentose-floccose, especially along the leaf margins.

185 SENECIO

102 Pil YT) Or Te0eG: Dyk Vol. 23, 0maane

Leaves entire to dentate, oblanceolate, the upper much
reduced. Mid summer. Alpine ridges in Waterton. -- swAl-
ta-seBC, nwUS.

6. S. Fremongii T. & G. var. Fremontii -- A tufted

“_
perennial, somewhat fleshy, 1-2 dm high, with a taproot.
Herbage glabrous. Basal leaves lacking, the stem leaves
fairly uniform, narrowly obovate, dentate. Heads few,
mostly 2-3 per stem. Longer tegules of two kinds; every
other one broadly hyaline-margined, the others with 4
narrow margin in pale green. Second half of summer. Roc-
ky alpine slopes. --swAlta-seBC, wUS.

The californian var. occidentalis Gray is more
slender and less sharply toothed, while var. blitoides
(Greene) Cronq. of the southern Rockies is more robust
and its broader leaves are more sharply toothed.

7. S. trianguleris Hooker -- Stem leafy with nume-
rous triangular leaves 5-12 cm long, sometimes broadly
cuneate but usually truncate at base, serrate at margin.
Ligules few and rather long. Otherwise pretty much like
S. eremophilus. Mid summer. Open mountain woods and
low alpine meadows. --Mack-Aka, swAlta-BC, nwUS.

8. §. foetidus Howell (S. hydrophiloides Rydb.)
-- Similar to the next but the lower and basal leaves
closely and sharply serrate. Generally larger, especial-
ly the basal leaves. Herbage quite glabrous. Tegules
with a conspicuous, triangular, black tip. First half of
summer. Along low montane creeks -- swAlta-sBC, nwUS.

S. hydrophiloides was some years ago reduced in
rank as S. foetidus var. hydrophiloides (Rydb.) Barkley
and discussed in Leafl. West. Bot. 9: 103-4, 1960¢eR Lhe
more southern and more western parts of the range are
said to be restricted to var. foetidus with more numerous
heads in an irregular compound corymb of small clusters,
the peduncles mostly shorter than the heads. Allowing
for a broad zone of overlap, the more northern and the
more eastern parts of the range are reputedly occupied
by var. hydrophiloides with fewer heads borne in 4@ near-
ly simple corymb, most peduncles longer than the heads.

Of the four Canadian collections examined, the one
from Rossland, B.C., (CAN) fits the distributional pat-
tern by having the morphology of var. hydrophiloides, but
the Alberta collections from Milk Range (CAN), Camp Impee-
sa (DAO) and Waterton (CAN) have the more numerous and
clustered heads of var. foetidus. Obviously the Canadian
material does not fall into the proposed pattern of geo-
graphical varieties. The U.S. material at hand is not
sufficient to enable us to form a firm opinion on the
tenability of these variations south of our borders.

9. S. integerrimus Nutt. var. integerrimus --
Leaves entire or remotely denticulate in the manner of

SENECIO 186

1972 Boivin, Flora of Prairie Provinces 103

some Arnica. Herbage + tomentose or villous, at least in
the axils and at the base of the heads. Lower and basal
leaves lanceolate, commonly around 1 dm long, the middle
and upper ones much reduced. Tegules green to the tip.
Légules yellow. Early summer. Wet or sandy meadows, in-
frequent and mainly more southern. --Man-BC, US -- Var.
exaltatus (Nutt.) Crong. (S. columbianus Greene; S. exal-
tatus Nutt.; S. Scribneri Rydb.)-- Tegules with a small
black patch at tip, the black patch. 1 mm long or less,
mostly lanceolate. --S-BC, wUS -- F. ochroleucus (Gray)
Boivin -- Ligules paler, white to cream: Manyberries.
--Alta-(BC, US) -- Var. lugeng (Rich.) Boivin (S. lugens
Rich.) -- Black patch larger and more conspicuous, trian-
gular to deltoid and about 2 mm long. -- Mack-Aka, Alta-
BC, nwUS.

Var. exaltatus (Nutt.) Cronq. f. gchroleucus (Gray) -
stat. n., S. lugens Rich. var. ochroleucus Gray. Syn. Fl.
1,2: 388. 1884.
~~ Var. lugens (Rich.) stat. n., S. lugens Rich., Bot.,
App. to Franklin's Narrative 747-8, 1823.

Var. Parryi D.C. Baton (=var. exaltatus) was repor-
ted by Dawson 1875 for west of the Turtle Mountain towards
the first crossing of the Souris river, but the correspon-
ponding collection (DAO) belongs to var. integerrimus.

See also Scoggan 1957 sub S. pauperculus. To our know-
ledge, all Manitoba specimen of S. integerrimus belong to
the typical variety.

10. S. canus Hooker (S. Purshianus Nutt.) -- Herba-
ge more or less grayish-tomentose. Basal leaves all or
mostly entire. Otherwise similar to S. pauperculus var.
thompsoniensis. First half of summer. Steppes on hill-
sides, frequent. -- (0O)-Man-BC, US.

Reputedly introduced eastward in Ontario, but we
have yet to see a specimen.

ll. §. streptanthifolius Greene (S. cymbalarioi-
des Nutt., var. borealis (T.& G.) Greenman; S. obovatus
AA.) -- Lower and basal leaves slightly fleshy, entire
except for 3-(5) apical teeth. Otherwise similar to the
next. Drier prairies. Late spring and early summer. --
sMack-sAka, nwS=-BC, wUS.

S. aureus var. borealis T. & G. was reported by
Macoun 1884 for the North West Angle of the Lake of the
Woods. This locality is in Minnesota, the Angle being
@ small geographical inset along the Ontario-Manitoba
boundary. The specimen (CAN) was correctly identified
and would to-day be called S. streptanthifolius; it was
so revised by T.M. Barkley in 1960. However the locali-
ty is far out of range and has never been confirmed, rai-
sing the suspicion that the label data could be erroneous.

187 SENECIO

10) Pony T OFL101G TA Vol, 23, moe 1

12. §. pauperculug Mx. var. pauperculus (S. Balsa-
mitae Muhl.) =-- middling type, forming an intergrading
series with the last two and the next four species, lLoo-
sely tufted, (2)-4-(6) dm high, glabrous of tomentose in
the leaf axils, Leaves strongly dimorphic; the lower and
basal oblanceolate to obovate, mostly 1 cm wide or sligh-=
tly less, petiolate, crenate or serrate and often pinna-
tifid towards the base; middle and upper leaves sessile,
pinnatifid towards the base. Involucre 4-5 mm high. Ear-
ly summer. Wet sandy soils and limestone flats in open
places. --K-(Mack-Aka), L-NF, NS-Man, US -- Var. firmifo-
lius Greenman (var. flavovirens (Rydb.) Boivin, var.
thompsoniensis AA.; S. Tweedyi Rydb.) -- Heads larger, the
involucre 5-7 mm high. Herb tending to be larger through-=
out. Wet meadows. --K-Mack, NF, Q-BC, US -- Var. thomp-
soniensis (Greenman) Boivin (S. plattensis Nutt.) -- Her-
bage more or less floccose-tomentose. Stem leaves more
often pinnatifid to pinnatipartite for their whole length.
Basal leaves usually larger, commonly 1-2 cm wide, cre-
nate to lobed. Heads larger, as in var. firmifolius. -=
sMack, O-BC, cUS.

Our Canadian material is fairly readily divisible
into three geographical variants, give or take a few in-
termediates. The more eastern plants are generally
smaller and smaller-headed, they constitute the typical
variety. The common type in our forested areas, common
westward and northward, becoming very local eastward,
is a somewhat larger plant with larger heads; it may be
known as var. firmifolius. More pubescent plants from
the prairie regions and southward are referable, sometimes

“arbitrarily so, to var. thompsoniensis. All specimens
under S. pauperculus at DAO, MT, MTMG, and QPA from west
and north of Manitoba were revised to other taxa, mainly
to var. firmifolius, Perhaps the western material in
other herbaria should be similarly revised and the Mack-=
Y-Aka reports discounted, just as we are discounting all
reports of S. pauperculs proper from Saskatchewan and
westward.

Canadian reports of, and identifications as, S. Bal-
samitae generally refer to somewhat larger plants from
any of our three varieties, mostly of var. pauperculus.

Var. firmifolius (1905) is apparently the earlier
name for the larger-headed plant which we had previously
called var. flavovirens (1948) and was called var. thomp-
soniensis (1911) by Cronquist 1955.

Macoun 1884 reports Senecio aureus L. var. obovatus
(Muhl.) T. & G. (S. obovatus Muhl.) as occuring from Nova
Scotia to B.C. and throughout the prairie regions. Modern
collections show that in Canada, S. obovatus is restric-
ted to two limited areas in Ontario and Québec and Macoun's
report for our area is no doubt to be discounted, even if

SENECIO 188

1972 Boivin, Flora of Prairie Provinces 105

we have not yet investigated its specimen basis. We ex-
pect that most of his western specimens will belong to
S. pauperculus var. firmifolius or to S. aureus.

13. §, aureus L. (S. so Rydb., var. se=
micordatus (Mack. & Bush) Barkle -- Spring-Avens, Squaw-=
Weed -- Basal leaves large and short, reniform or broadly
ovate to cordate, mostly 2-5 cm wide and broadest at the
base which is mostly cordate to truncate or broadly cu-
neate. Involucre 5-8 mm high. Otherwise similar to the
var. firmifolius of the last. First half of summer. Wet
meadows. -- NF-SPM, NS-sMan-swS-Alta(se, sw)-BC, US.

A troublesome name, often misapplied, so that lite-
rature records should not be trusted too eagerly. Thus
the long-standing Labrador record turned out to be based
on two Forteau collections (QK) which proved to belong to
S. pauciflorus and S. pauperculus respectively.

A highly variable species, difficult to define, not
always clearly distinct from its relatives. Numerous va-
riants have been defined and many of these will form the
dominant facies of the species in a particular region,
but as far as we can determine these variants have pri-
marily a statistical value, being quite common in a4 par-
ticular part of the range, sporadic or local elsewhere.

Bast of us nearly all specimens of S. aureus have
obviously cordate basal leaves and are thus readily dis-
tinguished from S. pauperculus; the latter is also noted
for its smaller heads and narrower leaves + cuneate at
base. But in our area where the local variant of S. pau-
perculus is the larger-headed var. firmifolius and where
the local facies of S. aureus is a somewhat smaller plant
with basal leaves less often cordate than not, the dis-
tinction is less obvious and at times merely arbitrary.

The latest student of the group, Barkley 1962, sub-
divided our Canadian material into three taxa as follows.
The more deeply cordate basal leaves characterize the eas-
tern S. aureus, while in the western S. pseudaureus they
are truncate or merely subcordate at base. The eastern
phase of the latter is distinguished as var. semicorda-
tus (= S. aureus var. semicordatus (Mack. & Bush) Green-
man) in . which the basal leaves have rounded teeth. This
occurs in southern Manitoba and southward. The more wes-
tern and typical phase of S. pseudaureus has acutely ser-
rate basal leaves and occurs from Alberta westward. Gros-
so modo, these distinctions can be applied to our speci-
mens, and so can Fernald's subdivision of the eastern ma-
terial into five varieties. But either classification
leaves behing a large residue of atypical or out-of-range
specimens.

To illustrate the low level of conformity between

_the actual specimens and the described standards, we are

189 SENECIO

106 PibvEs? OLLsO0G TeA Vol. 23, moccL

reproducing a count of heads on @ series at hand of 18
Alberta and B.C. collections comprising 40 flowering
stems. In that area only typical S. pseudgaureaus is
supposed to occur and it should bear 12-20 heads, as
contrasted with the more eastern var. semicordatus bea-
ring only 6-12 heads per plant. The result is as fol-
lows in which the first figure is the number of heads,
and the figure in brackets is the number of plants from
Alberta and B.C. bearing said number of heads: 2(1)--
3(6)-- 4(4)-- 5(2)-- 6(6)-- 7(4)-=- 8(3)-- 9(4)-- 10(1)
14(2)-- 15(2)-- 16(2)-- 17(1)-- 20(1)-- 22(1). Clearly
we have two series here: 31 plants have 2 to 10 heads per
inflorescence and would be better placed with the mani-
toban var. semicordatus; the remaining, a minority of

9 plants, have 14 to 22 heads and conform roughly to the
standard of S. pseudaureus as expected for the area. Ob=
viously, the number of heads per plant has no diagnostic
value in the present case. Other criteria have proved

to be equally unsatisfactory.

Alleged differences in root system are equally un-
convincing, granted that the more western plants tend to
have a somewhat thicker and more horizontal rhizome than
the eastern plants.

The species is of discontinuous distribution across
our area, being apparently restricted to southern Manito-
ba, the Cypress Hills and southwestern Alberta. We have
seen three of the Saskatchewan collections cited by
Breitung 1957 as S. pseudoaureus, but we place these in
the radiate form of S. indecorus (or S. discoideus), a
species not later recognized by Breitung. On the other
hand another Cypress Hills collection, Breitung's 4513
(or "4313") falls within our concept of S. aureus. It
was originally named and distributed as S. pseudoaureus,
but cited by Breitung 1954 as S. indecorus, then in 1957
included in S. pauperculus var. thompsoniensis, finally
in 1959 returned to S. pseudoaureus.

Some Alberta and B.C. specimens have oblong-lan=
ceolate basal leaves and would probably have been iden-=
tified as S. Robbinsii Oakes if they had been collec-
ted in the east.

S. aureus grades into the next species, but the
two are largely allopatric and generally quite distinct.
Typically S. aureus has + cordate basal leaves, radiate
heads and the involucre is green, although frequently
tipped in red, while S. indecorus has ovate basal leaves,
discoid heads, and a purplish=-tinged involucre. However,
smaller plants of S. aureus is our area will quite often
have ovate basal leaves, while on occasion these may be
subcordate in S. indecorus. Exceptional individuals may
be discoid in S. aureus while the radiate form of S. in-=

SENECIO 190

1972 Boivin, Flora of Prairie Provinces 107

decorus is not infrequent. And exceptional specimens of
S. indecorus may have 4 green involucre. On the basis of
the distributional patterns of the typical specimens and
of field associations, we judge that specimens that are
morphologically intermediate are more likely to relate

to S. aureus if the head is discoid and the involucre
green, but to S. indecorus if the head is radiate and the
involucre purple-tinged. The latter also runs to higher
heads. In practice the purplish condition of the involu-
cre is a more reliable characteristic of S. indecorus than
its discoid presentation.

14. S. indecorus Greene (S. discoideus (Hooker)
Britton) -- Mostly like a larger var. firmifolius and
transitional to the next species. Generally a larger
plant, usually 6-8 dm high. Leaves broadly lanceolate,
mostly pinnatipartite towards the base, the basal ones
broadly oval, mostly 2-3 cm wide. Heads discoid. Tegu-
les purplish. Involucre 7-10 mm high. Mid summer. Moist
meadows. --(seK)-Mack-Aka, Q-BC, nUS--F. Burkei (Green-
man) Fern. (S. pauciflorus Pursh f. fallax (Greenman) Boi-
vin) -- Heads radiate, hence resembling a smaller S. au-
reus, but the involucre purplish and the ligules rather
short, usually not over 5 mm. Occasional and sometimes
nearly as common as the discoid form, --Mack-Y, O, S-BC.

Barly reports of S. discoideus will be found to ap-
ply indifferently to this or the next species. The dis-
tinction in two taxa was introduced by Fernald, Rhodora
26: 116-122, 1924, but we disagree with his interpretation
of the name S. discoideus, The type of S. discoideus was
collected by Richardson at Fort Franklin on the Great
Bear River and, working from a photograph, Fernald con-
cluded that it was intermediate but best placed with S.
pauciflorus on the basis of the leaf shape, although it
resembled S. indecorus in its more numerous heads. Bark-
ley 1962 accepted Fernald's disposal of S. discoideus, but
Richardson's track, as mapped by Hooker 1840, lies outsi-
de the range S. uciflorus, Raup's 1947 map as well as
Barkley's finely dotted distribution maps show clearly
that Hooker's type came from the northern edge of the
area of S. indecorus and some 250 miles away from the
nearest known occurrence of S. pauciflorus. We judge the-
refore that the type of S. discoideus cannot but belong
with the polycephalous S. indecorus. Further Raup 1947
cited an apparent isotype (CAN) of S. discoideus under
S. indecorus. Thus is justified our disposal of S. dis-
coideus as a synonym of S, indecorus, while the more re-
cent reports of S. discoideus should be interpreted mos-
tly as S. pauciflorus.

As a binomial, S. discoideus first appeared in T.

& G. 1843 in the discussion of the synonyms, hence was

191 SENECIO

108 PAHVIAT OfL/08G Tok Vol. 23, no. i

not validly published at the time. Its valid publica-
tion by Britton 1898 is antedated by that of S. indecorus
Greene 1897, hence the present choice of correct name.

15. S. pauciflorus Pursh (S. discoideus AA.) --
Involucre 6-8 mm high and nearly always more or less pur-
plish, the ligules nearly always lacking. Mostly 1-4 dm
high. Basal leaves less than 2 cm wide, + ovate, roun-=
ded to broadly cuneate at base. Heads few, mostly 3-5.
Disk florets tending to red-orange. Mid summer. Arctic
or alpine meadows. -- K-Aka, L-NF, Q-nMan, Alta-BC, US.

This and the last are quite closely related and
their intraspecific variability is sufficiently wide that
an interspecific hybrid would be difficult to detect and
even more so to define. Two of Calder's collections
from B.C. and Yukon were distributed as such a hybrid,
but by their more numerous heads and higher involucres
we judge them to be better placed with S. indecorus.

16. S. tridenticulatus Rydb. (S. densus Greene;
S. manitobensis Greenman; S. plattensis AA.) -- Some-
what fleshy and the leaves all similar, all lobed to
pinnatifid, and not more deeply so towards the base.
Tufted, glabrous, about 2 dm high. First half of sum-
mer. Wind-eroded sands, very local. --swMan-(scS),
cUS.

A collection from Stewart's lake Mountain, B.C.
(CAN) was a syntype of S. manitobensis Greenman, Ott.
Nat: 25: 117, 1911 and was mentioned again as S. triden-
ticulatus in Ann. Miss. Bot. Gard. 3; 180, 1916. The
specimen was recently reexamined and revised to S. strep-
tanthifolius.

17. S. resedifoliug Less. (S. cymbalarioides
Buek; S. subnudus DC.) --Monocephalous perennial with
at least some of the leaves deeply lobed to pinnatifid.
Glabrous, 1-3 dm high. Basal leaves ovate, mostly den-
tate. Lower stem leaves deeply cut, the upper greatly
reduced, Tegules green or purplish, glabrous. Ligules
often lacking. Early to mid summer, Alpine boggy mea-=
dows and shale slides, --wF, wMack--Aka, (NF, seQ),
swAlta-BC, (nwUS, eHur).

Further west, there is an endemic variant in the
Queen Charlotte Islands: var. moresbiensis (Calder &
Taylor) stat. n., S. cymbalarioides Nutt. ssp. mores-=
biensis Calder & Taylor, Can. Journ. Bot. 43: 1399.
1965, somewhat more pubescent, the involucre being
slightly lanate at base, and the rosette leaves more
uniformly crenate or serrate right to the base. It was
by mistake that in 1967 this variety was listed under

S. streptanthifolius.

49. CALENDULA L. POT MARIGOLD
Achenes all peripheral, strongly curved and either

CALENDULA 192

1972 Boivin, Flora of Prairie Provinces 109

winged on either side or strongly rugose or acicular dor-
sally. Heads radiate, the ligulate flowers fertile, the
tubular ones sterile and long stipitate, the stipe not
dehiscent. No chaff, no pappus.

1. C. ARVENSIS L. == Gools (Souci des champs,
Fleur de tous les mois) -= Head fairly large and radiate,
like an Aster, but yellow and the rather unusual achenes
borne only at the periphery. Herbage strongly glandular,
Tegules isomegueth, abruptly caudate. Summer and fall.
Rare and fleeting escape from cultivation: Brandon.--NB,
Man, BC, swUS, Eur.

50. ECHINOPS L. GLOBE-THISTLE

Head compound, made up of a large number of primary
heads, each one reduced to a single floret and its involu-
cre.

a. Stem tomentose, the tomentum becoming white and
compact in the upper part ....soecseece Lo BE. exaltatus

aa. Tomentum mixed with numerous long, coloured and
Glandular hairs ....c.se.ecreeoe 2. EB. sphaerocephalus

1. E. BXALTATUS Schrader -= Rather closely simi-
lar to the next. Little if at all glandular. Leaves lar-
ger and more narrowly cut, the lobes + lanceolate. Tegu-
les glabrous on back. Second half of summer. Persisting
after cultivation in a city garden at Swift Current. --
swQ-O, swS, swBC, Eur.

2. 5B. SPHAEROCEPHALUS L. -- Globe-Thistle (Boulet-
te, Chardon-boulette) -- Head globular with the recepta-
cle at the centre of the sphere. Leaf resembling Cirsium
by its cutting and its excurrently spinescent nerves.
Heads few, 3-6 cm across, bluish, spinescent, borne on
long peduncles. Tegules puberulent dorsally. Mid summer.
Sometimes cultivated and rarely spreading to waste places:
Otterburne. -- swQ-seMan, BC, (US), Bur.

Reported from Regina and Saskatoon by Bussell 1937,
1944, 1954 and Breitung 1957. We have not been able to
tie these reports clearly to any herbarium specimen. Of
the two possible sheets located, Dr. G.W. Argus commen-
ted(in litt., 1964) on the first one (SASK) "there is no
locality or date on this specimen and it seems unjusti-
fied to assume that it is of Saskatchewan origin", The
other sheet came from Landis (SASKP) and we have revised

it to Eryngium planun.
51. ARCTIUM L. BURDOCK
Fruits very catchy as follows: the tegules are at-
tenuate to a fine, hooked point, they are also fused at

base and divergent at tip to form a globular unit which
becomes very readily detached from its peduncle.

193 ECHI NOPS

110 PH TIT OF LOG I-A Vol. 23, no.cd

a. Inflorescence broadly corymbiform, the lower
heads on long peduncles.
b. Involucre glabrous, 3 cm wide or more ooo.
ais 0 diein state se Oo Sie wlsseielvisieleleie Clee aja w'ele a nici ).L aN mentees
bb. Narrower and densely tangled with an ara-
chnoid tomentum .....caccccececce 2. A. tomentosum
aa. Inflorescence a broad panicle of racemes of small
clusters; lower heads on peduncles usually short,
rarely over 3 cm long.
ce Involucre glabrous or glandular ..... 4. A. minus
cc. Tegule tips more or less tangled with an
arachnoid tomentum ....-.-+cccacecoo 3a Ao nemorosum

1. A. LAPPA L. -- Great Burdock, Cukle-Buttons
(Grande Bardane, Glouteron) -- Heads largest, 3-4 cm
wide, hemispheric and glabrous, forming one or more broad
corymbs. Tegules green with ivory tips. Otherwise simi-=
lar to A. minus. Second half of summer. Waste places
and foothpaths, a rare weed with us. -- (NS), NB-Man,

Be, US, Bux.

2. A. TOMENTOSUM Miller -- Also similar to A. mi-=
nus, but the inflorescence + corymbose and each head is
wrapped in a loose cocoon of arachnoid tomentum ancho-
red near the tips of the tegules. Heads subglobose, 2-3
cm wide. Second half of summer. Roadsides and foot-=
paths. --NS, NB-Alta, (US), Eur.

3. A. NEMOROSUM Lej. & Court. -= Heads tangled with
an arachnoid tomentum and often larger, up to 3.5 cm wide,
and broadly globular, i.e. slightly broader than high.
Otherwise pretty much like the next and perhaps not spe-=
cifically distinct from it. Mid summer. Waste places
and footpaths; rare. -- NF, NS, (NB)-Q-Man, Alta-BC, US,
Eur.

We have checked specimens from Otterburne (MSM),
Saint-Pierre-Jolys (DAO), and Edmonton (DAO).

4, A. MINUS (Hill) Bernh. == Burdock (Bardane)
Coarse herb with very catchy fruits which readily beco-
me detached from their peduncle to attach themselves
firmly to the clothing of the passerby. Basal rosette
similar to Rhubarb, but the leaves somewhat smaller,
ovate, arachnoid below. Heads 2-3 cm wide, glabrous to
glandular, globular-ovoid. Tegules at first green with
ivory tips, becoming purplish at maturity. Second half
of summer. Waste places and footpaths; infrequent. -=
NF, NS-BC, US, Bur.

Some conspicuous foliar anomalies may be found.
They are apparently related to tramping or herbicide ac-
tion.

52. SAUSSUREA DC.

Heads discoid. Pappus long and plumose. Leaves

ARCTIUM 194

1972 Boivin, Flora of Prairie Provinces i

alternate and not spiny.

a. Tegules acuminate at tip, isomegueth..... 1. S.- nuda
ee. Dilated at tip into a suborbicular and
petaloid segment ...cecescccccecseeee 2e Se glomerata

1. 3, nuda Led. var. densg (Hooker) Hultén --

Short perennial with a small terminal corymb of large
discoid heads. Usually less than 2 dm high. Leaves ma-
ny, crowded, lanceolate, + dentate, somewhat arachnoid,
especially along the margin. Heads 1.5-2.0 cm high. Te-
gules usually dark purple. Mid summer. High alpine on
rocky slopes. --swAlta-sBKC.

The typical phase is alaskan and eurasian; its stem
leaves are rapidly smaller above, those from the upper
half entire and narrowly linear or nearly filiform; it
inflorescence overtops the foliage. In our var. densa the
upper leaves are much less reduced, being at least half
as large and half as long as the lower ones; and the in-
florescence is + overtopped by the upper leaves.

2. S. GLOMERATA Poiret -- Inner tegules with a4 pe-
taloid terminal segment, 1-2 mm wide, pink, + fimbriate
at margin. Stoloniferous perennial up to 4 dm high. Lea-
ves entire, densely glandular-punctate in yellow-brown be-
low. Outer tegules many times shorter than the inner.
Late summer. Rare farmyard weed: Debolt near Grande-Prai-
rie. ==wcAlta, Eur.

53. CARDUUS L. PLUMELESS THISTLE

Resembling Cirsium, the leaves similarly spiny.
However the pappus is not plumose, but merely short-bar-
bellate.

1. C. NUTANS L. var. VESTITUS (Hal.) Boivin (var.
Petrovicii Arénes, ssp. leiophyllus (Petrovic) Arénes;
C. Thoermeri Weinm.) -- Musk-Thistle, Nodding Thistle
(Cardinal, Chardon aux @nes) -- Large and ferociously
spiny herb, usually monocephalous, the head very large.
Stem 1-3 m high, spiny from decurrent wings. Leaves
spiny and cut in the manner of a Cirsium, glabrous dor-
sally. Head purple, discoid, 4.0-5.5 cm wide. Tegules
large, spiny tipped, becoming reflexed. Lateral heads,
if present, smaller, Mid summer. Obnoxious weed of
roadsides and pastures, still local but spreading. --
Q-S, BC, US, Bur.

Another variety also occurs west of us at Alexis
Creek in B.C.; var. macrocephalus (Desf.) stat. ne, C.
macrocephalus Desf., Fl. Atl. 2: 245. 1798-1800, heads
larger, 5-6 cm wide; leaves somewhat arachnoid-pubescent
dorsally.

The shape and coloration of the tegules vary and
specimensin which the upper and more colourful half is

195 SAUSSUREA

112 Paha ORL AO8G: 7A Vol. 23, no. 1

broader than the lower pale green half have been distin-=
guished as var. macrolepis (Peterm.) Rouy (=C. Thoermeri).
This phenotype has been recognized in our area, but its
significance, if any,eludes us.

54. CIRSIUM Miller THISTLE

Very spiny from the leaf=-nerves long=-excurrent into
needlelike points. Heads large, discoid. Pappus plumose.

a. Leaves decurrent with spiny wings from node
to node and up to the base of the headSecece
Coc ccscccreccecccescccecesecccsssescace Le C. vulgare
aa. Leaves not decurrent, or at least not in the
upper half of the plant.
b. Inner tegules ending in a twisted scarious
appendage, the outer tegules spine-tipped..
coer er ce sree sccececesseseecacce Of Co Drummondii
bb. All tegules sharp=pointed, or spinetipped;
heads smaller.
ce Involucre 1.0-1.5 cm high .aeoe 9. Co. arvense
cc. Heads larger.
d. Heads overtopped by the uppermost
or subtending leaves,
e. Tegules straight, their tips
appressed; herbage very long-=-
Villous c.sarceeceeeee 7o Co foliosum
ee. Tegules squarrose, ending in
spines directed outward; her-=
bage thinly arachnoid-tomen=-
tOSE sorceress eeeeees 8. C. Hookerianum
dd. Heads overtopping the foliage.
f. Stem white-tomentose.
go Leaves + flat, the lobes
lanceolate and narrower
than the sinuses..2. C. Flodmanii
gg. Leaves strongly crisped,
the lobes + deltoid and
+ overlapping se. 3. C. undulatum
ff, Stem green, not tomentose.
h. Tegules ending in a de-~
flexed spine; leaves
white-tomentose below..

eaeceeseceeeeacncond 4, Co altissimum
hh. Tegules merely sharp

pointed; leaves only
paler green below .oce
eccesesececeacvaa0aa00 Sle C. muticum

1. C. VULGARE (Savi) Tenore (C. lanceolatum AA.) --
Bull-Thistle, Scotch Thistle (Gros chaudron, Piqueux) r+
Herbage spiny throughout, even the upper leaf surfaces

CARDUUS 196

1972 Boivin, Flora of Prairie Provinces 113

acicular-hispid. Biennial, mostly about 1 m high and
unépproachable. Leaves + arachnoid below. Heads ten-
ding to be overtopped by the upper leaves. Florets pur-
ple. Mid to late summer. Rare weed, usually near dit-
ches or creeks, --NF-(SPM), NS-BC, (US, Eur).

The range was extended to Alaska by Hultén 1950
and Anderson 1952, but the main justifying specimen, An-
derson 5573, Hyder, 1939 (S) turned out to be a sterile
shoot of C. arvense (L.) Scop. Reports from Sitka and
Salmon River Glacier have not been investigated.

2. C. Rlodmanij (Rydb.) Arthur (C. oblanceolatum
Rydb.; C. plattense AA.) -- Long-stoloniferous perennial
with the rosette-leaves polymorphic, some of them unlo-
bed and merely spinulose-margined. Stem and lower leaf
surfaces covered by a thin and compact tomentum. Upper
leaves smaller and less deeply lobed, often unlobed even.
Involucre narrowly campanulate, 2.0-2.5 cm high, 1-2 cm
wide at base in the herbarium. Florets pinkish. Seeds
3.5=-5.0 mm long. Mid summer. Common in prairies. --
swQ-Alta, US -- F. albiflorum D. Live -- Florets white or
cream-coloured. -- Man-Alta, US.

Rather variable and herbarium specimens seem to be
readily confused with the next. In the field the diffe-
rence is quite striking. The few rosette-leaves of C.
Flodmanii are in part flat and unlobed. The heavily
crisped leaves of C. undulatum are all equally lobed and
gathered into dense rosettes. Both species may seem to
be biennial, but some patient digging (or a handy road
cut) will reveal deeply buried and rather extensive root-=-
connections.

3. C¢. undulat (Nutt.) Sprengel var. undulatum
(var. megacephalum (Gray) Fern.; C. Engelmannii AA.) --
Woolly Thistle -- Much like the last and not so obvious-
ly stoloniferous, but all the leaves cut alike and all
strongly contorted in the sinuses. Upper leaves are gra-
dually smaller. Heads larger, campanulate to hemispheric,
2.5=-3.0 cm high and 2.0-3.5 cm wide at base in the herba-
rium. Florets purplish-red. Seeds 5-7 mm long. Summer.
Drier prairies and steppes, from Dalny westward. --SwMan-
BC, US -- F. album Farw. (C. brevifolium AA.) -- Heads
white: Milden, Maple creek. --swS, US.

Often mentioned for Manitoba but all collections
examined appear to have been misidentified except the
following: Boivin 13434, riviére Souris 4 l'ouest de Dal-
ny, €corre de la coulee, 3 juin 1960 (DAO).

A more western var. Franktonis Boivin has pink flo-
wers and smallish heads as in C. Flodmanii, but the seeds
a@re larger as in C. undulatum. The technical justifica-
tion: C. undulatum var. Franktonis var. n., capitulis modo
minoribus; involucro 2.0=2.5 cm alt.; corolla rosea, post

197 CIRSIUM

114 Pulp Yr T Op LsQ0G> Lod Vol. 23, no. 1

anthesim albescens; sed semina + 6 mm long. et ceteris
praecipue ad C. undulatum vergens, attamen foliis sae=-
pius minus crispatis. Typus: Calder & Savile 9838, at
foot of Mt. Anarchist, just east of Osoyoos, common in
open sagebrush slopes above lake; June 29, 1953 (DAO).
Paratypi omnes ex DAO: Calder & Savile 10397, Spences

Mulligan & Woodbury 1931, Pentiction; Calder, Parmelee &
Taylor 19115, Williams Lake.

Named after Dr. C. Frankton, a long time student
of the genus.

4, C. altissimum (L.) Sprengel var. disgolor
(Muhl.) Fern. (C. discolor (Muhl.) Sprengel) -=- Leaf
surfaces strongly contrasted: white-tomentose below, dark
green and lightly villous above. A rather middling spe-
cies with pinnatitid to nearly pectinatipartite leaves,
strongly scabrous above. Stem green, mostly 1-2 m high,
with very large basal leaves. Late summer. Wet meadows
and marshy shores: Emerson, =-swQ-scMan, US.

The Bmerson (DAO) collection is the only one seen.
Old reports for our area of Cnicus altissimus var. disco-
lor and of Cirsium altissimum should probably not be in-=
terpreted in the sense of the above var. discolor. Daw-
son's 1875 report for the Rockies undoubtedly meant so-
mething else. And his Turtle Mountain report is probably
based on C. Flodmanii, if we are to judge from a Burgess
collection (TRT) made 5 days later. The Winnipeg report
by Macoun 1897 was based on a sheet that either did not
survive or else has been revised since to something else,
possibly C. Flodmanii.

In the typical and more southern var. altissimum
the leaves are less deeply cut, being merely serrate to
lobed. The two varieties are reported to be completely
intergradient in their area of sympaty, but all the Cana-
dian material examined was clearly referable to var. dis-
color.

Se C. muticum Mx. -= Dunce=Nettle, Horsetops -=- Te-
gules not ending in a squarrose spine, but the middle and
outer ones merely mucronate while the inner ones are atte=
nuate into petaloid and scarious tips. Main leaves pin-=
natipartite, arachnoid below, weakly acicular-ciliate.
Heads purplish. Tegules arachnoid. Mid to late summer.
Marshy or boggy places. -=L-NF-(SPM), NS=-cS, US -- F. lag-
tiflorum Fern. -- Flowers white. Wallwort. == NF, Q, S.

6. ¢. Drummondii T. & G. (C. foliosum AA.; C. Hil-
lii AA.) -- Head largest, 5-8 cm across and typically so-

CIRSIUM 198

1972 Boivin, Flora of Prairie Provinces 115

litary, its involucre + 4 cm high. Stem low, thick, fis-
tulous, easily crushed, sometimes lacking, often short,
always very leafy and the leaves rather long, the upper
overtopping the inflorescence. Herbage long villous.
Heads sometimes more than one, then 3-5 in a terminal
cluster, the lateral heads smaller. Shortly before mid
summer. Chernozems at forest margin; infrequent. --
sMack, wO-eBC, ncUS.

7e Q- foliosum (Hooker) DC. var. foliosum -- So-
mewhat like the last, but the heads not quite so large
and the corrollas whitish. Herbage also quite similar to
the last and similarly long villous. Leaves very nume-
rous and the upper much overtopping the inflorescence.
Heads always many in a crowded terminal cluster. Involu-
cre 2,0-2.5 cm high, the tegules all spine-tipped. Pap-
pus pale brown or grayish and conspicuous, overtopping
the corollas. Mid summer. Mountain meadows, down to
lowland meadows northward. --sMack-(sY), sAlta-BC, US.

Leaves mostly green and pilose below, or sometimes
+ white-tomentose. Tegules all appressed. Further east
there is a highly isolated var. minganengs (Vict.) stat.
no, C. minganense Vict., Mém. Soc. Roy. Can. 19: 81.

1925, which has the pappus only as long or slightly shor=
ter theta the pink corollas. Also the rosette leaves are
white-tomentose below; stem leaves variable, mostly pi-
lose below; inner tegules squarrose, twisted and slight-
ly dilated towards the tip. Known only from the shores

of some of the Mingan Islands in the Gulf of Saint-Lawren-
ce.

Var. minganense has also been treated as &n out-=
right synonym of C. scariosum in Can. Journ. Bot. 45:
1742, 1967, althoug it presents itself more like @ va-
riety iiteniediate between C. foliosum and C. Hookeria-
num. It is quite close to c. foliosum because of its
habit, its denser inflorescense and its grayish pappus.
It shown some affinity to C. Hookerianun in its tomento-
se pubescence, its short pappus, its squarrose inner te-
gules and somewhat larger seeds.

8. €. Hookerianum Nutt. var. Hookerianum - Heads
also whitish like the last but the pappus not so conspi-
cuous, being overtopped by the corollas. Pubescence mo-
re tomentose and at least the lower leaves white-tomento-=
se below. Inflorescence very variable, corymbose or pani-=-
culate to monocephalous, typically racemosely paniculate
with a terminal cluster of + 3 subsessile heads and many
axillary clusters of 1-3 heads on short peduncles. Te-
gules glandular and the middle ones somewhat villous and
long ciliate, the outer and middle ones arachnoid-tomen-=
tose. Pappus white and shorter. Summer. Shale slides
and alpine or subalpine meadows. -=-swAlta-BC, US -=- Var.

199 CIRSIUM

116 PuBy YT OFLYOuG. ToA Vol. 23, novel

scariosum (Nutt.) Boivin (C. scariosum Nutt.) - Tegules
not squarrose, except the inner, and less pubescent,
merely glandular or the outer somewhat arachnoid=tomen=
tose, From the Crowsnest southward. --swAlta, nwUS.

Var. scariosum (Nutt.) state ne, C. scariosum
Nutt., Trans. Am. Phil. Soc. 7: 420. 1841.

9. C. ARVENSE (L.) Scop. (var. integrifolium
Wimm. & Grab., var. mite Wimm. & Grab., var. vestitum
Wimm. & Grab.) -- Canada Thistle (Chadron, Chaudron) --=
Heads smallest. About 1m high and growing in dense
colonies. Heads at first few and corymbose, becoming
many and narrowly paniculate. Inner tegules not spiny,
the outer very short spiny. Florets purplish. Mid to
late summer. Common and invading weed. -= G, Mack, Aka,
NF-SPM, NS-BC, US, Bur, (Afr) -- F. ALBIFLORUM (Rand &
Redf.) R. Hoffm. -- Heads white-flowered -- NF, NS-BC,
US, Eur.

54, SILYBUM Adanson MILK-THISTLE

Resembles Cirsium but the pappus bristles are not
plumose, not even barbellate, and the tegules are cons-
tricted towards the middle to delimitate two segments,
the upper spiny-tipped as in Cirsium, the lower acicular-
ciliate.

1. S. MARIANUM (L.) Gaertner -=- Milk-Thistle, La-=
dy's Thistle (Chardon-Marie) -- Like a huge Cirsium and
the leaf nerves,except the midnerve, outlined by a broad
white strip. Foliage spiny in the Cirsium manner, but
huge, the lower leaves 5-10 dm long. Heads very large,
the terminal 5-8 cm wide. Mid summer. Rare and flee=
ting weed of cultivated ground, sometimes seeded in as
an ornamental, --NS, NB-O, S, BC, US, SA, Eur.

56. CENTAUREA L. STAR-THISTLE

Tegules, at least the inner ones, more or less
clearly differenciated into a lower and an upper segment
in the manner of the last genus. Terminal segment pal-
mately lobed or fimbriate, the fimbriae sometimes spines=
cent. Herbage otherwise not spiny. Pappus variable, chaf-=-
fy or bristly or none. This genus not readily defined
except that the achenes are attached obliquely to the re-
ceptacle.

@. Involucre longespiny «<«.s.«scoccce 2a, CG. solstitialis
aa. Not spiny, sometimes with short acicules.

b. Leaves pinnatipartite to bipinnatipartite;

EVOWERS, YL LOW, 6) <.cier icberenens <) ovals 0's) 0 of etal |G my CUerem tees

bb. Entire to dentate; flowers pink or blue.

c. Florets blue, the peripheral ones

much enlarged acosececccerseeeee Se ©

cc. Pink and all alike .c.cecesscreece 4. Co. Depens

SILYBUM 200

1972 Boivin, Flora of Prairie Provinces 117

1. C. DIFFUSA Lam. -= Tip of tegule pectinate, the
lobes very stiff, the central one 2-3 times longer and
almost acicular. Biennial, often tufted, very scabrous.
Leaf segments 1-2 mm wide. Heads narrow, the involucre
about 1 cm high. Corollas sometimes fading pink. Mid
summer. A railway weed at Grassy Lake. --sAlta-BC, US,
Bur.

2. C. SOLSTITIALIS L. -= Barnaby's Thistle, Yel-
low Star-Thistle (Chardon doré, Auriole) -- Non spiny
herb except for the heads ferociously armed with yellow
spines. Herbage tomentose. Stem winged from the decur-
rence of the linear leaves. Spines widely divergent, the
main ones longer than the body of the head. Florets yel-
low. Late summer and fall. Rare garden weed: Shellmouth,
Ogema, Scott. --sO-S, US, (Eur).

3. C. CYANUS L. -=- Cornflower, Bluebottle (Bleuet,
Barbeau) -- Peripheral florets much longer and much larger
than the inner ones, simulating a blue head radiate in
blue. Narrow-leaved annual. Middle and inner tegules nar-
rowly lobed at tip. Summer and fall. Casually reseeding
itself along roadsides and waste places after cultivation.
--NF, NS-Man, Alta-BC, US, Eur.

4. C. REPENS L. (C. Picris Pallas) -- Russian Knap-
weed, Turkestan-Thistle -- Tegules ciliate at tip, the in-
ner ones abruptly contracted into a long, plumose bristle.
Perennial from long and deeply buried stolons. Main lea-
ves dentate, the others entire and much smaller. Heads
pink, few, in a corymbose inflorescence at the end of long
and leafy branches. Mid summer to early fall. Uncommon
weed of fields. --O-BC, US, Eur.

57. CICHORIUM L. SUCCORY

Florets all ligulate, but the pappus not plumose,
@ mere ring of small scales. Flowers blue.

a. Heads all or mostly longer than their bracts ..
eoeeeeveeveveeeeveoeseeeeeeeeoeee eos ooeeogese ee ed le Cc. Intybus
ea. Bracts much longer ecssccccccecsecacces 4e Ce Endivia

1. C. INTYBUS L. -=- Chicory, Blue Sailors (Chicorée,
Chicorée sauvage) -- A branchy and nearly squeletic pe-
rennial with large, blue, ligulate flowers. Milky juice
white. Rosette leaves large, + runcinate, the cauline
ones few and much reduced. Flowers in small, distant
glomerules with bracts mostly under 1 cm long. Outer te-
gules reflexed, at least after flowering. Pappus minute.
Mid to late summer. Casual weed of roadsides, waste pla-
ces and neglected gardens. =-- L-SPM, NS-BC, US, SA, Eur.

2. C. BNDIVIA L. -- Endive (Chicorée endive) -- Simi-
lar but the inflorescence more leafy, the glomerules being

201 CENTAUREA

118 PHT O'L/0°G Tk Vol. 23, no. 1

subtented by triangular bracts mostly 2-5 cm long. Pappus
up to 1/2 as long as the seed. Mid summer. Rare escape:
Kinistino, Lethbridge --S-Alta, Eur.

58. LAPSANA L. NIPPLEWORT

Like the last, but the flowers pale yellow and the
pappus lacking or vestigial.

1. L. COMMUNIS L. == Nipplewort, Swine's Cress
(Herbe aux mamelles, Saune blanche) -- Tegules dimegueth,
the inner about 8 in number and 5-6 mm long, the outer
about 5 and + 1mm long. Herbage hirsute. Upper leaves
lanceolate and sessile, the middle ones ovate with a nar=
rowly winged petiole, the lower ones lyrate-pinnatipartite.
Heads small in an open inflorescence. Mid summer to fall.
A rare weed of shaded places, reported from Winnipeg. ==
G, Aka, NF, NB-O-(Man), BC, US, Eur.

59. MICROSERIS D. Don

Scapose or nearly so and generally resembling Ago-
seris, but the pappus subsessile. One species is atypi-=-
cal, the achenes being tapered at tip.

a. Leaves all basal; tegules isomegueth .cacec

Rol ae © ass o\p's © cin wele © e okcienavebetae evele ainrwie! = cia Me) Mle ech eeee
aa. At least one stem-leaf; tegules strongly

Gimegueth ercccoccccccsccsensccccccccee Le M. nUtans

1. M.-. nutans (Meyer) Schultz-Bip. -- Herbage some=
what farinose=-puberulent with small vesicular hairs re=-
sembling those of Chenopodium. Habit of Crepis, but the
stem leaf (or leaves) borne towards the base. Stem often
becoming branchy. Leaves eciliate, long=linear and entire
or more commonly pinnatipartite, the lobes few, narrow
and remote. Involucre 1=2 cm high, the inner tegules at
least twice as long as the outer, First half of summer,
Open slopes in the mountains: Waterton. --swAlta-BC, wUS.

2. M. cuspidate (Pursh) Schultz-Bip. -- (Agoseris
cuspidata (Pursh) Raf.; Nothocalafis cuspidata (Pursh) Gree-
ne) -- Leaves tomentose=ciliate. Peduncle + tomentose
towards the summit. Pretty similar to Agoseris but the
latter flowers later and its leaves are eciliate, Mid
spring. Steppes on hillsides. --swMan-sS-sAlta, US.

Rather uncommon, and often confused with Agoseris,
the latter eciliate. Most early reports are to be taken
with @ grain of salt. We know of only one Alberta sheet,
@ Dawson collection from the Milk River (CAN), but we
have checked 5 or 6 from Saskatchewan. For Manitoba we
have checked collections from south of Minto (CAN) and
Brandon (DAO). An early report from Fort Ellice (MTMG)
by Macoun 1884 was more recently listed as Agoseris
agrestis by Scoggan 1957; it has since been revised to
A. glauca. A report from Kleefeld by LYve 1959 has not

LAPSA NA 202

1972 Boivin, Flora of Prairie Provinces 119

been checked.
60. KRIGIA Schreber DWARF DANDELION

The pappus appendages dimorphic, the 5 outer ones
being very short, hyaline and inconspicuous scales, whi-
le the numerous inner ones are capillary bristles. Ache-
ne beakless.

1. K. biflora (Walter) Blake (K. amplexicaulis
Nutt.) -- Cynthia, False Dandelion -- Subscapose perennial,
the leaves mostly basal, but with 1 stem leaf or at least
with 1-3 bracts subtending the forks. Herbage glabrous
or the peduncles glandular. Leaves resembling Agoseris.
Heads few, yellow, the upper 3 on subequal peduncles. Te-
gules about 10, isomegueth, 7-10 mm long. Late spring.
Open sandy woods, rare: Teulon and region. -=-swO-Man,

US.

61. HYPOCHAERIS L. CAT'S EAR

Receptacle chaffy. Pappus of plumose bristles at
the end of a thin long beak. Tegules strongly imbrica-=
ted.

1. H. RADICATA L. -= Cat's Ear, Fall-Dandelion,
(Salade de porc, Herbe & 1'épervier) -- Stems and bran-
ches bearing many very small bracts; the leaves all ba-
sal and very coarsely hirsute. Resembles Crepis runci-
nata, but the latter has a sessile pappus and nearly
glabrous leaves, or at least less pubescent than the in-
florescence. Peduncles very long, glabrous, slightly
thickened upwards. Tegules dark green with a pale green
midnerve which becomes purplish and thickened or barbed
towards the tip. Outermost tegules very small and for-
ming an ill-defined calycule. Ligules yellow, but the
outer ones green dorsally. Late summer and early fall.
Rare garden weed: Scott. --(Aka, NF)-SPM, NS, NB-O, S,
BC, US, SA, Eur.

62. PICRIS L.

Leafy-stemmed herb with a pappus of plumose bris-
tles borne at the end of a very long beak. Tegules di-
megueth or dimorphic.

1. FP. BCHIOIDES L. var. ECHIOIDES --Ox-Tongue
(Langue de boeuf) -- Outer tegules rather large, trian-
gular-cordate and acicular=hispid, especially acicular-
ciliate. Herbage acicular=hispid throughout, almost
like some Borage. Stem leaves alternate, becoming sub-
opposite or subverticillate in the inflorescence. Outer
tegules 5-8 mm wide and 1-2 cm long, the inner narrower
and lanceolate. Late summer and fall. Rare weed of
waste places: Prince Albert, Grande=Prairie. --(NS), NB,
O, S-Alta, US, SA, Bur.

203 PICRIS

120 Po YeT OPLiOiG TA Vol. 23, no. 1

Ours is the typical plant with an involucre + 1 cm
high and dimorphic pubescence, the longer and black hairs
being + 1 mm long. The beringian var. kamtschatica (Led.)
stat. n., P. kamtschatica Led., Mem. Ac. Imp. Sc. St. Pet.
5: 557. 1815 is a generally coarser plant, the coarse
black hairs + 2 mm long, and the involucre 12-14 mm high.

63. STEPHANOMERIA Nutt.

Pappus bristles plumose, otherwise quite similar to
Lygodesmia and perhaps better united with it. Barbs of
the pappus 0.5-1.0 mm long.

1. S. cuncinata Nutt. (S. tenuifolia AA.) -- Much
like the common Lygodesmia juncea, but a bit more leafy
and the main stem leaves runcinate=pinnatifid. Pappus
pure white. Early to mid summer. Rolling steppes and
badlands, rare. --swS-swAlta, nwUS.

S. tenuifolia (Torrey) Hall occurs in Canada only
in British Columbia. Its leaves are filiform and entire
or merely denticulate. All specimens so named from our
area turned out to have the broader and more deeply cut
leaves of S. runcinata.

S. minor (Hooker) Nutt. was also reported by Daw-
son 1875 and Macoun 1884, but the justifying collection
from south of Wood Mountain (DAO, TRT) has since been re-=
vised to S. runcinata. See the Blue Jay 23: 41-42, March
1965.

64. TRAGOPOGON L. GOAT'S BEARD

Barbs of the pappus bristles very long and crinky,
becoming entangled at tip with the barbs of the next
bristle. Seed rather large, long beaked, its bristles
spreading horizontally into a conspicuous little parachu-
te.

a. Flowers purplish red ...eseeecccooe Lo To porrifolius
aa. Yellow; seeds with shorter beak.
b. Peduncle gradually thickened upward,
becoming about twice thicker near the
SUMMA Li 6, < ire «0.00 neve ele obieimis Gmjaiwie ss stay 7 (op eS
bb. Peduncle of uniform thickness... 3. IT. pratensis

1. T. PORRIFOLIUS L. -- Salsify, Oyster-Plant
(Salsifis) -- Largely similar to the next two. Ligules
purplish red, drying dark purple blue. Heads larger, up
to 8 cm in flower or fruit. Peduncle + enlarge upward=
ly. Involucre 3.5-5.5 cm high, at least in fruit. Seed
2.5-4.0 cm long, excluding the pappus, but including the
beak which is longer than the body. Pappus very light
brown, nearly concolourous with the achene. Late spring
to mid summer. Rare weed of rights of way. --(NS), Q=Man,
Alta-BC, US, (Eur).

STEPHANOMERTIA 204

1972 Boivin, Flora of Prairie Provinces 121

2. TT. DUBIUS Scop. (T. major Jacq.) -- Flowers
yellow like the next, but the peduncle gradually enlar-
ged upward to 4-8 mm across. Leaves not so strongly
falcate. Involucre 3-6 cm high. Seed 2.5-3.5 cm long,
the beak shorter than the body. Pappus whitish, lightly
tinted gray. Summer. Frequent weed, even invading na-
tive prairies in places. --Mack, NS, Q-BC, US, (Bur).

3. T. PRATENSIS L. -=- Goat's Beard, Jack-Go-to-
Bed-at-Noon (Salsifis blanc, Barbe de bouc) -- Leaves
longest, attenuate; tegules relatively longest; parachu-
te-seed largest. Leaves grass-like, falcate-recurved,
the long attenuate tip longer than the + lanceolate base.
Peduncle elongate, about 2 mm thick. Involucre 2-4 cm
high, at least equalling the florets. Fruiting heads +
5 cm wide. Achene 1.5=-2.5 cm long, the beak shorter than
the body. Summer. Rare weed of disturbed soils: North
Kildonan, Calgary. --NS-Man, Alta=-BC, US, Eur.

65. TARAXACUM L.

Scapose herb with a rosette of runcinate leaves and
a globose head of umbrella-like seeds. Ribs of the seed
covered towards the summit with short acicules. Pappus
bristles minutely scabrous.

Both in Europe and in America, the species concept
in this genus has been miniaturized. About 1000 micros=
pecies were described and named during the 1940-65 period
alone. In our 1962 survey of literature and major herba-
ria, we found the Taraxaca of Canada, Greenland and Alaska
filed under 112 different specific names. No overall
treatment exists for our area and none seems forthcoming.
The effective recognition of these finer segregates is
practically restricted to a few skilled botanists with
access to the specialized literature of the genus and a
good collection for comparison. We have found the reco-
gnition of the segragates to be a very fascinating and
sometimes frustrating herbarium exercise. But the intel-
lectual import of the exercise has eluded us.

As far as our experience goes, three names will ac-
count satisfactorily for the variation to be encountered
in our area, all the native plants being versed into T.

ceratophorum.

a. Tegules ascending or appressed ....3. IT. ceratophorum
aa. The outer ones strongly squarrose and
reflexed.
b. Achene stramineous to brown; leaves more
shallowly lobed upward ...cecscooe le To. officinale
bb. Achene becoming reddish brown at maturity;
leaves uniformly and more deeply lobed ..
Cece ec cccccc oc ccceccccccccecceos oe Ile laevigatum

205 TRAGOPOGON

122 PET OL0°G TA Vol. 23, no. 1

1. T. OFFICINALE Weber -- Dandelion, Faceclock
(Pissenlit) -- Seeds umbrella-like in a globose head.
Scapose perennial with abundant milky juice, a rosette
of runcinate leaves and monocephalous scapes. Leaves
more deeply and more narrowly lobed towards the base,
the upper lobes shorter and broader, the terminal one by
far the largest. Beak 0.8=1.5 cm long. Mid spring to
frost, mainly late spring. Common weed of lawns and
tramped or grassy places. -=-Mack, L-NF, NS-BC, (US, Eur).

2. T. LAEVIGATUM (W.) DC. (I. erythrospermum
Andrz.) -- Seed red brown, the base of the beak also red
brown. Resembles the above species. Leaves pinnatifid
to pinnatipartite, the lobes rather narrow and fairly
uniform in size, the terminal one not particularly lar-
ger. Beak 0.4=-1.0 cm long. Second half of spring. In-=
frequent weed of shaded or tramped places. =-=Mack, NS,
NB-BC, (US, Eur).

3. I. ceratophorum,(Led.) DC. (I. dumetorum Gree-
ne; T. eriophorum Rydb.; T. lacerum Greene; T. lapponi-
cum Kihlm.; T. lyratum (Led.) DC.) -- Like the above two
but the tegules neither squarrose nor reflexed. Leaves
variable. Tegule tip varying from flat and acute to ir=
regularly shaped or verrucose or bullate or corniculate.
Early to mid summer. Native in semi-open ground of open
places. --(G)-F-K, (Aka), L-(NF), Q-BC.

66. SONCHUS L.

Achenes strongly flattened. Pappus sessile, of
smooth capillary bristles. Stem leafy, the leaves auricu-
late-clasping, acicular-toothed.

a. Terminal leaf lobe deltoid; auricles triangular,
ACUTE crcccccccccarcccecesecccscsescs Se Se Dleraceus
aa. Terminal lobe ovate to lanceolate; auricles broa-
dly rounded.
b. Perennial; leaves mostly borne near the base
Of the Stem cecccusscescornaseces Le Se ArVenoie
bb. Annual; stem fairly uniformly leafy ....
were ccc c cere e neces cccccereccccesacceaa So Soe ASper

1. S. ARVENSIS L. var. ARVENSIS -- Sow-Thistle
(Chaudron=jaune, Créve-z-yeux) -- A coarse herb with spi-
nulose-margined leaves and yellow heads 3-5 cm wide in
flower. Perennial from deeply buried rhizomes. Leaves
mostly borne near the base of the stem, runcinate-lobed,
the middle and upper unlobed, much smaller and much more
remote. Inflorescence glandular hispid. Involucre 1.5
cm high or more. Summer, Common weed of cultivated
ground and wettish places. -=Aka, NF-SPM, NS-BC (US, Eur)
-- Var. GLABRESCENS C., G. & W. (S. uliginosus Bieb.) --
Inflorescence glabrous or at least not glandular, but only

TARAXACUM 206

1972 Boivin, Flora of Prairie Provinces 123

finely tomentose in places. --Mack, NS-BC, (US, Eur).

Despite a difference in chromosome number (54 for
arvensis and 36 for glabrescens) our two variants fall
short of the minimum morphological differentiation to
justify specific rank. The intervarietal hybrid, named X
var. Shumovichii Boivin, has been found in Ontario and
is likely to turn up in our area; it has intermediate
vestiture and chromosome count, it backcrosses readily
with the parental types to produce a sliding scale of
chromosome counts and pubescence density.

2. S. OLERACEUS L. --Milk-Thistle (Chardon blanc,
Laiteron) -- Terminal leaf lobe deltoid, about as wide
as long. Stem about evenly leafy, the middle leaves
often largest, mostly pinnatifid, the margin not quite
so sharply acicular as the first, the basal auricles nar-
rowed to acute tips. Annual. Involucre about 1 cm high.
Achenes finely rugulose, the rugosities in transverse
rows and about as abvious as the weak longitudinal nerves.
Mid summer to early fall. A weed, mainly of backyards,
waste places and gardens. --Mack, (Aka), NF-SPM, NS-BC,
US, Eur.

There are many reports for Saskatchewan, but their
basis remains largely obscure. A Saskatoon (SASK) col-
lection in 1917 originally identified as S. arvensis var.
glabrescens proved to be our first definite sheet of S.
oleraceus for the province. Also reported as a greenhou-
se weed at Regina. The Langham report (SASKP) has been
revised to S. asper. Other reports could not be substan-
tiated, but a recent collection by Hudson at Saskatoon
(DAO) is confirmed herewith.

3. S. ASPER (L.) Hill -- (Chaudronet) -- Resembles
S. arvensis, but annual and the involucre only 1.0-1.4
cm high. Leaves more evenly spaced, although the upper
are smaller and the internodes longer. Auricles strongly
recurved, almost spirally coiled. Achene with 3 nerves on
each side, otherwise smooth or nearly so. Mid summer to
early fall. Weed of waste places and beaches. --Y~-Aka,
L=-(NF)-SPM, NS-BC, US, Eur.

67. LACTUCA Le LETTUCE

Differs from Sonchus by the beak (or top) of the
achene being dilated into a disk on which the pappus is
borne. Habitally quite similar to Sonchus.

a. Perennial with large blue heads ..... 5. L. tatarica
aa. Annual or biennial with narrow heads. a
b. Leaves acicular dorsally along the midnerve.
ce Involucre 10-13 mm high ..... 1. L. Serriola
cc. Larger, 17-23 mm high ..e.. 4. Lo ludoviciana

207 SONCHUS

12h Prk 2) 0) ElOnG TA Vol. 23, no. 1

bb. Midnerve smooth.
d. Pappus dirty gray to brown..e.e. 7o Le biennis
dd. White.
e. Leaves, and especially the bracts,
broadly cordate clasping at base a.
Bleteleltels slieleteiene o's Gisleitela eletelere eieie (2 eta emmemet eves
ee. Bracts, and also usually the leaves,
winged=-petiolate or tapered at base.

f. Pappus on a long and thin beak;
panicle narrow, crowded, + lan-
ceolate o...scae-eeo Sa Le Canadensis

ff. Pappus sessile; panicle ample,
broad and open ....e. 6 Lo floridana

1. L. SERRIOLA L. (f. integrifolia (Bogenh.) G.
Beck, var. integrata G.& G.; L. Scariola L.; L. virosa
AA.) -- Prickly Lettuce (Plante boussole, Escarole) -=
Leaves with a row of stiff acicules on the back of the
midnerve. Leaves lobed or not, spinulose=-margined, on
sunny days becoming twisted into a common vertical plane.
Inflorescence ample, heads very narrow, yellow, often
drying blue. Mid summer to early fall. Waste places, un-
common, --(NS)-PEI-BC, US, Bur, (Afr).

2. L. SATIVA L. -- Lettuce (Salade, Laitue) -- Lea-
ves broadly flabellate or obovate, Cordate=clasping at
base, passing into the numerous cordate bracts. Somewhat
spinulose=-toothed along the leaf margins but not along
the midnerve. Heads small, numerous, tending to become
corymbose, Late summer and early fall. Commonly culti-
vated; rarely and fleetingly spontaneous: Fort Saskatche=
wan. --O, Alta, (US, Eur).

3. J. canadensis L. (var. latifolia Kuntze, var.
longifolia (Mx.) Farw., var. montana Britton; L. integri-
folia Big.) -- Devil's Weed (Chicorée blanche) -- Very
variable, the leaves sometimes narrowly lanceolate, en=
tire and clasping at base, but typically they are pinna-
tifid with a winged petiole. Inflorescence leaves and
bracts attenuate at base. Involucre 10-15 mm high. Pap-=
pus white, borne on a filiform beak, Early summer. Dry,
open places. --NS-seMan, seBC, US.

Reported by Groh 1950 for Eastend, Sask. The jus-—
tifying specimen (DAO) has since been revised to L. tata-
rica var. heterophylla. The source of the Saskatchewan
reports by Fernald 1950, Gleason 1952 and Scoggan 1957 is
still to traced. These may rest on specimens, such as
those of Bourgeau, with outdated or vague geographical
documentation. A listing by Russell 1937, 1944 was mere-
ly speculative. An Alberta report by Rydberg 1932 has not
been investigated.

4. L. ludevicjana (Nutt.) Riddell -- Like the last,
but the leaves spinulose dorsally along the midnerve and

LACTUCA 208

1972 Boivin, Flora of Prairie Provinces 125

the heads longer. Inflorescence an open panicle. Invo-
lucre 15-23 mm high. Mid summer. In the shrubby zone
around bluffs. --(O-Man)-seS, US.

S. J. tatarica (L.) C.A. Meyer var. heterophylla
(Nutt.) Boivin (var. pulchella (Pursh) Breitung; L. pul-
chella (Pursh) DC.) -- Blue Lettuce -- Especially conspi-
cuous along roadsides, a virgate herb with large blue
heads of ligulate flowers. Leaves narrowly lanceolate,
entire, or the lower remotely lobed. Heads 2-3 cm wide.
Midsummer. Scattered on the prairie, becoming conspi-
cuous when the soil es disturbed. --seK~Mack~(Y) -Aka,
Q-BC, US.

Stat. n., Mulgedium heterophyllum Nutt., Trans,

Am. Phil. Soc. Z, 7: 441. 1841; Lactuca pulchella
(Pursh) D.C. var. heterophylla (Nutt.) Farw., Ann. Rep.
Mich, Ac. Sc. 6: 214, 1904. The latter combination es-
tablishes the priority of heterophylla at varietal rank.

Our plant is weakly differentiated from the sibe-
rian var. tatarica in which the leaves bear smaller, more
remote and somewhat spinulose teeth.

A white-flowered form is known from Minnesota and
probably occurs in our area, merely awaiting a sharp-eyed
collector. It may be designated as f. Steyvensii f.n.,
floribus albis. Typus: O.A. Stevens 2514, Felton, Minn.
Aug. 10, 1961 (DAO). ARPA SANRL TS hE

6. L. flocidana (L.) Gaertner -- Resembles L. cana-
densis, but the inflorescence is more open and the achene
is beakless, the pappus sessile. Leaves pinnatifid to pin-
natipartite, the terminal lobe broadly deltoid. Flowers
blue. Mid summer. Edge of woods, rare: Otterburne.--
sO-seMan, US.

7. Le biennig (Moench) Fern. (L. spicata AA.) --
Pappus tinted grayish to pale brown, otherwise similar to
L. canadensis. Main leaves pinnatipartite, the upper ones
narrow, entire auriculate-clasping at base. Involucre +
1 cm high. Mid to late summer. Low and wet places. --
(Aka), L-NF-(SPM), NS-BC, (US).

Closely related to L. floridana and perhaps only va-
rietally distinct.

68. LYGODESMIA D. Don
Ligules pink, the heads very narrow and containing
only about 5 florets. Achene tapered at beak. Pappus of
smooth bristles. Tegules dimegueth, the outer ones many
times shorter.
@. Perennial; involucre 1.0-1.5 cm high .. 1. L. juncea
aa. Annual; involucre 1.5-2.0 cm high .... 2. L. rostrata

1. L. juncea (Pursh) D. Don -=- Skeleton-Weed -- A
skeleton weed with pink ligules. Rhizomes deeply buried.
Very branchy from near the base. Branches longitudinally

209 LYGODESMIA

126 PLY TD Oj iGag: EA Vol. 23, no. 1

striate. Leaves many but narrow, small and appressed,
not very conspicuous and mostly shorter than the interno-
des. Heads terminal, solitary. Mid summer. A common
prairie and steppe species. --sMan-sBC, US.

2. L. rostrata Gray -- Leaves and heads longer
than the last. Blending in its surroundings and very hard
to see. Leaves very narrow and very long, 2-6 times lon-
ger than the internodes. Branching mainly near the top.
Heads terminal and axillary. Late summer. Bare or semi-
bare sands, mainly in blowouts. --swMan-sAlta, cUS.

69, AGOSERIS Raf.

A basic type, scapose with monocephalous scapes
and a pappus of non=-plumose bristles on a beaked achene.
Bristles minutely scabrous. Tegules imbricated. Resem-=
bles Taraxacum, but the achene is not acicular-muricate
towards the top.

a. Beak 2-4 times as long as the body of the achene;
tegules strongly dimorphicec..-coee 3. Ao grandiflora
aa, Beak shorter; outer tegules shorter, but otherwise
similar to the inner.
b. Achene with beak half as long as the body or
less; ligules yellow, drying yellow..1. A. glauca
bb. Beak longer; ligules deep orange, often drying
PUrpLISh” sco ce cece ee cc owocccsc! Se) Ne Ube mnt eee

1. A. glau (Pursh) ref. var. glauca (var. agres-
tis (Osterh.) a Jones, var. dasycephala AA., var. parvi-

flora (Nutt.) Rydb.; A. agrestis Osterh.; A. parviflora
(Nutt.) Dietr.; A. scorzonerifolia AA.; A. turbinata Rydb.)
-- A native resembling the common, weedy Dandelions, but
the leaves entire to narrowly lobed and the tegules not
squarrose, Also resembles Microseris, but the latter has
tomentose-ciliate leaves. Herbage entirely glabrous or
quite often tomentose towards the summit of the scape and
on the involucre, more rarely pilose on the stem and/or
the leaves irregularly retrorse=-ciliate towards the base,
but glabrous on both faces, yet the midnerve sometimes pi-
lose. Early to mid summer. Common in prairies. --O-BC,
US--Var. dasycephala (T.& G.) Jepson (A. scorzonerifolia
(Schrader) Greene -- Herbage pubescent throughout, beco=
ming lanate on the involucre and towards the summit of the
stem. Leaves pubescent on both faces, at least towards
the edges. -- Cypress Hills, Grande-Prairie and Rockies.
--Mack, Alta-BC, US.

A range extension to Yukon by Anderson 1949, repea-=
ted by Hultén 1950, querried by Boivin 1967, was based on
a British Columbia collection from mile 611 along the A-
laska Highway. See under Aster conspicuus above.

2. Ae aurantiaca (Hooker) Greene var. aurantiaca
-- Closely similar to the first, the top of the peduncle den-

AGOSERIS 210

1972 Boivin, Flora of Prairie Provinces 127

sely lanate. Beak of the achene thin, up to twice as
long as the body. Leaves more commonly dentate to remo-
tely lobed. Mid summer. Alpine and sub-alpine prai-
ries. --Mack-Y-(Aka), Q, swAlta-BC, wUS,.

In @ more southern var. purpurea (Gray) Cronq.
the tegules are larger, more strongly imbricated and
purple-spotted,

3. A, grandiflora (Nutt.) Greene -- Herbage light-
ly villous below to more densely so above, sometimes gla-
brescent. Leaves entire to more commonly pinnatifid. Ou-
ter tegules rhomboid-ovate, acuminate, broader than the
inner. Ligules yellow, drying yellow or sometimes pur-
plish. Early summer. Lowland meadows. Lake Saskatoon.
--wcAlta-swBC, wUS.

70. CREPIS L.
Much resembling Agoseris, but with the stem more or
less leafy and the inflorescence of more than one head.
Achene beakless or short-beaked.

a. Annual or biennial weed.
b. Leaves mostly cauline ..ceccaseeee Se Co tectorum
bb. Rosette leaves more numerous cee Oe Cao “capillaris
€@a. Perennial natives with a strong taproot;
leaves mostly in a basal rosette.
c. Depressed and nearly stemless plant
with the basal leaves overtopping the
NEAdS cecccccecccccacerevescccesescoce “eo Co nana
cc. Stem well developed and the heads much
overtopping the foliage.
d. Glabrous; stem branchy and leafy
throughout eeoesseccoeerecseeee 1. C. elegans
dd. At least the rosette tomentose or
hirsute; stem leaves, bracts and
heads mostly borne in the upper
third of the plant.
e. Plant glabrous above, the roset-
te coarsely hirsute ..... 3. C. runcinata
ee. Herbage lightly to densely tomen-
tose throughout ecceseo 4. C. occidentalis

1. €. elegans Hooker -- Glabrous throughout, some-
wheat glaucous and slightly fleshy. Tufted and very bran-
chy, 1-2 dm high. Lower and basal leaves petiolate, lan-
ceolate, entire or nearly so, the upper linear and sessi-
le. Heads numerous, 6-8 mm high. Summer. Gravel flats
of braided glacial streams, --Mack-Aka, wAlta-eBC, (nwUS).

2. C. nang Rich. var. nana (ssp. ramosa Babcock)
Like a dwarf version of the last, the stem so short that
the basal leaves usually overtop the inflorescence. Ta-
proot long and thick, rather large in relation to the

aboveground parts. Involucre dark green, 9-13 mm high.
211 CREPIS

128 Paboyerl OpLsOnG Lea Vol. 23, no. 1

Mid summer. Alpine shale slides, --F-(K)-Mack-Aka, L-NF,
swAlta-BC, wUS, (eBur).

Occasional specimens have a more elongated stem
(=var. elongata). These are readily distinguished from
C. elegans by the size of the heads.

In the alaskan and east-asiatic var. lyratifolia
(Turcz.) Hultén the leaves are + pinnatifid.

3. C. runcinata (James) T.& G. var. pauncinata
(C. glaucella Rydh;C.perplexans Rydb.) -- Very much like
a Taraxacum, the leaves + runcinate and all basal, but
the stem bearing more than one head and bracteolate in
the inflorescence. Rosette leaves sometimes entire,
coarsely hispid, not glandular. Tegules glandular-his-=-
pid, the glands yellow; sometimes also finely puberulent.
Early to mid summer. Common in wet places; alkali tole=
rant. --sMan-BC, US -- Var. glauca (Nuttall) Boivin (C.
glauca (Nutt.) T.& G.) -=- Involucre not glandular, merely
finely puberulent, or more commonly glabrous. Mainly on
alkaline prairies and shores of playas. --(sMan)-S-Alta,
US -- Var. hispidulosa Howell (C. platyphylla Greene) --
Larger and the leaves usually glandular along the midner-=
ve. Pubescence otherwise as in var. runcinata. Leaves
ovate to lanceolate, usually larger, 1 dm long or more,

4 cm wide or more. Heads numerous, usually more than 10.
Grassy highlands: Cypress, Waterton. --swS-Alta, wUS.

The range had been extended eastward to Timmins,
Ontario, in Nat. Mus. Can. Bull, 156: 246, 1958, but the
justifying collection (MT, TRT) has since been revised
to Leontondon autumnalis L. var. pratensis(Link)W.D.J.Koch,.

4. C; oggidentalis Nutt. var. ogcidentalis (var.
costata Gray; C. atribarba Heller; C. intermedia Gray) --
Leaves deeply lobed, pinnatifid to pinnatipartite, the
lobes triangular to filiform and entire to dentate. In-
florescence corymbose. Heads few to many, usually gra-
yish-tomentose, commonly 1.0-1.5 cm high, bearded with
thick, black hairs about 0.5 mm long. Early summer.
Montane prairies and Milk River Valley. --swS-sBC, US.

More western plants in which the involucre is devoid
of thick black hairs are distinguished as var. cytotaxo-
nomicorum (Boivin) stat. n., C. atribarba Heller var.
cytotaxonomicorum Boivin, Nat. Can. 87: 31. 1960 (and ul-
timately ssp. originalis Babc. & Stebb.)

This species varies within unusually broad limits;
the more obvious phenotypic variant has the leaves pec-=
tinately dissected into remote, narrow and usually entire
segments; it has been called C. atribarba, but it is not
discretely separable from the run-of-the-mill C. occiden-
talis and the rank of variety (i.e. var. gracilis D.C.
Baton) would be more realistic. We have not given it re=
cognition at any rank because, at least in the canadian
part of its range, it seems to present itself as an ex-=

CREPIS 212

1972 Boivin, Flora of Prairie Provinces 129

treme of variation of sporadic occurrence.

Canadian reports of C. acuminata Nutt. and C. an-
gustata Rydb. were based, at least in part, on speci-
mens of var. cytotaxonomicorum. This remark may possi-~
bly apply also to some of the earlier reports of C. in-
termedia discussed below.

C. modocensis Greene, & more southern species, ap-
pears to grade into C. occidentalis on the one hand and
also into the more southern C. acuminata Nutt. on the
other. Neither C. modocencis var. C. acuminata are known
in Canada, but intermediates to C. occidentalis do occur.
It is customary to use C. intermedia Gray to designate
such intermediate specimens. We are not too clear as
to their significance; they do not appear to be hybrids,
yet we are not able at present to offer a classification
that would reflect taxonomically and account realistical-
ly for the existence of such extraneously related inter-
mediates in our flora. Over a period of years we have
tried now to consolidate the members of this series into
a single species (C. occidentalis), now to treat them as
so many species, but we have not been able to achieve a
satisfactory treatment either way.

Se C. TECTORUM L. -- Tegules pubescent on both
faces, being strigose on the inner. Annual (or biennial)
and very variable. Stem simple, becoming very branchy.
Rosette leaves evanescent, being usually wilted by flowe-
ring time. Stem leaves numerous, lanceolate to long li-
near or filiform, entire to pinnatifid. Involucre 7-9 mm
high, tomentose and glandular=pubescent. Seeds 2.5-4.5 mm
long, with a short thin beak and a white pappus. Summer.
Common weed of roadsides, railways, etc. --G, sMack-Y,
(NF, NS)-PBI-BC, US, Eur, (Oc).

6. C. CAPILLARIS (L.) Wallr. -- Somewhat like the
last but biennial and retaining its abundant rosette all
summer. Branchy and often many-stemmed, the lowermost in-
ternode(s) usually quite short. Stem leaves mostly sub-
tending branches and smaller than the rosette leaves, the
latter mostly 1-2 dm long. Tegules pubescent on the outer
face only, or glabrous on both faces. Heads small, the
involucre only 5-8 mm high. Seeds beakless and only 1.5-
2.5 mm long. (Late summer?). Rare weed of drier and open
places: Calgary.--NS, NB-O, swAlta-BC, US, (SA), Bur,
(Oc).

Our only collection (MIMG) is undated. It was made
by M.E. Moodie about half a century ago.

71. PRENANTHES L. RATTLESNAKE-ROOT

A middling type with a leafy stem and beakless seeds
bearing a white and smooth pappus. But the flowers are
nearly white or pale pink and the inflorescence is race-

213 CREPIS

130 PHY ‘T O'L-0'C FA Vol. 23, now. 1

mose or paniculate. Habitally often similar to Lactuca,
but the seeds not flattened.

a. Inflorescence abundantly hirsute .oeee le Pe racemosa
aa. Glabrous.
be Tegules green; pappus lightly tinged ...
wi big), & elisy'ni eosi'ape oh oVe) alain Sal we Gia mie wish ele)s\e | im kt inane ees
bb. Tegules purple; pappus chestnut brown .o

e@eoeerreeroeeseoe eer eoeeoeeeeeeeee79228289900 oye P. alba

1. BP. gacemosa Mx. (Nabalus racemosus (Mx.) DC.)=-=
Copiously hirsute in the inflorescence, glabrous and
glaucous below. Lower leaves oblanceolate, petiolate, the
upper ones much smaller, sessile and cordate-clasping.
Involucre purple. Ligule pink to nearly white. Pappus
yellow. Late summer. Wettish prairies, infrequent. --
(seK), NF, NS, NB=-neBC, US.

2. PB. sagittata (Gray) Nelson -- Leaves sagittate,
remotely dentate, the upper successively rhomboid then lan-
ceolate. Lower leaves often opposite. Petiole winged.
Inflorescence narrow. Ligules white. Pappus straw-colou-
red. Mid summer. Mountain woods: Rockies and Swan Hills.
--Alta, (nwUS).

3. P. alba L. (Nabalus albus (L.) Hooker) -- Rattle-
snake-Root -- Main leaves deltoid, remotely dentate to
deeply lobed. Lower petioles not winged. Pedicels very
short. Ligules white. Pappus deep brown. Second half of
summer. Low woods. --Q-cS, (US).

72, HIERACIUM L. HAWKWEED

Like Prenanthes, but the flowers typically yellow
and the inflorescence commonly umbellate.

a. Leaves mostly basal.
b. Leaves glabrous or nearly 50 .-eacooe 3o H. triste
bb. Copiously long hirsute.
c. Flowers orange-red ...-cecoo. 1. H. aurantiacum
CCo WHITE coeeerccceceeeceecaces Co H. albiflorum
aa. Leaves all or mostly borne on the stem.
d. Leaves fairly uniformly distributed on the
stem, the lower ones wilted or deciduous
by flowering time ...-eece..-.ee0 Ze H. umbellatum
dd. Leaves borne mostly in the lower third of
the stem, the others few and much smaller ...
wee cere weer ese cescaceececes 4. H. cynoglossoides
1. H. AURANTIACUM L. == Devil's Paint-Brush, King-
Devil (Marguerite rouge, Saint-Louis) -- Heads red-orange,
tending to dry deep red. Herbage copiously very long hir-=
sute throughout and more or less purplish. Involucre den-
sely pubescent with a mixture of long hirsute hairs, shor-
ter glandular ones and very small stellate hairs. Mid
summer. Rare and recent roadside introduction. --NF-

PRENANTHES 214

1972 Boivin, Flora of Prairie Provinces 131

(SPM), NS-O, Alta-BC, US, Eur.

Has been repeatedly reported for Manitoba by Lowe
1943, Frankton 1955, 1970, Scoggan 1957, Budd 1957 and
1964, Best 1964, and Boivin 1966 on the basis of a Winni-
peg collection. Yet in 1966 no such collection could be
located at WIN or elsewhere and we are now speculating
that, if any specimen ever existed, it may have been re-
vised to some other taxon.

2. H. umbellatum L. (var. canadense (Mx.) Brei-
tung; H. canadense Mx.; H. columbianum Rydb.; H. scabri-
usculum Schwein.) --(Accipitrine) -- Leaves typically re-
motely dentate. Pubescence variable. No basal rosette
and the lower leaves early deciduous. Leaves broadest
below the middle. Heads often subumbellate. Mid and
late summer. --Mack-(Y-Aka), L-NF-(SPM), NS-Alta-(BC),
US, Eur.

A somewhat variable species, found on both sides of
the Atlantic. Mainly on the basis of pubescence varia-
bility, the Canadian material has been subdivided into
about 20 taxa. In Europe, where the genus is dealt with
on the basis of national monographs, this species has
been further subdivided into a host of microspecies.

3. H. triste W. var. gracile (Hooker) Gray--Smal-
lest and green below, but conspicuously black=pubescent
in the inflorescence. Commonly less than 3 dm high. Her-
bage glabrous or nearly so below, becoming densely long
pubescent in black in the inflorescence, the pubescence
mixed with much shorter and partly glandular hairs. Youn-
ger plants are sometimes finely pubescent to the base.
Leaves less than 1 dm long, oblanceolate, rounded at tip.
Involucre 6-8 mm high. Mid summer. Meadows towards tim-
berline. --Mack-Aka, wAlta-BC, wUS, (SA).

In the more western typical phase the longer hairs
reach 2-4-(5) mm and are usually not glandular. In our
variety they are only O0.5-1.5 mm long and often partly
glandular.

4. H. cynoglossoides A.-T. (H. albertinum Farr;

H. Rydbergii Zahn; H. Scouleri AA.) -- General habit of
H. a@urantiacum and H. albiflorum, but the ligules yellow
and the leaves mostly borne towards the base of the stem.
Leaves and lower part of plant densely long-hirsute, the
upper part variously hirsute or glandular or stellate-pu-
bescent. Larger leaves mostly 1-2 dm long. Involucre
mostly 8-10 mm high. Mid summer. Montane prairies and
light woods; Cypress and Rockies. --Alta-sBC, nwUS.

H. Scouleri is listed by Porsild 1959 for the Roc-
kies, but all specimens so named at CAN,GH and V have been
revised to H. cynoglossoides.

Se. H. albifloryn Hooker -- Ligules white and the
herbage devoid of stellate pubescence. Otherwise quite

215 HI BRACI UM

132 Pon YT OFL{0%G TA VoL. 23, nese

similar to H. aurantiacum but the long hairs very dense
towards the base, becoming very sparse in the inflores-
cence. First half of summer. Lodgepole forests. -=-
(Y)-Aka, (seMan)-swS-BC, wUS.

HI ERACIUM 216

1972 Boivin, Flora of Prairie Provinces 133
ADDITIONS AND CORRECTIONS

The following information became available only
after the corresponding text had be given its final form
for printing.

Page 9 »- Cuscuta Gronovii W. -- It must be reco-
gnized that the U.S. material at hand is much more varia-
ble than ours. Some of the U.S. specimens examined do
have much longer and lanceolate corolla lobes, other spe-
cimens do have much smaller capsules loosely enclosed by
the marcescent corolla, etc. Therefore we are not pre-
cluding that some of the distinctions rejected for our
Canadian material could be applicable and pertinent to
more southern populations.

Page 15 -- Penstemon albidus Nutt. -- Dawson's 1875
report of P. glaucus Graham from the Second Crossing of
the Souris River (DAO) was ignored by Macoun 1884, but
referred to P. gracilis by Scoggan 1957. It proved to be
based on @ specimen of P. albidus.

Page 17 -- Penstemon gracilis Nutt. -- The stem is
minutely retrorse-puberulent, at least towards the base.

Page 19 -- Limosella -- The leaf width is not a
fully reliable distinction between L. aquatica and L. su-
bulata, but these may be further contrasted as follows:

L. aquatica L. -- Stoloniferous, the stolons green,
superficial, and usually present in the herbarium. Flo-
wer 1.0-2.0 -(2.5) mm long, the corolla only slightly
(or not at all) exserted; typically only the lobes are
exserted. Ripening capsule with a white line along the
suture, eventually opening by two valves, these finely
pencil-margined in white.

The last character is transposed in Fernald 1950.

L. subulata Ives -- Stolons thinner, white, sligh-
tly buried, fragile and usually absent in the herbarium,
Flower larger, 2.5-4.0 mm long, the corolla being 1 ti-
mes the length of the calyx; typically the corolla lobes
are fully exserted along with part of the tube. Capsule
not lined in white, rupturing irregularly at maturity.

The reports of L. subulata for our area by Boivin
1967 and above are to be discounted as they were based
on collections from Granum and Ponoka (both DAO) with
filiform leaves but with the smaller flowers, etc., of
L. aquatica and they have been revised accordingly. Re-
ports from Alberni, B.C. and Keewatin are also to be
referred to L. aquatica on similar grounds. L. subulata
is then apparently restricted in its distribution to the

tidal shores of the east coast of North America.
217 ADDITIONS

134 PsHVPtT OGLZOtG Tek Vol. 23, no. 1

Page 20 -- Veronica longifolia L. -- Mostly 5-10
dm high. Leaves all opposite, or the uppermost often
alternate or verticillate. Calyx green, much less den-
sely puberulent than the rest of the inflorescence, the
lobes finely ciliate and lightly puberulent on back.
Style 7-9 mm long after the fall of the corolla.

The above criteria will bring out the differences

with V. spicata.
Page 20 -- Add the following species:

la. VERONICA SPICATA L. -- (Perse brunette) -=
Quite similar to V. longifolia but generally smaller,
mostly 2-4 dm high. More densely puberulent, becoming
grayish in the inflorescence. All leaves opposite. Ca-
lyx as densely puberulent as the rest of the inflorescen-
ce and + grayish. Style 4-6 mm long after the fall of
the corolla. Infrequent ornamental, rarely spreading to
roadsides; Lacombe. --Q, Alta, (US), Bur.

Page 23 -- Agalinis purpurea and its variety par-
viflora should be eliminated from our area.

First reported by Hooker 1838 as Gerardia purpurea
from Saskatchewan where collected by Drummond. This was
repeated by many later authors, but the use of Saskatche-
wan in Hooker does not coincide with the modern meaning
of Saskatchewan as a province. Reports in the latter sen-
se were justifiably discounted by Breitung 1957. Indeed
the label of Drummond's specimen merely reads ''Norway
House to Canada" (K). Pennell 1935 studied this sheet,
and cited it under Gerardia paupercula borealis as coming
from "Manitoba (?)'". However Drummond's geographical data
was essentially vague and in the absence of a later con-=
firmation it seems unwarranted to assume that his speci-
men was actually collected in Manitoba rather than fur-
ther to the east.

A report from Morden by Lowe 1943 was discounted by
Scoggan 1957 as being based on a specimen of G. aspera.
Similarly the collection Garton 3537, Stony Mountain
(DAO) cited by Scoggan 1957 as G. paupercula has since
been revised to Agalinis aspera. A Dawson collection
from Lake of the Woods (MTMG) was listed by Dawson 1875
as G. purpurea and by Scoggan 1957 as G. paupercula. It
belongs with A. tenuifolia var. parviflora as do all col-
lections studied from the southeastern part of the pro-
vince.

Thus we are left without any unquestioned voucher

to the presence of A. purpurea (or G. paupercula) in Ma-

nitoba,.

Page 24 -- Agalinis tenuifolia (Vahl) Raf. -- Axil-
lary fascicles are usually present in our var. parviflora

ADDITIONS 218

1972 Boivin, Flora of Prairie Provinces 135

(Nutt.) Pennell, and the capsule is 5-7 long. In the more
eastern and less boreal typical phase, var. tenuifolia,
axillary fascicles are usually lacking, the calyx lobes

do not exceed 1 mm, and the smaller capsule is only 3-5 mm
long.

Page 25 -- Castilleja lutescens (Greenman) Rydb. --
Also in Waterton and the Pincher Creek areas. Older Al-
berta reports of C. pallida var. septentrionalis by Ma-
coun 1884 and others were primarily based on specimens of
C. occidentalis, but the Dawson collection seems closer
to C. lutescens.

Page 29 -- Pedicularis flammea L. -- Add: F. fle-
vesceng, Pol. -=- Corolla monochrome in yellow. Cadomin,.
-- (nQ)-nO, swAlta.

Page 29 -- Pedicularis Oederi var. albertae is to
be versed into the synonymy of P. flammea of which it is
only a@ more abundantly villous extreme.

The difference in pubescence between P. flammea and
P. Oederi is not sharply marked, despite some keys, in-
cluding ours above.

On the basis of the more abundant material now at
hand, the variation in pubescence runs as follows:

P. Oederi. Herbage more or less villous throughout
or at least in the inflorescence. But sometimes the her-
bage is completely glabrous except for the ciliate bracts
and calyx lobes.

P. flammea is typically glabrous except for the ci-
liate bracts and calyx lobes. Varies to completely gla-
brous and eciliate, or again to more or less villous in
the inflorescence.

The two species are obviously close to each other,
yet quite distinct, and may be contrasted as follows:

P. Oederi. Usually 1-2 dm high in flower, elonga-
ting to 2-3 dm in fruit, Flowers larger by half, (16)-
20-(25) mm long. Calyx 8-11 mm long, it lobes + dilated
and + toothed at tip. Corolla exserted from the calyx
by + lcm. Galea 2-3 mm wide and tinged or spotted in
red towards the tip. Lower corolla lobes spreading +
horizontally. Style exserted by (0.1)-0.5-1.0-(2.0) mm.

P. flammea is generally a smaller plant with smal-
ler and more deeply coloured flowers. Usually less than
1 dm high at flowering, elongating to 2 dm in fruit.
Flower (11)-14-(16) mm long. Calyx 6-9 mm long. Corol-
la exserted by 5-7 mm. Galea 1.5-2.0 mm wide, deep red
for about half of its length. Lower corolla lobes di-
vergent by 30-45°, Style not protruding from the hood

219 ADDITIONS

136 PLAY Op 0.G EA Vol. 23, no. 1

of the galea.

Page 33 -- Orobanche fasciculata Nutt. -- The range
extension to Ontario is based on a single plant recently
collected at La Cloche Island (OAC). For Anaplanthus read
Anoplanthus.

age 33 -- Orobanche uniflora L. -- On the basis of<
the length and shape of the calyx lobe this species is di-

visible into a pair of geographical variants as follows:

Var. uniflora -- Calyx lobes less than twice as long
as the tube, gradually narrowed into an attenuate tip
which is shorter than the main triangular portion of the
lobe. Cypress Hills. --(Y)-Aka, NS-(PEIL)-NB-O, swS, US.

Var. minuta (Suksd.) Beck -- Calyx lobes longer and
more abruptly narrowed, at least twice as long as the tu-
be, the main portion of the lobe tending to be deltoid and
shorter than the caudate tip. Waterton. --soAlta-BC,
wUS.

More than 90% of the specimens will conform well to
the above criteria relative to their geographical origin,
but quite a few are transitional and the odd one (igno-
red for the purpose of the above ranges) will be comple-
tely atypical.

A further subdivision of the western phase is so-=
metimes attempted in which var. minuta (=var. Sedi(Suksd,)
Achey) is restricted to the smaller-flowered plants, whi-
le var. purpurea (Heller) Achey denotes the rather showy
larger-flowered plants. Both varieties appear to have
the same range; the rank of form would probably be more
appropriate, if the distinction is deemed desirable.

Page 38 -- Geranium pratense L. var. erianthum
(DC.) Boivin -- The flowers are typically bluish-mauve,
but a white-flowered mutant is known: f. leucanthum f.
ne, petalis albis. Typus: W.B. Schofield 2489, Alaska,
Cold Bay, flowers white, rare among typically blue-flo-
wered plants on sheltered tundra slope, July 28, 1952
(DAO). Still known from a single alaskan collection, but
expected to be sporadic throughout the range of the spe=
cies.

Page 56 -- Mertensia lanceolata (Pursh) A. DC. var.
lanceolata -- Our plant grows in closed tufts and its
stems bear (8)-10-(15) leaves. It is barely distinct
from the rare arctic var. Drummondii,(Lehm.) stat. n.,
Lithospermum Drummondii Lehman, Nov. Stirp. Pug. 2: 26.
1828; Mertensia Drummondii (Lehm.) G. Don. The latter
is more loosely tufted, the caudex branches more or less
elongated and abundantly clothed with stubs of old pe-
tioles, the stem leaves only 5-8-(10) to a stem. No con-

ADDITIONS 220

1972 Boivin, Flora of Prairie Provinces 137

sistent difference could be detected in the floral parts.

Page 70 -- An undetermined species of Mentha has
been recorded as persisting in a long abandoned garden at
Glenevis (Pegg 1213, DAO). By its ovate leaves, its long
and dense pilosity, etc. it resembles M. rotundifolia
(L.) Hudson, a species known to persist occasionally in
southern Ontario. However, our plant differs in a num-
ber of respects: its pubescence is longer in the inflo-
rescence, its spikes are leafy-bracted, the bracts being
large, its calyx is somewhat longer with longer pubes-
cence, etc.; we have yet to find a satisfactory name for
it.

Page 70 -=- For Elscholtzia read Elsholtzia.

Page 210 -- Agoseris glauca (Pursh) Raf. -- The
herbage will vary from completely glabrous to + lanate
on the involucre and at the summit of the stem. The pu-
bescent phase has been commonly identified and reported
upon as var. dasycephala or as A. scorzonerifolia. But
both forms are generally common, they have been reported
throughout our area and do not appear to be taxonomically
Significant. If however var. dasycephala be defined ina
somewhat more restrictive manner, as we have done above,
it does become a geographically restricted variety.

Because of our more restrictive definition of var.
dasycephala, it seems preferable to discount all previous
reports except such as were checked and conformed to our
criteria. All Manitoba sheets examined, including Mars-
hall's from Brandon (DAO), belonged to var. glauca. Ear-
lier Saskatchewan reports of A. scorzonerifolia were dis-
counted by Breitung 1957 and all specimens examined, in-
cluding Breitung 4442 (DAO), were placed with var. glauca.
Similarly most Alberta specimens were referred to var.
glauca. But all Mackenzie sheets examined were closer

to var. dasycephala.

The leaves vary from entire to dentate, or more ra-
rely pinnatifid with the lobes narrow and somewhat remote.
The last phenotype (=var. agrestis) seems to occur throu-
ghout our range and accordingly it has been submerged into
the typical phase.

Page 215 -- Hieracium triste W. var. triste -- Now
known to our area on the basis of the following: G. Scot-=

ter 16857 & 16950, Tonquin Valley, Jasper Park, 1971 ~ (DAO).

The general range: swMack-Aka, swAlta-BC, wUS Wares uae
(eBur).

221 ADDITIONS

138

PoB: Yo? OGiOG TA

Vol. 23, no. 1

INDEX OF GENERA IN PART III

Achillea, 165
Actinella, 164
Adenocaulon, 149
Adoxa, 5

Agalinis, 23, 218
Agastache, 62
Agoseris, 202, 210, 221
Allocarya, 52
Ambrosia, 150
Amsinckia, 53
Anaphalis, 147
Anaplanthus, 33, 220
Androcera, 8
Antennaria, 139
Anthemis, 165, 168
Anthopogon, 74
Aplopappus, 106
Arctium, 193

y Neg ols irot= Wel OF E4/
Artemisia, 171
Asperugo, 53
Aster, 109

Bahia, 163
Balsamorhiza, 154
Bartsia, 27
Besseya, 23
Bidens, 160
Boltonia, 108
Borago, 54
Boschniakia, 32
Brickelia, 96
Buchlo#, 149
Calendula, 192
Campanula, 80
Carduus, 195
Cartholinum, 37
Castilleja, 24, 219
Centaurea, 200
Chaenactis, 164
Chaenorrhinum, 14
Chamaesaracha, 6
Chamomilla, 168
Chelone, 14
Chondrophylla, 73
Chrysanthemum, 169
Chrysopsis, 98, 105
Chrysothamnus, 108
Cichorium, 201

222

Cirsium, 196
Cnicus, 198
Collinsia, 14
Collomia, 43
Conopholis, 32
Convolvulus, 9
Conyza, 138
Coreopsis, 159,
Cota, 165
Crepis, 211
Cryptantha, 52
Cuscuta, 9, 217
Cyclachaena, 150
Cynoglossum, 50
Dasystephana, 71
Datura, 8
Diplopappus, 106
Doellingeria, 105
Downingia, 83
Dracocephalum, 63
Echinacea, 154
Echinops, 193
Echium, 58

Ellisia, 46
Elscholtzia, 70, 221
Erechtites, 177
Erigeron, 127
Erodium, 40
Eryngium, 193
Eupatorium, 94
Euphrasia, 27
Franseria, 151
Gaillardia, 164
Galeopsis, 65
Galinsoga, 161
Gentiana, 70
Gentianella, 73
Geranium, 38, 220
Gerardia, 23, 218
Gilia, 43

Glechoma, 63
Gnaphalium, 148
Gratiola, 18
Grindelia, 97
Gutierrezia, 98
Hackelia, 51
Halenia, 75
Haplopappus, 106, 107

182

1972

Hedeoma, 68
Helenium, 164
Helianthus, 155
Heliopsis, 152
Heliotropium, 50
Hieracium, 214
Homopappus, 107
Hydrophyllum, 45
Hymenopappus, 162
Hymenoxis, 163
Hyoscyamus, 6
Hypochaeris, 203
Hyssopus, 68
Impatiens, 41
Iva, 149

Knautia, 84
Krigia, 203
Laciniaria, 97
Lactuca, 207
Lallemantia, 63
Lamium, 65
Lappula, 50
Lapsana, 202
Leontodon, 212
Leonurus, 66
Lepachys, 154
Leptilon, 138
Leucanthemum, 169
Leucophysalis, 6
Liatris, 96
Limosella, 19, 217
Linanthus, 43
Linaria, 13
Linum, 36
Lithospermum, 57, 220
Lobelia, 82
Lomatogonium, 75
Lycium, 5
Lycopsis, 54
Lycopus, 68
Lygodesmia, 204, 209
Machaéeranthera, 126
Madia, 162
Marrubium, 62
Matricaria, 167
Megalodonta, 161
Melampyrum, 26
Melissa, 68
Mentha, 69, 221
Menyanthes, 75
Mertensia, 56, 220

Boivin, Flora of Prairie Provinces

223

139

Microseris, 202
Mimulus, 17
Moldavica, 63
Monarda, 67
Mulgedium, 209
Myosotis, 55
Myzorrhiza, 32
Nabalus, 214
Navarretia, 43
Nemophila, 46
Nepeta, 63

Nonea, 55
Nothocalafs, 202
Odontites, 27
Oligoneuron, 103
Onosmodium, 58
Oreocarya, 52
Orobanche, 32, 220
Orthocarpus, 26
Oxalis, 40
Pedicularis, 28, 219
Penstemon, 14, 217
Petasites, 175
Phacelia, 46
Phlox, 42
Physalis, 6
Physostegia, 64
Picradeniopsis, 163
Picris, 203
Pinguicula, 33
Plagiobotrys, 52
Plantago, 76
Pleurogyne, 75
Polemonium, 44
Prenanthes, 213
Proboscidea, 35
Prunella, 64
Psilocarphus, 139
Pyrethrum, 168
Pyrrocoma, 107
Ratibida, 154
Rhinanthus, 28
Romanzoffia, 48
Rudbeckia, 153
Salvia, 67
Saussurea, 194
Scrophularia, 14
Scutellaria, 61
Senecio, 177, 183
Sideranthus, 106
Silybum, 200

1,0 Bev ORION TcA

Solanum, 7
Solidago, -98
Sonchus, 206
Stachys, 66
Stenotus, 107
Stephanomeria, 204
Symphytum, 54
Tanacetum, 170

Tonella, 13
Townsendia, 108
Tragopogon, 204
Unamia, 105
Utricularia, 34
Valeriana, 4
Verbascum, 12
Vernonia, 94
Veronica, 19, 218

Vol. 23, nos

Taraxacum, 205
Tetraneuris, 163
Teucrium, 61
Thelesperma, 160

Veronicastrum, 22
Xanthium, 151
Xanthoxalis, 40

CONTENTS OF PART III

GONNATAS 6 caieec bese Sa eee ee cle os ees be celle eee) 6 Gets resol ene netenaamns

(Valerianaceae - Compositae)

Keys to Herbaceous Dicots with fused petals ....-.ececece 1
Additions and Corrections ......... © ose ee 6 e oie c ee raietemnemne

indexvole Genera ea sce ee cise cle.c eis 222

224

A NEW VARIETY OF POIRETIA LATIFOLIA
AND A BRIEF RESUME OF THE GENUS POIRETIA VENT. (LEGUMINOSAE)

Velva E. Rudd, Smithsonian Institution

Among recent collections made in Brazil by H. S. Irwin, et al.
is a unifoliolate Poiretia. Other known taxa of the genus are 3-
or 4-foliolate. The new taxon appears to be a variety of P. lati-
folia Vog. and is described below.

The faboid genus Poiretia Vent., with its lomentaceous fruit,
is included in the tribe Hedysareae DC., Prodr. 2: 307-353. 1825,
subtribe Poiretiinae Burkart, Darwiniana 3: 125. 1939.

POIRETIA Vent. Mem. Cl. Sc. Math.-Phys. Inst. Nat. France 1807(1)
:4. July 1807, nom. cons., non J. F. Gmelin 1791, nec Cav.
1797, nec J. E. Smith 1808.

Turpinia Pers. Syn. Pl. 2: 314. Sept. 1807, nom. rej., non Humb.
& Bonpl. Apr. 1807 ('1808'), nec Vent. Jul. 1807, nec Raf.
1808, nec Lex. 1824.

Plants herbaceous or suffrutescent, erect or scandent, to
about 3-4 m. high; stipules lanceolate, usually glandular, cadu-
cous; stipels present at base of lower pair of leaflets, linear
to lanceolate or ovate, usually glandular, caducous; leaves l-,
3-, or 4-foliolate; leaflets chartaceous to coriaceous, linear to
ovate, obovate, or suborbicular, glandular-punctate below, the
glands sometimes visible above; inflorescences axillary or ter-
minal, racemose, sometimes paniculate or spicate, few- or many-
flowered; bracts stipule-like; bracteoles lacking; flowers small
to moderate in size, 4-20 mm. long; calyx glandular-punctate,
campanulate with 5 subequal lobes shorter than the tube, the vex-
illar lobes smaller than the others and somewhat connate; petals
yellow, glandular-punctate, the vexillum reflexed, glabrous or
nearly 80; stamens 10, monadelphous, the anthers dimorphic, al-
ternately oblong and basifixed or elliptic and dorsifixed; ovary
sessile, 5-8-ovulate, glabrous or pubescent, the stigma terminal,
minutely capitate; fruit sessile, oblong, laterally compressed,
1-8-articulate, glandular-punctate, sometimes verruculose; seeds
brown, reniform, compressed, the hilum elliptic, lateral.

Type species: Poiretia scandens Vent., a synonym of P. punc-
tata (Willd.) Desv.

11

12 Per OL OG I A Vol. 23, mo. 2

Plants scandent; flowers 4-8 mm. long in axillary racemes; fruit
about 3 mm. wide; leaves 4-foliolate, the leaflets obovate
with margin often glandular but not completely hyaline.

Flowers 4-5 mm. long; calyx, fruit, and leaflets glabrous or
rarely pubescent. Mexico, West Indies, South America to

Perusand Brazel: < s . + « sl) euesrlehicns « « Le hempuncbanen

Flowers 6-8 mm. long; calyx, fruit, and leaflets pubescent or
the leaflets rarely glabrous. Brazil. . . 2. P. pubescens.

Plants usually erect; flowers 6-20 mm. long in axillary or ter-
minal racemes or panicles; fruit 5-6 mm. wide; leaves 1, 3-,
or 4-foliolate, the leaflets linear to ovate, obovate, or
suborbicular, the margin sometimes hyaline.

Flowers 6-8 mm. long; margin of leaflets not obviously hyaline.
Leaflets 3 or 4, linear to lanceolate or oblanceolate.

Leaves 4-foliolate; leaflets linear to linear-oblanceolate;

flowers about 6 m. long in axillary racemes. Brazil.
3. P. angustifolia.

Leaves 3-foliolate; leaflets linear-lanceolate to lanceo-
late or lanceolate-oblong; flowers about 7 mm. long in
terminal panicles. Brazil. ... .. . 4. Ps \longipes.

Leaflets 4, obovate to elliptic or suborbicular; Brazil, Uru-
Ruay, Argentinaele je «me sic) cle) ie 5. P. tetraphylla.

Flowers 10-20 mm. long; margin of leaflets glandular and usual-
ly hyaline.

Leaflets 4, ovate-elliptic, rounded at the apex and base.
Brazil, Paraguay. ... 6a. P. latifolia var. latifolia.

leaflets 1 or 3, ovate or elliptic, cordate or acute at the
base or sometimes rounded.

Leaves 3-foliolate; leaflets usually ovate, sometimes el-
liptic, the apex rounded or acute, the base cordate to
rounded. Brazil. . . 6b. P. latifolia var. coriifolia.

Leaves 1-foliolate; leaflets elliptic, acute at apex and
base. Brazil. . .. 6c. P. latifolia var. unifoliolata.

1. POLRETIA PUNCTATA (Willd.) Desv. Journ. Bot. Agric. 1: 122,
tab. 5, fig. 17. 1813; Mem. Soc. Linn. Paris 4: 308. 1826.

Glycine punctata Willd. Sp. Pl. 3: 1066. 1802. Type: Poiteau,
Haiti (B herb. Willd. no. 13469).

Poiretia scandens Vent. Mem. Cl. Sc. Math.-Phys. Inst. Nat.
France 1807(1): 4. July 1807; Choix, tab. 42. Nov. 1807.
Type: Turpin, "Domingo", "Saint-Dominique" (P).

Turpinia punctata (Willa. ) Pers. Syn. Pl. 2: 314. Sept. 1807.

Poiretia densiflora Mart. Flora 20, beibl. 2: 124. 1837. Type:
Martius, "in montibus apricis, saxosis, ad Rio de Janeiro, in
prov. minarum passim," Minas Gerais, Brazil (BR).

Poiretia multiflora Mart. & Gal. Bull. Acad. Roy. Brux. 10(2):
179. 1843. Type: Galeottii 3270, "savanes de Zacuapan," Vera-
cruz, Mexico (BR).

Poiretia refracta Griseb. Cat. Pl. Cuba 72. 1866. Type:

C. Wright 2311, Oriente, Cuba (GOEFT 7).

i

1972 Rudd, The gems Poiretia 13

ho?
42)

[Ventenat, Choix de Plantes. Plate

POIRETIA scandens.

hh P eYeCOM, OGM A Vol. 23, nose

Herbaceous or suffrutescent vine; stems to about 3 m. long,
generally glabrous or puberulent; stipules linear-lanceolate, at-
tenuate, 1-3 mm. long, 1 mm. wide; stipels linear-lanceolate, 1-2
mm. long, caducous; leaves 4-foliolate, the axis (0.5-) 1-4 (-6)
em. long; leaflets chartaceous, glabrous or nearly 80, obovate,
5-50 mm. long and wide, obtuse, sometimes emarginate, the base
acute, the margin not conspicuously hyaline; inflorescences axil-
lary, racemose; flowers 4-5 mm. long; calyx glabrous or, in South
America, sometimes puberulent, about 2 mm. long; fruit glabrous
or rarely somewhat puberulent, 2-3 cm. long, 3 mm. wide, the ar-
ticles 3-5 mm. long; seeds about 2.5-3 mm. long, 1-1-5 mm. wide.

Distribution: In wooded areas, usually climbing on shrubs,
Mexico, Central America, eastern Cuba, Hispaniola, Colombia, Ec-
uador, Peru, Bolivia, Venezuela, Brazil.

Representative collections:

MEXICO: Sinaloa: Mazatlan, Gonzalez Ortega 6396. Nayarit: Near
Mazatan on road to Varas, Feddema 1053. Mexico: Temascaltepec ,
Tenayac, Hinton 5142; Hinton 6751. Oaxaca: Rio Cascobel, Mell
2227, 2286. Oaxi Oaxaca ? or Veracruz ?: Consoquitla, iebmann “h696.
Veracruz: Zacuapan, Purpus 1879, 3046, 16215. Chiapas: Cascada,
Matuda 1618. Montecrista, Matuda 1976. Mun. Venustiano Carranza,
above above Soyatitan, Ton 3140. GUATEMALA: Huehuetenango: near Bee
tenango, Nelson 3574. pla tae Volcan Flores, Shannon 3677. Jal-
apa: Laguna de Ayarza, Heyde & Lux 174. EL SALVADOR: San Salva-
dor, Calderon 65; Standley 19190. | Ahuachapan , Standley 19926;
Padilla 567. HONDURAS: Morazan: San Antonio, Molina 735, 1159;
Swallen 10925. Copan: Between Hacienda Grande “and 0: Ostuman, Molina

24762. COSTA RICA: San Jose: Alto de Santa Barbara, Pittier 1650.
Escasu, Standley 32352. N of Santa Ana, A. Jimenez M. 1299. Ala-
juela: Alajuelita, Tonduz 9079. Panama: Cocle: Ola, Pittier 5082.
CUBA: Oriente: Punton de Cuero, N of Imfas, Morton & Alain 8871.
HAITI: Massif des Malheux, Thomazeau, Ekman H.1066. Massif de la
Selle, Petionville, Fourcy, Ekman H.1140. Morne 1a Seller lerre
Froid, Holdridge 1806. Marmelade, Leonard 8431. St. Michel de 1'
Atalaye, Leonard 7176. Cabaret, Leonard & & Leonard 11897. Port de
Paix, Leonard & Leonard 1111. eae Rabel, Leonard & Leonard 4313,
4572. Bombardopolis, Leonard & Leonard 13373. 73. DOMINICAN REPUBLIC:
Without exact locality, Wrigl Wright, "Parry, & Brummel 99. Santiago:
Igua, J. Jimenez 1189. Monte Colorado near La Placeta, Jd. Jiménez
3818. San Juan: Piedra del Aguacate to Rfo del Oro, Howard & How-
ard 9395. Barahona: La Hotte, Howard 12266. COLOMBIA: Cund Cundina-
marca: Fusagasuga to Pandi, Pennell | 2ejey7. Cauca: Popayan, Lehmann
4752. Huila: Neiva, Mosel ands Caicedo 4. VENEZUELA: Distrito to Fed-
eral: Near Caracas, Pittier 7430, 7431. Between Antimano and Las
Adjuntas, Pittier 12253. Rio Macarao, Jahn 431. Between Baruta
and Sabana Grande, Ll. Williams 10589. Between Caracas and La
Guaira, Ll. Williams & Alston 34. Miranda: Las Mostazas, beyond
Los Teques, Allart 195. Oripoto, Gines 4562. Carabobo: Valencia,

1972 Rudd, The genus Poiretia 1h5

Pittier 8684. Bolfvar: Between La Paragua and El Cristo, Killip
37617. ECUADOR: Barraganetal, Stevens 303. PERU: Cajamarca: Be-
tween Santa Cruz and Tamborapa, Lopez, Sagastegui, & Collantes
4122, BOLIVIA: Santa Cruz, Kuntze 6. n., May 1892. BRAZIL: Per-
nambuco: Tapera, Pickel 3104. Caruaru, Lima 50-524. Bahia: Mach-
ado Portello, Rose & e & Russell 20008. Goids: Between Posse and Al-
vorado, Irwin et al. 14956. - Distrito Federal: Brasflia, Pires et
al 9549. Minas Gerais: Lagoa Santa, Warming s. n. Belo Horizonte,
Capoeirinha, Magalhaes 3031. Paraopeba, Heringer [RB no.] 88225.
Rio de Janeiro: "St. Salvador," Riedel 451.

2. POIRETIA PUBESCENS Vog. 12: 52. 1838.
Type: Sellow, "ad Rio Jan.", Brazil (B destroyed; Field Mus.
Photo 2146 ex B; F fragment).

Herbaceous vine with stems to about 4 m. long, generally pub-
erulent; stipules lanceolate, attenuate, 3-6 mm. long, 1-1.5 m.
wide; leaves 4-foliolate, the axis 2-5.5 cm. long; leaflets char-
taceous, puberulent or rarely glabrous, obovate, 1-4 cm. long,
5-30 mm. wide, rounded at the apex, the base cuneate, the margin
not hyaline; inflorescences axillary, racemose or spicate;
flowers 6-8 mm. long; calyx puberulent, about 2 mm. long; fruit
puberulent, 2-3 cm. long, 3 mm. wide, the articles 4-6 mm. long;
seeds about 2.5-3 mm. long, 1 m. wide.

Distribution: In wooded areas, climbing on shrubs, Brazil:
Goias, Minas Gerais, Sao Paulo.

BRAZIL: Goias: Chapada dos Veadeiros, ca 5 km NE of Francisco Sd,
road to Salinas, Irwin et al. 23194. Minas Gerais: Mun. Santa
Luzia, Fazenda da Chicaca, Assis 118. Carandai, Quebra-Cambao,
Duarte 654. Estrada dos Borges, near Belo Horizonte, Agronomic
Institute, Irwin 2336. Serra do Espinhaso, ca 15 km N of Sao Joao
da Chapada, Irwin et al., 28123. Sao Paulo: Ypanema, Riedel 2043.

3. POLRETIA ANGUSTIFOLIA Vog. Linnaea 12: 53. 1838.
Syntypes: Sellow, Minas Gerais, Brazil (B destroyed, Field Mus.
Photo 2143 ex B); Manso & Lhotzky, Minas Gerais, Brazil.

Erect, suffrutescent herb, to about 70 dm. high, generally
glabrous; stipules linear, attenuate, 5-7 mm. long, 0.5 mm. wide;
stipels linear, minute, fugacious; leaves 4-foliolate, the axis
1-2 cm. long; leaflets chartaceous, glabrous, linear to linear-
oblanceolate, 5-20 mm. long, 0.5-3 mm. wide, mucronulate, the
base cuneate, the margins revolute; inflorescences axillary, few-
flowered but in aggregate appearing spicate; flowers about 6 m.
long; calyx glabrous, about 2 m. long; fruit glabrous, 2-3.5 cm.
long, 5-6 mm. long, the articles about 5-6 mm. long; seeds about
2-2.5 mm. long, 1.5 mm. wide.

Distribution: In cerrado and margin of gallery woods, south-
eastern Brazil: Goias to Parana.

146 P HOY JBeOrT O/GeT & Vol. 23, noQor

BRAZIL: Without exact locality: "Mugy," Riedel 1553. Distrito
Federal: Bras{lia, Irwin et al. 11083. Goias: Serra dos Pirineus,
ca 15 km S of Corumba, Irwin et al. 10811. Serra do Fac&o, ca 25
km NE of Catalao, Irwin et al. Minas Gerais: Lagoa Santa, Warming
Ss. n. Uberava, Regnell III. hee, Uberlandia, Macedo 948. Morro das
Pedras, ca 26 km NE of Patrocinio, Irwin et al. 25573. Sao Paulo:
Sao Paulo, Vila Morais, Handro 45309. Parana: Jaguariaiva: Ponta
Grossa, Parque Vila Velha, Hatschbach 13106. "Rod. Jaguariaiva-
Senges, Hatschbach 8654.

4, POIRETIA LONGIPES Harms, Fedde Repert. Sp. Nov. 9: 440. 1911.
Type: Peckolt, July 1911, Sao Paulo, Brazil (B destroyed;
Field Mus. photo 2144 ex B; F encase

Poiretia longipetiolata Hoehne, Bol. Inst. Brasil. Sc. 2: 2k7,
pl. 3. 1926. Type: Gehrt 4043, Sao Paulo, Brazil, "Pedregulho,
interior do Estado" (SP 7).

Erect subshrub, generally glabrous; stipules and stipels not
known; leaves 3-foliolate, the axis 3-12 mm. long; leaflets sub-
coriaceous, glabrous, linear-lanceolate to lanceolate or lanceo-
late-oblong, 7-15 mm. long, 1-2 mm. wide, mucronulate, the base
cuneate; inflorescences terminal, paniculate and axillary, race-
mose; flowers about 7 mm. long; calyx glabrous, 2.5-3 mm. long;
fruit glabrous, 1.5-2.5 cm. long, 3-4.5 mm. wide, the articles
5-8 mm. long.

Distribution: Sao Paulo, Brazil.

5. POLTRETIA TETRAPHYLLA (Poir.) Burkart, Darwiniana 3: 224. 1939.
Psoralea tetraphylla Poir. in Lam. nee Gee Meth. Bot. suppl.

589. 1816. Type: Commerson, Buenos Aires, Argentina (P).
Poiretia psoraloides DC. Prodr. 2: 315. 1825. Based on Psoralea

tetraphylla Poir.

Erect, suffrutescent herb, to about 1 m. high, generally glab-
rous or subglabrous; stipules lanceolate, attenuate, 5-10 mm.
long, 1 mm. wide; stipels about 2-4 m. long, 0.5 mm. broad, at-
tenuate, caducous; leaves 4-foliolate, the axis about 2-5 cm.
long; leaflets chartaceous to subcoriaceous, glabrous, obovate to
suborbicular, about 5-25 mm. long, 5-20 mm. wide, the apex and
base usually rounded, the margin not conspicuously hyaline; in-
florescences racemose, terminal or axillary; flowers 6-8 mm.
long; calyx glabrous or subglabrous, 2-3 mm. long; fruit glabrous,

1-4 om. long, 5-6 mm. wide, the articles about 5-7 mm. long;
seeds 3 mm. long and 2 mm. broad.

Distribution: In cerrado, grassland and open woods, southeast-
ern Brazil: Goids, Minas Gerais; Uruguay; northeastern Argentina.

1972 Rudd, The genus Poiretia 147

BRAZIL: Goias: Serra dos Pirineus, 15 km N of Corumbd de Goids
on road to Niquelandia, Irwin et al. 18594. Minas Gerais: "C.
Verde," Macedo 653. Ituiutaba, Macedo 3194. Uberava, Regnell III.
43a. URUGUAY: Cerro Largo: R{o Negro, Rosengurtt et al. A-782.
Canelones: Atlantida, Rosengurtt B-2821. ARGENTINA: Chaco: Las
Palmas, Joergensen 2138. Las Brenas, Venturi 9748. Santa Fe:
Mocovi, Venturi 52. Corrientes: Empedrado, rado, Rfo Parana, Estancia
"Las Tres 6 Marias,” Pedersen 3013. Concepcion, Rincon de Luna,

Pedersen 4483.
6. POLRETIA LATIFOLIA Vog. Linnaea 12: 54. 1838.

Plants usually erect, sometimes scandent, suffrutescent, to
about 3 m. high, generally glabrous; stipules obliquely ovate to
lanceolate, acute, 5-15 mm. long, 3-5 m. wide; stipels 3-7 mm.
long, 1-1.5 mm. wide; leaves 1-, 3-, or 4-foliolate, the axis
1-7 cm. long; leaflets coriaceous or subcoriaceous, glabrous,
ovate to elliptic, about 1.5-7.5 cm. long, 1-5 cm. wide, acute or
obtuse, the base acute, rounded to cordate, the margin usually
densely glandular, hyaline; inflorescences commonly terminal,
paniculate, but also axillary; bracts obliquely lanceolate-oblong
about 2-10 mm. long, 1-2 mm. wide; fruit glabrous, 3-4 cm. long,
5-6 mm. wide, the articles 6-8 mm. long; seeds 2.5-3 mm. long and
2 mm. wide.

Distribution: In cerrado thickets, Brazil, Paraguay, Argentina.

6a. POIRETIA LATIFOLIA var. LATIFOLIA
Type: Sellow, Montevideo Uruguay (B destroyed).

The typical variety has 4-foliolate leaves, the leaflets ovate
to elliptic, 1-5 cm. long, 1-3 cm. wide, rounded at the apex and
base.

. Distribution: Brazil: Minas Gerais, Parana, Santa Catarina;
Paraguay.

BRAZIL: Minas Gerais: Serra do Espinhago, ca 4 km N of Sao Joao
da Chapada, Irwin et al. 28183. Paranda: Balsa Nova, Bugre, Hatsch-
bach 9526. Palmas, “S80 . Agostinho, Hatschbach 15429. Santa Catari-
na: W of Guritibanos toward Campos Novos, Smith & Klein 8340.
Chapeco, Fazenda Campo Sao Vicente, 24 e W of Campo Eré, Smith
et al. 9352. Lajes, Estrada da Rodagem S of Llajes, Smith & - Reitz
9972. Agua Doce, Fazenda Esperanga, Campos de Palmas, Sm: Smith wd t
Klein 13496. PARAGUAY: Tapyta, Joergensen 4550.

148 PHY T Od OGD A Vol. 23, no. 1

6b. POIRETIA LATIFOLIA var. CORIITFOLIA (Vog. ) Benth. in Mart. Fl.
Bras. 15(1): 79. 1859.

Poiretia coriifolia Vog. Linnaea 12: 55. 1838. Type: Sellow, "In
Brasilia meridionali: Sellow leg. (in Montevideo?)", , Uruguay
or Brazil (B destroyed).

This variety is readily distinguished by its 3-foliolate leaves,
the leaflets usually ovate, sometimes broadly elliptic, 2-7.5 cm.
long, 2-5 cm. wide, rounded or acute at the apex, rounded or cor-
date at the base.

Distribution: Brazil: Goias, Minas Gerais, Mato Grosso.

BRAZIL: Goigs: Chapada dos Veadeiros, Irwin et al. 12341, 24134,
24707. Serra dos Pirineus, 20 km NW of Corumba de Goids, Irwin et
al. 19237. Distrito Federal: Corrego Paranoazinho, near Sobradin-
ho, Irwin et al. 11500. Riberao Torto, ca 10 km NE of Brasf{lia,
Irwin et al. 13082. Mato Grosso: Serra do Roncador, ca 60 km N of
Xavantina, Irwin et al. 16666. Upper Rio Araguaia, S of Xavantina
Irwin et al. - 16827, 17343. Minas Gerais: Sitio, Dorsett et al.
299d. "C. Verde," Macedo 634. Ituiutaba, Macedo 1614. Bento Pires
near Belo Horizonte, e, Williams & Assis 5926, 6110. Diamontina,
Pereira 1595; Irwin 2459. Ca 10 km SE of Paracatu, Irwin et al.
26234.

6c. POIRETIA LATIFOLIA var. UNIFOLIOLATA Rudd, var. nov.

Type: H. S. Irwin, R. H. Maxwell, D. C. Wasshausen 20431,
Brazil, "western n slopes, cerrado wi with “exposed sandstone cliffs
and outcrops, Serra do Cipo, Minas Gerais, ca. km. 112 (ca. 135
km. N. of Belo Horizonte), elev. 1200 m., 18 February 1968.
Holotype US 2590985 A; isotype NY.

A varietate typica foliis unifoliolatis, foliolis ellipticis,
apice basique acutis differt.

This variety is characterized by 1-foliolate leaves, the leaf-
lets elliptic, 1.5-3 cm. long, and about 1 cm. wide, acute at the
apex and base.

Distribution: Known only from the type collection.
The collections cited above are at US unless otherwise noted.

Additional collections from South America were cited by Burkart
in his treatment of the genus in Darwiniana 3: 222-228. 1939.

THE STATUS OF THE GENUS CRYPHIDIUM
(BRYOPHYTA: MUSCI)

Harold Robinson
Smithsonian Institution, Washington, D.C. 20560

During recent studies of South American mosses, I have
rather incidentally gathered information on the genus Cryphidium.
I have seen material of a number of the species that have been
placed in the genus, and the more recent transfer of most of
these species to other genera has been noted. At this time
I would complete the reduction of the genus.

Cryphidium was first established by Mitten (1869) as a
section of Neckera with one species, N. leucocoleum Mitt., from
Uruguay. The section with its single species was raised to
generic rank by Jaeger (1877). The extension of the genus to
include five Australian and Pacific Island species was the work
of Brotherus in the first edition of Engler and Prantl (1905).

A more proper alinement of these other species was indicated
first by Fleischer (1914) and then summarized by Brotherus in the
second edition of Engler and Prantl (1925).

In the Brotherus treatment of 1905, the concept of the genus
was altered greatly by the addition of the Australian and
Pacific species which had the capsules terminal on unspecialized
branches. These latter species are now placed in a separate
genus, Cyptodon, and the relationship is closer to Dendro-

cryphaea of Chile. Cyptodon and Dendrocryphaea come closest to
each other geographically in Chile where C. crassinervis Broth.
of Juan Fernandez has been mistaken for D. cuspidata (Sull.)
Broth. of the Chilean mainland. The differences between these
two species include the following, of which some are generic
characteristics. Cyptodon crassinervis has a more acute leaf
apex, lacks the minute papillae at the ends of the leaf cells,
has a smoother abaxial surface on the costa, lacks the small
area of slightly larger cells below the larger area of trans-
versely elongate alar cells, and has a flatter rather than
conical operculum.

There remains the question of the generic distinctions of
Cryphidium as represented by the type species. In the recent
treatment in Musgos Sul-Brasileiros II by Sehnem (1970), the
genus is separated from Cryphaea by the portion of the Conspecto
stating "arquegoniarios terminais em ramos mais ou menos
alongados" versus "Arquegoni4rios em ramos curtissimos laterais".
This incorrect characterisation of Cryphidium seems to be
derived from the earlier work of Brotherus (1905).

Examination of recent material collected in Uruguay (Zorron,
3933) shows that Cryphidium differs from Cryphaea only by the

149

150 PAY PO: OG a8 VoL. 23, no. 1

completely rounded leaf apices. The continued separation of the
two genera seems to be a relict of century-old thinking where
pleurocarpous mosses with blunt leaves were rather mechanically
thrust into the genus Neckera. The existance of some members
of the Neckeraceae with similar leaves such as Leptodon smithii
(Hedw.) Web. & Mohr and species of Pinnatella must have helped
perpetuate the separation.

On the basis of my observations I do not consider
Cryphidium generically distinct from Cryphaea. The following
new combination is necessary.

Cryphaea leucocolea (Mitt.) H.Robinson, comb. nov. Neckera
Teucocolea Mitt., J. Linn. Soc. Bot. 12: 457. 1869.

Literature Cited

Brotherus, V. F. 1905. Hedwigiaceae-Rhacocarpeae (Schluss),
Fontinalaceae, Climaciaeceae, Cryphaeaceae, Leucodontaceae,
Prionodontaceae und Spiridentaceae. Die Natitirlichen
Pflanzenfamilien 1(3). 223: 721-768.

1925. Musci (Laubmoose). Die Natiirlichen Pflanzen-
familien, Ed. 2. 11: 1-52.

Fleischer, M. 1914. Kritische Revision von Carl Miillerschen
Laubmoosgattungen I. Hedwigia 55: 280-285.

Jaeger, A. 1877. Adumbatio florae muscorum. Sectio IV.
Pleurocarpi. Trib. XXVI. Leucodontaceae - Trib. XXVIII
Hookeriaceae. Bericht St.Gallischen naturw. Ges.

1875-76: 201-371.

Mitten, W. 1869. Musci austro-americani. Enumeratio muscorum
omnium austro-americanorum auctori hucusque cognitorum.
J. Linn. Soc. Lond,m Bot. 12: 1-659.

Sehnem, S. J. Aloysio 1970. Musgos sul-brasileiros II.
Pesquisas Bot. 28: 1-117.

STUDIES IN THE EUPATORIEAE (ASTERACEAE). LXV.

A NEW GENUS, NEOCABRERTA.

R. M. King and H. Robinson
Smithsonian Institution, Washington, D.C. 20560.

There is a small group of South American species that
because of superficial differences have not previously been
placed together in systems of classification. Microscopic
examination proves that these species are very closely related
to each other, sharing hairs on the inner surface of the corolla
and deeply bilobed anther appendages. In general characters
the species are Critonoid though the stylar appendages are
slightly more papillose than most members of that group. In
general habit the group is more like some members of the Gyptoid
series which is prominent in eastern South America, but none of
these have hairs on the inside of the corolla.

One of the reasons the species have been separated in
treatments is because of the pronounced hairyness of one of the
species, a condition that extends on to the surface of the
receptacle. The real relationships of the species seem very
remote from anything else which has been placed in the section
Hebeclinium, however. The two primary species also differ in
the number of flowers per head. Neocabreria serrulata is usually
cited as having 10-12 flowers while N. malacophylla usually has
20-25 flowers.

With this distinctive genus we wish to honor the noted
South American botanist and author of many papers on the
Eupatorieae, Angel L. Cabrera, Director of the Museo de La Plata,
Argentina.

Neocabreria R.M.King and H.Robinson, genus novum Aster-
acearum (Eupatorieae). Plantae herbaceae vel suffrutescentes
erectae pauce ramosae. Caules subglabri superne parce vel dense
pubescentes. Folia opposita breviter petiolata angusta ellipti-
ca basi cuneata margine serrulata superne parce pubescentia
inferne villosa. Inflorescentiae corymboso-paniculatae;
pedicelli pubescentes. Involucri squamae imbricatae 25-30
inaequilongae 3-4 seriatae oblongae extus parce pubescentes
intus glabrae vel hirsutae; receptacula plana vel pauce convexa
glabra vel dense hirsuta. Flores 10-25 in capitulo; corollae
anguste infundibulares 5-lobatae extus inferne glabrae intus
hirsutae, cellulis oblongis parietibus interioribus plerumque
Sinuosis, lobis aequilateraliter triangularibus vel longioribus
extus glabris vel glanduliferis ad apicem vix mamillosis intus
laevibus; filamenta antherarum in parte superiore angusta,

151

152 PHY? OL Oj/G A Vol. 23, (m0.

cellulis infernis plerumque quadratis, parietibus annulis
plerumque transversis ornatis, thecis basi non hastatis cellulis
exothecialibus subquadratis, appendicibus antherarum valde
bilobatis. styli inferne non incrassati glabri, appendicibus
late linearibus dense mamillosis vel sublaevibus; achaenia
prismatica 4-5-costata setifera inferne angustata, punctis
plerumque in seriebus transversis; carpopodia indistincta;
pappus setiformis uniseriatus, setis 30-40 usque ad apicem
rigidis non dilatatis scabris persistentibus, cellulis apical-
ibus acutis.

Species typica: Eupatorium serrulatum A.P.Decandolle.

Our studies of the genus indicate that it contains the
following three species.

Neocabreria concinna k.M.King & H.Robinson,
nom. nov. Eupatorium concinnum A.P.Decandolle, Prodr. 5:
156. 1836. not E. concinnum Hook. & Arn. Brazil.

Neocabreria malacophylla (Klatt) R.M.King & H.Robinson, comb.
nov. Kupatorium malacophyllum Klatt, Jahrb., Hamburg. Wiss.
Anstalt., 9: 125. 1892. Eupatorium niederleinii Hieronymus,
Bot. Jahrb. 22: 763. 1897. S. Brazil to N. Argentina.

Neocabreria serrulata (A.P.Decandolle) R.M.King & H.Robinson,
comb. nov. Eupatorium serrulatum A.P.Decandolle, Prodr. 5:
147. 1836. Eupatorium acuminatum Hook. & Arn., Comp: Bot.
Mag. 1: 241. 1835 (1836). S.Brazil.

Acknowledgement

This study was supported in part by the National Science
Foundation Grant GB- 20502 to the senior author.

STUDIES IN THE EUPATORIEAE (ASTERACEAE). LXVI.

THE GENUS, PACHYTHAMNUS .

R. M. King and H. Robinson
Smithsonian Institution, Washington, D.C. 20560.

Careful reconsideration causes us to raise the rank of a
subgenus which we described some years ago (King & Robinson,
1970) under the genus Ageratina. The single species involved is
one of the most specialized toward xeric habitats of any in the
Eupatorieae. The thickened stems and the seasonally leafless
condition are reminiscent of other Mexican plants such as
Cnidoscolus and Senecio praecox A.P.Decandolle.

The striking differences in habit are reinforced by two
anatomical features of the flower. The corolla lobe is more
papillose on the outer surface than other species that we have
treated in Ageratina. This characteristic is of considerable
significance when the consistently thickened and smooth outer
surfaces of the lobes of other Ageratinas are considered. The
second distinction is the lack of enlargement at the base of the
style. Almost all other species in Ageratina and the closely
related genera have a distinct enlargement just above the
nectary. Sometimes this consists of simply a ring of enlarged
cells but in some cases a very large node is present. There is
a small group of species without an enlarged base on the style
that we retain in the genus Ageratina. This last group consists
of A. pentlandiana and its relatives in Peru, and these show
none of the other specialized features of the Mexican species
that we are placing here in a separate genus.

The genus Pachythamnus is closely related to Ageratina and
the carpopodium is of the type found in Ageratina subgenus
Neogreenella. It is the latter subgenus of Ageratina that we
feel contains the more primitive elements of that genus and there
is no reason to suggest that Pachythamnus has to be of particular
recent origin.

Pachythamnus(R.M.King & H.Robinson) R.M.King & H.Robinson,
new status. Ageratina subgenus Pachythamnus R.M.King & H.Robin-
son, Phytologia 19:228. 1970.

Perennial shrubs or small trees. Stems swollen, glabrous,
terete when fresh, leafless during dry season, with inflorescence
terminal on stems or branches and withering and falling after
flowering, new leafly branches by subfloral innovation. Leaves

153

154 PHY TOLOGIA Vol. 23, no. 1

opposite distinctly long petiolate; laminae broadly ovate to
deltoid, margin with a few blunt teeth. Inflorescence rather
densely corymbose; heads ca. 15 flowered; involucre of ca. 15
rather narrow usually acute subimbricate mostly subequal
phyllaries in 2-3 series; receptacle slightly convex, with
minute scattered hairs. Corolla narrowly funnelform; outer
surface and margins of lobes smooth below, with projecting cells
at the tip, without hairs or glands, without stomates; inner
surface of lobes papillose with long projecting cells; cells of
backs of lobes and of tube mostly narrow with sinuous walls.
Anther collar composed of numerous quadrate cells below, elongate
cells above, all with little or no ornate thickening on the walls;
exothecial cells in part usually lax and somewhat longer than
wide; anther appendage large, pollen spherical, tricolpate,
spinose, type II sometimes present. Style base glabrous, with-
out swollen node; surface cells of stvylar appendage densely long
projecting. Achene prismatic, usually 5-costate, bearing setae;
carpopodium distinct but without distinct upper limit, rounded,
with rather lax quadrate cells having thin beaded walls; basal
vasculature of achene united to upper level of carpopodium;
pappus of ca. 25 slender scabrous setae rather easily deciduous,
with pointed apical cells.

Type species: Fupatorium crassirameum B.L.Robinson

The genus is monotypic.

Pachythamnus crassirameus (B.L.Robinson) R.M.King & H.Robinson,
comb. nov. Eupatorium crassirameum B.L.Robinson, Proc. Am.
Acad. 35: 332. 1900. Mexico, C. Amer.

Acknowledgement

This study was supported in part by the National Science
Foundation Grant GB— 20502 to the senior author.

DISCOMYCETE FLORA OF ASIA, PRECURSOR IV:
A NEW SPECIES OF BIFUSELLA (RHYTISMATACEAE, HYPODERMATEAE )

ON CUNNINGHAMIA IN OKINAWA}

CHOEI OGIMI
Department of Forestry, College of Agriculture
University of the Ryukyus, Naha, Okinawa

and

RICHARD P. KORF2
Plant Pathology Herbariun
Cornell Untversity, Ithaca, New York

Cunmninghamia lanceolata Hook., a handsome Taxodiaceous species,
has been introduced in experimental plantings in the Yona Demonstra-
tion Forest on the island of Okinawa in the Ryukyus. In the course of
a survey of forest fungi, the authors noted a rather spectacular dis-
ease of this tree in that planting (FIG. 1), and the senior author has
twice since recollected the fungus. The causal agent is a species
referable to the tribe Hypodermateae of the family Rhytismataceae (us-
ually called the Hypodermataceae). It has the characteristic dumbbell
shaped spores which Darker (1932) termed "bifusiform." As is typical
of all members of the family, the ascospores are surrounded by a gela-
tinous sheath both within the ascus and on discharge (FIG. 5), and in
this species are often also coiled or twisted (FIG. 7).

The glistening black ascocarps are elongate ellipsoid, oriented
longitudinally on the leaf, a few occurring scattered on the upper
surface (FIG. 2), with a larger number occurring on the lower surface
where they are almost completely restricted to three rows that are
separated by peculiar whitish bands characteristic of the leaves of
the host species (FIGS. 3, 4). The ascocarps open by a longitudinal
cleft to expose the hymenium in wet weather, closing again under drier
conditions. Such ascocarps have been termed hysterothecia or uniapo-
thecial stromata.

I Previous articles in this series appeared in Lloydia 26: 16-20, 21-
26 (1963), 28: 113-119 (1965).

2. Supported in part by National Science Foundation Grant GB-8548, and
also by the U.S. Office of Education, the University of the Ryu-
kyus, and the U.S. Civil Administration of the Ryukyu Islands
while the junior author was a Consultant in their Forest Diseases
program.

155

156 PB ot (0) .0hG Ek Vol..23, no. a

In a recent revision of the family (Darker, 1967) several new
genera and new generic concepts have been advanced. Bifusiform asco-
spores occur in five genera. In Btfusepta Darker, on Vacctntum, the
spores are septate; in the others, as in the Okinawan fungus, the
spores are non-septate. In Duplicaria Fuckel the ascocarps are sub-
spherical, open irregularly, and are not on conifers. The species of

FIGS. 1-4. Photomicrographs of Bifusella cunninghamticola on the dead
leaves of Cunninghamia lanceolata. FIG. 1. Portion of a branch with
ascocarps, x 1. FIG. 2. Upper surface of a leaf with scattered asco-
carps, x 2. FIG. 3. Lower surface of the same leaf with ascocarps in
three longitudinal rows, x 2. FIG. 4. Ascocarps at a greater magni-
fication, longitudinally oriented clefts visible, developing almost
wholly between the two longitudinal whitish bands on the lower leaf
surface, x 16. All photographs from the holotype specimen, taken by
Howard H. Lyon, Department of Plant Pathology, Cornell University.

1972 Ogimi & Korf, Discomycete flora of Asia 157

the remaining three genera were all at one time or another referred to
the genus Bifusella von Hthnel, have ellipsoid to linear ascocarps,
and occur on coniferous hosts. It is to this "Bifusella complex"

FIGS. 5-8. Camera lucida drawings of Bifusella cunninghamiicola, from
the holotype specimen, x 1000. FIG. 5. Mature ascospore showing the
thick gelatinous sheath and various inclusions within the spore. FIG.
6. Apices of paraphyses. FIG. 7. An ascus containing eight, mostly
vermiform, bifusiform ascospores. FIG. 8. Vertical freezing micro-
tome section through the extreme edge of an ascocarp, the thick-walled
epidermal cells at the base, with even thicker-walled hypodermal cells
below them, the host cuticle (shown shaded) raised by the ascocarp and
adhering closely to the upper surface of the ascocarp, the cells of
the stroma mostly gelatinized at the margin and difficult to differen-
tiate, at the upper portion of the section at the left with cells more
typical of the stroma, forming a brown-walled textura angularis.

158 Palpta Ts Op, LsOuGe TA: Vol. 23, nos 1

(Darker, 1967) that the species on Cumninghamia belongs.

Btfusella was erected by von Hthnel (1917) for a single species,
B. linearis (Peck) v. H&hn., on the needles of Pinus strobus L. He
noted as generic distinctions that the ascocarps are subcuticular (ra-
ther than intraepidermal as in Hypoderma), the bifusiform shape of the
spores, and the absence of paraphyses. A second species, on Abtes,
was added to the genus by Dearness (1926), and three new species, on
Abies, Picea and Pinus, brought the number of species recognized by
Darker (1932) to five. That critical monograph of the species of this
family on conifers resulted in a much broadened generic concept.

Darker's (1967) more recent treatment of Btfusella differs signi-
ficantly from that in his earlier monograph. The subepidermal B.
stritformts Darker, on Pinus, is now removed to become the monotype of
a new genus, Soleella Darker. The two "nervisequious" species on
Abies are now both treated in the new genus Isthmiella Darker as I.
abtetts (Dearn.) Darker and I. faulti (Darker) Darker. The non-nervi-
sequious species on Picea became I. crepidiformis (Darker) Darker.
These four species differed from the type species, B. linearts, in
having persistent paraphyses and in not being subcuticular.

The newer treatment, having removed all but the type species from
the genus, then proceeded to add two species Darker (1932) had earlier
placed in Hypoderma. Neither of these agree very well with the type
species in spore morphology, for they have cylindrical or rod-shaped
ascospores which at most could be termed "tending toward a bifusiform
condition" as mentioned in the revised generic diagnosis for Btfusel-
la. Both B. saccata (Darker) Darker and B. pini (Dearn.) Darker are
said, however, to be subcuticular?, unlike typical members of Hypoder-
ma, and both occur on soft pines as does the type species. All three
of the species now recognized by Darker also possess large, conspicu-
ous pycnidia.

Our fungus from Okinawa agrees well with the type species of Bz-
fusella both in spore characters and in being strictly subcuticular.
Unlike the three species on Pinus, however, pycnidia are unknown. The
presence or absence of paraphyses in those species has been variously
reported. For B. linearts von Hthnel (1917) reported: "Paraphysen
fehlen. Doch findet man hier und da einzele 1 yp dicke Hyphen zwischen
den Schlduchen."" These interthecial hyphae are apparently analogous
to the paraphyses seen in our material (FIG. 6). Darker (1932) une-
quivocally stated "Aparaphysate" for the species, but later (Darker,
1967, p. 1413) notes that "the paraphyses disappear so early in the
development that the species has come to be regarded as aparaphysate."
Of the two species he transfers from Hypoderma, he states that in B.
pint the paraphyses disappear early, while in B. saccata "the paraphy-
ses though more persistent finally undergo lysis and at maturity may
be lacking." In our fungus the paraphyses certainly do not disappear
early, since they may be found profusely in apparently completely
mature ascocarps which open widely to expose the hymenium in moist
weather. While it is possible that the paraphyses undergo lysis at
maturity or post-maturity in the Okinawan species, this has not yet

3, B. pini was stated by Darker (1932) to be intraepidermal.

1972 Ogimi & Korf, Discomycete flora of Asia 159

been demonstrated.

Bifusella cunninghamiicola Korf & Ogimi, sp. nov.

Hysterothecia ellipsoidea, nigra, subcuticularia, 0.3-0.4 *
(0.5-) 0.6-0.75 mm, amphigena; stroma ex textura angulart formata,
parietibus brunneis, cellulitis 3-6 (-8) wm diam.; asct late elavatt,
octospori, J—, (17-) 19-22 x 90-110 (-120) wm; ascosporae bt fust-
formes, plerwnque vermiformes, ad isthmun (2-) 3-4 um latae, ad ex-
tremitates 4.5-5.5 (-6.5) um latae, 28-36 (-40) wm Longae, hyaltn-
ae; paraphyses filiformes, graciles, 1 um latae, persistentes, ctir-
einatae, apice clavato vel difformi. Intra folta Cunninghamiae
lanceolatae in Okinawa. Holotypus: CUP-JA 3437.

Ascocarps ellipsoid, hysterioid, black, 0.3-0.4 x (0.5-) 0.6-0.75
mm, opening widely when moist by a longitudinal cleft that is faintly
fimbriate and somewhat whitish to expose the pallid to beige disc,
amphigenous (fig. 1), sparse and irregularly disposed on the glabres-
cent upper surface of the leaf (fig. 2), more densely arranged into 3
longitudinal rows on the lower surface (fig. 3), the rows separated by
broad whitish bands characteristic of the host species (fig. 4), the
long axis of the ascocarp parallel to the long axis of the leaf, sub-
cuticular (fig. 8). Spermatia or conidia unknown, presumably absent.
In section: hymenium 110-120 um thick; subhymentum and medullary tis-
sue indistinguishable from one another, very reduced, 5-10 um thick,
of textura intricata seated on the scarcely modified epidermal cells,
somewhat gelatinized toward the basal edges, hyaline; covering stroma
mostly of textura angularis with somewhat thickened, brown walls, the
individual cells 3-6 (-8) um in diam, paler and more hyphoid at the
base, ca. 20 um thick at the sides, flaring to 4O um thick toward the
cleft, at the cleft suddenly enlarged to ca. 80 um thick and provided
there with subhyaline, glassy-walled apices ca. 15 x 3 um that form
the fringe at the cleft, covered nearly to the cleft by the closely
adherent host cuticle (fig. 8). Aset broad clavate, with a pore not
blue in iodine (J—), arising from gnarled, repeating croziers, 8-
spored, (17-) 19-22 x 90-110 (-120) um (Fig. 7). Ascospores bifusi-
form or dumbbell-shaped, hyaline, non-septate, often twisted or ver-
miform (fig. 7), (2-) 3-4 um wide at the central isthmus, ends swollen
to 4.5-5.5 (-6.5) um, length when not coiled 28-36 (-40) um, surround-
ed in the ascus and after discharge by a thick, hyaline, gelatinous
sheath (fig. 5), oil drops and other inclusions mostly polar and not
in the isthmus. Paraphyses about as long as the asci, gracile, mostly
unbranched below, ca. 1 um broad, remotely septate, often apically
branched one or more times with tips usually circinate, apices usually
clavately swollen or sometimes irregularly distorted, 1.5-2 (-3) um
broad, hyaline, persistent (fig. 6).

SPECIMENS EXAMINED: On leaves of Cunninghamia lanceolata, 20-50 m
elev., Yona Demonstration Forest, Yona, Okinawa, Ryukyus, 19. IV.
1969, Korf & Ogimi CUP-JA 3437, HOLOTYPE (CUP), ISOTYPES: DAOM, K,

160 PeHa<210eb 20 Gnbk Vol. 23, no. 1

TNS, Dept. Forestry, University of the Ryukyus, Naha, Okinawa. PARA-
TYPES: Same substrate, same locality, 10-100 m elev., 10. II. 1971,
Ogimt, Dept. Forestry, University of the Ryukyus, Naha, Okinawa; Same
substrate, same locality, 2. V. 1971, Ogimi, Dept. Forestry, Universi-
ty of the Ryukyus, Naha, Okinawa.

There are two other members of the Rhytismataceae reported on the
leaves of Cunninghamta lanceolata, but both should be easily distin-
guished on field characters as well as microscopic features. Neither
species has the constricted ascospores so characteristic of Btfusella
cunninghamticola. In the case of Lophodermium petrakii Durrieu (1957)
the long filiform ascospores (56-72 x 1.5-2.0 um) are borne in subcu-
ticular ascocarps which are scattered on the leaf with the clefts ran-
domly oriented. Darker's supposition that the reports of Lophodermium
pinastri (Schrad. per Hook.) Chev. (Teng, 1932), and of L. unetnatum
Darker (Teng, 1933), both on this host from China represent instead
Durrieu's species are at least partially confirmed by studies of the
junior author. The undestroyed portion of the Chinese National Fungus
Herbarium is on deposit in CUP (part of this collection is also on de-
posit at BPI). The report of L. pinastri is based on Teng 1005, but
that specimen was also cited when Teng (1933) apparently reidentified
it as L. unetnatum. While the Teng 1005 cannot be located at CUP, an-
other specimen cited in 1933 is deposited as L. unetnatum, Teng 1940.
This specimen is clearly L. petrakit. The intraepidermal ascocarps of
the second species, Hypoderma handelit Petrak*+, are also not regularly
alligned as in B. eunninghamitcola, and the bacillar ascospores are
shorter and l-septate (13-22 x 4-6 wm fide Petrak, 1947). Another
fungus which might conceivably be confused with these is Lophtosphaera
orientalis Hino & Katumoto (1964) which also occurs on the leaves of
Cunninghamia, but the ascostromata of that species are deeply immersed

4+. This species is referred to as H. cunntnghamtae (Keissl.) Teng by

Darker (1967), but Petrak's name is the correct one for the fun-
gus at species rank. Although H. strobtcola Tub. f. eunninghant-
ae Keissler (1924) was validly published, Teng (1936) did not
know of this paper and adopted Keissler's epithet only from a
herbarium label. Teng's name is not validly published under Art.
36 of the International Code of Botanical Nomenclature since he
provided neither a new nor a reference to a previously published
Latin description or diagnosis. When Petrak (1947) reworked
Keissler's Chinese material he cited primarily a later paper
(Keissler, 1937), decided the fungus deserved species rank, and
gave it a new name rather than adopting Keissler's epithet for
the forma (as he is permitted to do under the Code). The some-
what confusing synonymy is:

Hypoderma handelii Petrak (as "Handelii"), Sydowia 1: 371. 1947.
= Hypoderma strobicola Tub. f. Cunninghamtae Keissl., Akad.
Wiss. Wien Anzeiger, Math.-Naturwiss. Kl. 61: 13. 192k.
= [Hypoderma Cunninghamiae Teng (as "(Keissl.) Teng"), Sinensia
7: 261. 1936, not validly published. ]

1972 Ogimi & Korf, Discomycete flora of Asia 161

and the ascospores l-septate, fusoid, and appendaged; its asci are bi-
tunicate, as is characteristic of the Lophiostomataceae, to which the
species seems correctly referred.

As delimited and redefined here, the genus Bifusella is to be ac-
cepted for two, or possibly four, species, all of which occur on the
leaves of conifers. All of the other species assigned to this genus
which occur on conifers have been excluded by Darker (1967), and have
been discussed above. Two other species which do not occur on coni-
fers have, however, been referred to the genus, "B. vacctntt Tehon"
and B. acwninata (Ellis & Everh.) Bonar & Cooke. The first of these
is now the type species of Bifusepta, B. tehonit Darker, and occurs on
the stems of Vaccitniwn. The second, a species not mentioned by Darker
(1967), occurs on the culms of Juncaceae and Cyperaceae, and was ori-
ginally placed in the genus Duplicaria. Neither of these species now
appears to be congeneric with Bifusella linearis or B. cunninghamiico-
ta, and the junior author will discuss both at greater length in his
monographic study of Dupltcaria now in preparation.

It is not yet known how serious this disease is in Okinawa, nor
whether any successful control measures can be found. Though Cunning-
hamia lanceolata is widely grown in Asia, this disease has apparently
never been noted before its discovery in the experimental forest at
Yona. Whether the fungus was brought to Okinawa with the seeds of
Cunninghamia, or possibly exists also on some other Taxodiaceous or
Pinaceous host plant endemic to Okinawa, is also unknown. The senior
author is continuing his studies of the disease.

Be a
Cunninghamia lanceolata M SHXT Ii ROOM ZVWAIW H

RIN HED RERBRR RE UK. RAM cee
Seam, XXULT WA Bifuselta cunninghamiicola Korf & Ogimi sp.nov. 0”
HKG. SOT wipusctia Ba tyre TH} B. linearis &
Hee SMEG, —B. cunninghariicola (t FWTEOH) JB
Wat Cid fei, 2 tincaris it REWBIHK 77 HY AT,
Atk , Bifusetta Mit Hypodermatese 1c HX 109 DH, (dwii-

162 PsbsyeT JOybrO"G: Dee Vol. 23, no. 1

caria, Bifusepta, Isthmiella, Soleella) kb <c%21% rk Be SMT
VE a Bifusella cunninghamticola ¢t C. lanceolata (< REG

Ow Et & 22 iB) Sou °F V\ (092 Fe Paes Lophoder-

mium petrakit, Hypoderma handelii (= H. cunninghamiae) t& Lophio-

sphaera orientalis ‘4% An REY Me st ee a4 WG,

LITERATURE CITED

DARKER, G. D. 1932. The Hypodermataceae of conifers. Contr. Arnold
Miglooien, HG aalsik, 2i jaithe

DARKER, G. D. 1967. A revision of the genera of the Hypodermataceae.
Canad. J. Bot. 45: 1399-14hh.

DEARNESS, J. 1926. New and noteworthy fungi— IV. Mycologia 18:
236-255.

DURRIEU, G. 1957. Deux Hypodermataceae nouvelles sur coniferes.
Beih. Sydowia 7: 355-358.

HINO, I., and K. KATUMOTO. 1964. A new species of the genus Lophto-
sphaera on Cunninghamia lanceolata Hook. (Studies on fungi para-
sitic on introduced foreign trees in Japan. I.) Bull. Fac. Agr.

ia Yamaguti Univ. 15: 517-522.

HOHNEL, F. von. 1917. Mycologische Fragmente. CXXXVI. Uber Rhytts-
ma lineare Peck. Ann. Mycol. 15: 318-319.

KEISSLER, K. 1924. Fungi novi sinenses a Dre. H. Handel-Mazzetti
lecti. II. Akad. Wiss. Wien Anzeiger, Math.-Naturwiss. Kl. 61:
10-13.

KEISSLER, K. 1937. Micromycetes. Jn HANDEL-MAZZETTI, H. Symbolae
Sinicae 2: 1-36. Julius Springer, Wien.

PETRAK, F. 1947. Kritische Studien tier chinesische Pilze. Sydowia
UE SSyP=SiCl(s

TENG, S. C. 1932. Fungi of Nanking II. Contr. Biol. Lab. Chin. Ass-
oc. Advancem. Sci., Sect. Bot. 8: 5-48.

TENG, S. C. 1933. Notes on Hysteriales from China. Sinensia 4: 129-
4h.

TENG, S. C. 1936. Additional fungi from China I. Sinensia 7: 212-
265.

DESCRIPTIONS OF NEW SPECIES AND COMBINATIONS
IN ERLOGONUM (POLYGONACEAE)

James L. Reveal

University of Maryland, College Park 20742
and
Smithsonian Institution, Washington, D.C. 20560

During the course of a preliminary revision of the genus
Exiogonum (Polygonaceae), a number of new combinations were found
necessary, and a few undescribed entities were discovered. For
the most part these new taxa and combinations will remain unpub-
lished awaiting a final monograph. Others, however, must be made
at this time in connection with various floristic projects in the
western United States.

Eriogonum ammophilum Reveal, spec. nov.

Enxiogono kearneyt Tidestr. affinis sed caulibus glabris, fol-
iis satis basalibus et latioribus, et pedunculis obviis; a E.
nummulart M.E. Jones foliis non rotundis et caulibus non lanatis
differt.

Frauticukt 2-4 dm alti ex caudicibus subterraneis; folia dis-
persa secus caules ad basim, laminis ellipticis vel late ellipti-
cus, 1-2.5 (2.8) cm longis et 8-17 mm latis, subtus albo-tomento-
sis, supra subglabris et viridibus, petiolis 1-5 (15) mm longis,
basibus expansis petiolorum 1.5-2 mm longis et 1-1.5 mm latis, to-
mentosis; caules glabri, 5-15 cm longi; inflorescentiae cymosae,
glabrae, 1-2.5 mm longae; pedunculi glabri, (2) 5-10 (12) mm longi
et erecti; <nvolucra solitaria, turbinata, (2.5) 3-3.5 mm longa et
2-2.5 mm lata, glabra, lobis 5, acutis, 0.4-0.8 mm longis, bract-
eolis linearibus vel oblanceolatis, 1.5-2 mm longis, pedicellis
glabris, 2.5-4 mm longis; §20%e5 albi, 2-3 mm longi, glabri, tep-
alis + similaribus, anguste obovatis, (0.5) 1-1.7 mm latis; Stan-
Ana exserta, 2.5-3.5 mm longa, filamentis parce puberulis, anth-
eris roseis vel purpureis, 0.5-0.6 mm longis; achaents ignota.

Low herbaceous subshrubs, erect and spreading, 2-4 dm high,
2.5 dm across, the caudices woody, brown, deeply buried in sand,
leafless, the upper branches herbaceous, thin, glabrous and glau-
cous; £eaves solitary, alternate and rather closely congested a-
long the lower part of the branches just above the level of the
sand, the leaf-blade elliptic to broadly elliptic, 1-2.5 (2.8) cm
long, 8-17 mm wide, densely white-tomentose below, less so to sub-
glabrous and green above, the base obtuse, the apex obtuse to
rounded, the leaves soon deciduous, the plants leafless at matur-

163

16) PHY TO LOG Ta Vol. 23, no. 1

ity, the petiole slender, 1-5 (15) mm long, lanate to tomentose,
the petiole-base triangular, 1.5-2 mm long, 1-1.5 mm wide, lanate
or nearly so; 4Lowering stems slender, few to several per plant,
5-15 cm long, glabrous and glaucous, lanate amongst the leaves be-
low; <nglorescences cymose, trichotomous at the first node, dich-
otomous above, 5-20 cm long, up to 25 cm across, open, glabrous
and glaucous; bracts ternate, scale-like, 1-2.5 mm long, 1-2 mm
wide, triangular, glabrous without, tomentose within, connate at
the base; peduncles slender and erect, (2) 5-10 (12) mm long, gla-
brous and glaucous; involucres solitary, turbinate, (2.5) 3-3.5
mm long, 2-2.5 mm wide, glabrous within and without, the 5 acute
teeth 0.4-0.8 mm long, the bractlet linear to oblanceolate, 1.5-2
mm long, sparsely fringed with short capitate gland-tipped cells,
white, the pedicel glabrous, 2.5-3 mm long; (Lowers white with a
reddish-brown midrib and base, 2-3 mm long, glabrous within and
without except for a few microscopic glands scattered along the
midribs within, the tepals slightly dissimilar, the outer whorl
of tepals 1-1.7 mm wide, the inner whorl of tepals narrower, 0.5-
0.9 mm wide, united only at the base; stamens slightly exserted,
2.5-3.5 mm long, the filament sparsely puberulent basally, the
anther red to purple, 0.5-0.6 mm long, oblong; acheneds unknown.

TYPE: About |.3 miles northwest of Ibex Warm Point on a dry
sandy flat associated with Tetradymia, Onyzopsis and Ceratoides
[=Eurotia], T.22S., R.14W., sec. 30, at 5270 feet elevation, Mill-
ard Co., Utah, 4 August 1970, Holmgren & Holmgren 4650. Holotype:
US! Isotypes: BRY, NY, UTC!

ADDITIONAL SPECIMENS SEEN: UTAH: Millard Co.: Ibex Range
Exclosure just south of Ibex Warm Point on a sandy flat, T.22S.,
R.14W., sec. 32, 19 June 1965, R.C. Holmgren 479 (BRY, NY, USFS,
UNG

Extogonum ammophitum is a low herbaceous subshrub similar to
E. kReameyt Tidestr. and E. nummulare M.E. Jones. It resembles,
and is technically most closely related to, the Kearney Buckwheat
in its habitat, leaves and open inflorescence, but it is not as
woody in the above-ground parts as is typical of E. Reameyi. In
this respect, the new species approaches the rare, western Utah
species, E. nummulare. They share a similar inflorescence struc-
Ture and other features such as leaf pubescence type. However,
the two differ in shape and structure of the leaves, flowers and
stem vesture. As now seen, E. ammophilum represents an end line
in the evolutionary development of a small groups of species typ-
ified by E. Leptockadon Torr. & Gray.

These species may be separated as follows:
A. Leaves |inear-lanceolate to narrowly oblanceolate, |.5-4.5 am

long, 3-6 mm wide; branches tomentose or glabrate and flocc-
ose; flowers white or yellow; sandy deserts, Emery Co., Utah

1972 Reveal, New species & combinations in Eriogormum 165

southward to Coconino Co., Arizona, eastward into southwestern
Colorado and northwestern New Mexico....... E. Leptockadon

AA. Leaves oblanceolate to elliptic or orbicular to rounded, 0,5-
2.5 (3) cm long, 5-I5 mm wide; flowers white.

B. Leaves oblanceolate to elliptic, I-2.5 (3) cm long.

C. Stems and inflorescences tomentose, rarely glabrate;
leaves scattered along the lower stems; involucres
sessile, 2-2.5 mm long; common in western Utah from
Tooele Co. southward to northern Mohave Co., Arizona,
westward across southern and central Nevada to Washoe
Co., entering California in Mono Co. and extending
southward into Owens Valley of Inyo Co. €E. kearneyt

CC. Stems and inflorescences glabrous and glaucous; leaves
densely clustered near the base of the stems; involuc-
res peduncled, at least in the forks of the branches,
the peduncle (2) 5-10 (12) mm long, the involucre
(2.5) 3-3.5 mm long; local and infrequent, near Ibex
Warm Point, Millard Co., Utah...... E. ammophilum

BB. Leaves orbicular to rounded, 0.5-I| cm long and wide, clus-
tered at the base of the stems, densely white-tomentose
on both surfaces; stems densely lanate; involucres short
peduncled, the peduncles up to | cm long; rare, Known only
from sandy areas in extreme western Juab and Tooele cos.,
MN Ane Nat e.g n allssd a eisoulel w vigl cise iss a aes E. nummulanre

Eriogonum eremicum Reveal, spec. nov.

Ertogono osthundii M.E. Jones affinis sed foliis ovatis vel
rotundis, subtiliter tomentosis, laminis 1.2-2 cm longis et I-1.7
cm latis, involucris plerumque congestis, 2.5-4 mm longis.

PLantae herbaceae perennes 2.5-4.5 dm altae et caudicibus;
folia basalia, laminis ovatis vel rotundis, 1.2-2 cm longis et I-
1.7 cm latis, subtiliter albostomentosits, basi rotundis vel trun-
catis, petiolis 1-2.5 cm longis, tomentosis, bastbus expansis pet-
iolorum 2-3 mm longis et latis, tomentosis; caules glabri, 5-20
cm longi; <nflorescentiae cymae, glabrae, 12-25 cm longae; bxract-
eae ternariae, glabrae, 1-3 mm longae; pedunculé nulli; cnvolucra
plerumque congesta, 2.5-4 mm longa et 2-2.5 mm lata, lobis 5, ro-
tundis vel acutis, 0.4-0.8 mm longis, bracteolis linearibus, 1.5-
3 mm longis, pedicellis 2.5-4 mm longis, glabris; {lores albi,
2.5-3 mm longi, tepalis similaribus, obovatis, 0.8-1.5 mm latis;
Stamina exserta, 2.5-3.5 mm longa, filamentis parce puberulis,
antheris rubris, 0.5-0.6 mm longis; achaenta 3 mm longa.

Spreading herbaceous perennials 2.5-4.5 dm high and 1-2.5 dm
across, with short woody, somewhat congested, caudex branches 5-10
cm across, arising from a slender woody taproot; leaves strictly

166 P HEY GRsO Ores] A Vols 23, inde

basal, the leaf-blade ovate to round, 1.2-2 cm long, I-1.7 cm
wide, finely tomentose on both surfaces, the apex rounded, the
base rounded to truncate, the margin entire and plane, the pet-
iole 1-2.5 cm long, tomentose, the petiole-base triangular, 2-3
mm long and wide, tomentose to floccose without, subglabrous to
glabrous within; fLowering Stems erect, 5-20 cm long, glabrous
and green, remaining pubescent amongst the leaves at the very
base; inflorescences cymose, 12-25 cm long, 1-2 dm across, divid-
ed several times, trichotomous at the lower nodes, dichotomous a-
bove, the crown open, glabrous; bracts ternate, scale-like, 1-3
mm long, 0.8-1.3 mm wide, narrowly triangular, glabrous without,
sparsely tomentose within, connate at the base; peduncles lack-
ing; <nvoLucres solitary or more frequently clustered, turbinate,
2.5-4 mm long, 2-2.5 mm wide, glabrous within and without, the 5
rounded to acute teeth 0.4-0.8 mm long with a membranous margin,
the bractlet linear, fringed with minute gland-tipped cells, 1.5-
3 mm long, the pedicel 2.5-4 mm long, glabrous; flowers white
with a reddish-brown to greenish midrib and base, 2.5-3 mm long,
glabrous within and without except for minute glands along the
midrib within, the tepals essentially similar, obovate, 0.8-1.5
mm wide, the inner whorl of tepals slightly narrower, united less
than 1/5 the length of the flower; Stamens slightly to long ex-
serted, 2.5-3.5 mm long, the filament sparsely pubescent basally,
the anther red to rose, 0.5-0.6 mm long, oblong; achenes light
brown, 3 mm long, the broadly globose base tapering to a long 3-
angled beak.

TYPE: Common on scattered low sagebrush, Atriplex and OrAy-
Zopsds& covered hills on gravelly soil, I7 miles southeast of Gar-
rison, Millard Co., Utah, along Utah Highway 21, at 5650 feet el-
evation, 23 July 1965, Holmgren, Reveal & LaFrance 2247. Holo-
type: UTC! Isotypes: ARIZ, BRY, CAS, GH, MO, NY, OKL, RM, RSA,
UC! -- Distributed under the name Extogonum batemanti M.E. Jones.

ADDITIONAL SPECIMENS SEEN: UTAH: Millard Co.: Antelope
Valley, 21 August 1967, A&dex 73 (BRY, DRES!, USFS); West of Wa
Wa, 26 June 1906, M.E. Jones S5.n. (DS, NY, PH, POM, US); 2 miles
west of Cowboy Pass, 8 August 1941, Maguire 21228 (GH, NY, UC,
UTC, WTU); 25 miles southeast of Garrison on low l|imestone hills,
along Utah Highway 21, 6200 feet elevation, 28 July 1971, Reveak
2544 (NY, US, UTC); 10 miles east of Robinson's Ranch, | August
1963, Welsh & Moone 2251 (BRY).

This new species has been known for some time, being first
collected by Marcus E. Jones in 1906 and irregularly rediscovered
by various people associated with the United States Forest Service
and the Desert Range Experiment Station in Millard Co., Utah. It
is well isolated and separated from its two most closely related

1DRES stands for the Desert Range Experiment Station, U.S.
Forest Service, Milford, Utah.

1972 Reveal, New species & combinations in Eriogonum 167

species, Enxiogonum osthundii and E. batemanii, both originally
described by Jones in 1903.

Eniogonum batemanti is a wide spread species found in the
Colorado-Green river drainage system of eastern Utah and adjac-
ent western Colorado. Its range extends from the foothills of
the Uinta Mountains in the north, south to the Henry Mountains
of Garfield Co., Utah, and then eastward into Moffatt and Rio
Blanco cos., Colorado. It is similar to E. eremicun in the clu-
stered involucres, glabrous stems and strictly basal leaves. The
two differ in leaf shape and size, the type of the tomentum, the
flower length and general ecology of the sites - with E. bate-
manti most frequently being found on clay soils while E. erem-
4cum is encountered mainly on gravelly limestone soils.

The relatively restricted Erxtogonum ostlundii occurs among
pines (both Pinus ponderosa and P. edulis) and sagebrush along
the Sevier River drainage of Piute and Sevier cos., Utah. It is
clearly related to E. erxemicum, but the two differ in numerous
subtle ways in both ecology and morphology. They differ in leaf
shape, involucre size and number in any given cluster, flower
length and the branching pattern of the inflorescence.

These three species, plus the recently described Extogonun
cronquistii Reveal, all belong to a small group of species found
primarily in Utah. They may be distinguished as follows:

A. Leaves elliptic to spatulate, densely white-tomentose below,
floccose and green above, or if tomentose on both surfaces,
then plants of eastern Utah; leaf-blades tapering to the pet-
ioles.

B. Involucres 2-2.5 mm long, not clustered; leaf-blades |-1|.5
cm long, elliptic to spatulate; inflorescences with num-
erous, thin, branches making up more than half the height
of the plant; flowers |.5-2.3 mm long; peduncles, when
present, up to | cm long, erect; rocky slopes and clay
foothills and flats, Piute and Sevier cos., central Utah.
De Aes Se Wel TG oN Un el ISOM Tua ottde aati E. osthundii

BB. Involucres 2.5-4 mm long, usually clustered; eastern Utah
and adjacent western Colorado.

C. Leaves not crenulate, the leaf-blade (1.5) 2-4 cm
long, elliptic; plants 2-4 dm high, compact, not at
all spreading; flowers |.5-2.8 mm long; peduncles
lacking; clay slopes, mounds and washes, Duchesne and
Uintah cos. southward into Garfield Co., Utah, hence
eastward into western Colorado..... E. batemantii

CC. Leaves crenulate, the leaf-blade 0.5-2 cm long, elli-
ptic; plants (0.7) I-1.5 (2) dm high, spreading;

168 2 Yio Gi & Vol. 23, now 1

flowers (1.5) 2-3 mm long; peduncles lacking; steep
talus slopes, Bull Mountain, Henry Mountains, Gar-
fie drOad , wlitahes.<¢-.as'ate ewes enemeed E. cronquistit

AA. Leaves nearly rotund to ovate, densely white-tomentose on
both surfaces, the leaf-blade |.2-2 cm long, I-I.7 cm wide,
not tapering to the petiole; plants 2.5-4.5 dm high, the in-
florescence less than half the height of the plant; flowers
2.5-3 mm long; peduncles lacking; involucres 2.5-4 mm long;
dry slopes and rocky hillsides in gravelly limestone places
Of WesienueMililare Cor. Uirdlicsate sec ae cc ete = E. eremicwn

Eriogonum argophyllum Reveal, spec. nov.

A Extogono kingii S. Wats. differt foliis 4-8 mm longis et
2-5 mm latis, albo-tomentosis, petiolis 0.5-1 (1.5) mm longis,
scapis gracillimis, (4) 5-7 cm longis, involucris 2-2.5 mm long-
is, floribus luteis, 2.5-3 mm longis, tepalis oblongis.

Plantae humiles compactae perennes crassas tegetes formantes;
foltia oblanceolata vel elliptica, 4-8 mm longa, 2-5 mm lata, dense
utrinque albo-tomentosa, petiolis 0.5-1 (1.5) mm longis, tomento-
sis, basibus expansis petiolorum 1-3 mm longis et 172.5 mm latis;
Scapt erecti, gracillimi, (4) 5-7 cm longi, floccosi; bracteae
lanceolatae, 2-2.5 mm longae, membranaceae, sparse floccosae, 5-6;
AnvoLucta membranacea, 5-7, 2-2.5 mm longa, lobis lanceolatis,
6-7, extra sparse tomentosis et glandulosis, intra glandulosis,
bracteolis lineari-lanceolatis, 0.8-1.3 mm longis, pedicellis
glabris, 1.5-2.5 mm longis; {20x25 lutei, 2.5-3 mm longi, sparse
glandulosi extra secus costum, intus glandulosi, tepalis subaequ-
alibus, oblongis; stamina exserta, filamentis 3-3.5 mm longis,
sparse puberulis, antheris luteis; achaenia 2.8-3 mm longa.

Low compact caespitose herbaceous perennials forming a thick
spreading mat with numerous, spreading, woody caudex branches,
these densely covered with old persistent leaves and petiole-bases
over much of their length, arising from a slender, woody, brown-
ish taproot; Leaves basal, the leaf-blade oblanceolate to ellipt-
ic, 4-8 mm long, 2-5 mm wide, densely white-tomentose on both sur-
faces, not glandular, the margin entire and plane, the petiole
short, 0.5-1 (1.5) mm long, densely tomentose, the petiole-base
elongate-triangular, 1-3 mm long, 1-2.5 mm wide, sparsely pubes-
cent with long white hairs without, glabrous within, membranous
and + papery, light brown; flowering stems scapose, (4) 5-7 cm
long, floccose, erect and very slender; inflorescences capitate,
the head 5-10 mm across; bracts 5-6, lanceolate, 2-2.5 mm long,
sparsely floccose, membranous, connate at the base; peduncles
lacking; involLucres congested, 5-7 per head, membranaceous, tur-
binate-campanulate, 2-2.5 mm long, sparsely tomentose especially
at the apex without and with scattered glands along the ribs, min-
utely glandular within and often along the inner margin of the
tube, the 6-7 lanceolate, acutish lobes I-1.5 mm long, the lobes

1972 Reveal, New species & combinations in Eriogonum 169

and involucral tube totally hyaline, the bractlet linear-lanceo-
late, 0.8-1.3 mm long, the pedicel 1.5-2.5 mm long, glabrous;
flowers yellow with a greenish-yellow midrib and base, becoming
tinged with reddish-brown at maturity in some, 2.5-3 mm long,
sparsely glandular on the midribs without, glandular within es-
pecially along the midribs, the tepals subequal, oblong, the out-
er whorl 1-1.2 mm wide, the inner whorl 0.6-0.8 mm wide, united
about 1/3 the length of the flower; Stamens exserted, 3-3.5 mm
long, the filament sparsely puberulent basally, the anther yellow,
0.5-0.6 mm long, oblong; achenes brown, 2.8-3 mm long, the glob-
ose base tapering to a long, 3-angled beak.

TYPE: Common in sandy washes on crusty mineralized sand
with Senecto below Sulphur Hot Springs, Ruby Valley, Elko Co.,
Nevada, near Nevada Highway ||, 16.8 miles southwest of the jun-
ction with U.S. Highway 93, T.3IN., R.59E., sec. II, at 6050 feet
elevation, 7 July 1969, Holmgren & Kern 3661. Holotype: US!
Isotypes: BRY, NY, UTC!

ADDITIONAL SPECIMENS SEEN: NEVADA: Elko Co.: Sulphur Hot
Springs, 20 June 1941, A.H. Holmgren 1151 (NY, UTC); Sulphur Hot
Springs, 18 July 1971, Reveal & Reveal 2527 (BRY, NY, US, UTC).

One of the more fascinating features of Erxiogonum is the role
isolation has played in the development of species. Occasionally
the isolation barrier is geographic, where great distances have
separated entities and allowed them to develop independently of
each other along pathways which may be similar (no doubt due to
a relatively high percentage of similar genes) or quite different
due to the differences in selective pressures. This type of en-
demism is seen in E. gracthipes S. Wats. of the White Mountains
of California and Nevada, and in E. hofmgrenii Reveal, a species
that is restricted to the Snake Range of eastern Nevada. In this
case the morphological differences are not great, but the differ-
ences are consistent and definable. In the insular endemics off
the coast of California and northern Mexico, E. giganteum S.
Wats., E. zapatoense Moran and E. mokle Greene, here the morphol-
ogical differences are well pronounced, their geographical iso-
lation well enhanced, and yet some of these species can hybridize
when brought together in the garden.

Another feature of endemism in Extogonum is that determined
by edaphic specialization. A clear example of this is seen bet-
ween E. batemanti and E. erxemicum as noted above. However, there
is still a touch of spatial isolation here. Some species are
highly restricted in their distribution by the nature of the soil
they occur on. Exrtogonum saurinum Reveal is found only on Mowry
Shale in Dinosaur National Monument of extreme northeastern Utah
and adjacent Colorado. This vertical band of resistent rock is
only a few feet wide, and the species is not known to occur off
of this substrate.

170 P Bey ‘T1Oe LO Gor A Vol. 23, no. 1

In the case of Exiogonum argophylLum, both edaphic and spa-
tial isolation have played a role in the evolution of this spec-
ies. It is restricted to crusted mineralized sandy soil below
Sulphur Hot Springs at an elevation around 6050 feet. Its most
closely related species, E. kingti S. Wats., occurs less than 5
airline miles away, but at an elevation of from 9000 to 10,400
feet atop the East Humboldt Range and Ruby Mountains. Even given
this short distance, the two have evolved a number of morphologi-
cal differences, and these are of such a nature as to require a
specific rank to place each. At the upper elevations where E,.
kingit is found, it covers large areas of the slopes usually in
association with sagebrush, but with an occasional population
venturing out onto exposed ridges as near Angel Lake west of the
town of Wells. Enxtogonwn argophyllum occurs on more exposed sit-
uations, but at a lower elevation; even so, the overall nature
of the plants is similar.

The two species are easily distinguished on technical feat-
ures as noted in the key which followings the next species des-
CE Pilon.

Eriogonum scopulorum Reveal, spec. nov.

A Eniogono chrysopsidt Rydb. foliis (5) 6-9 (10) mm longis
et 2.5-4 mm latis, tomentosis et glandulosis, petiolis 2-4 mm
longis, scapis (3) 5-7 cm longis, glandulosis, involucris 2.5-3
mm longis, glandulosis et parce tomentosis, floribus stramineis,
2.5-3 (3.5) mm longis, tepalis oblanceolatis differt.

PLantae humiles compactae perennes crassas tegetes forman-
tes; folia oblanceolata vel elliptica, (5) 6-9 (10) mm longa,
2.5-4 mm lata, albo-tomentosa et glandulosa, petiolis 2-4 mm
longis, tomentosis et glandulosis, basibus expansis petiolorum
3-4 mm longis et (1.5) 2-2.5 mm latis; Scapi erecti, gracillimi,
(3) 5-7 cm longi, glandulosi; brzacteae triangulares, 1.3-1.6 mm
longae, membranaceae, sparse floccosae et glandulosae, 5-6; in-
voLucta membranacea, 5-7, 2.5-3 mm longa et lata, lobis 5-6, a-
cutis, extra glabris vel sparse tomentosis et glandulosis, intra
fere glabris, bracteolis linearibus, I-1.5 mm longis, pedicellis
glabris, 2.5-4 mm longis; 20x25 straminei, 2.5-3 (3.5) mm longi,
glabri, tepalis similaribus, oblanceolatis; stamina exserta, fil-
amentis 3-4 mm longis, sparse pilosis, antheris luteis; achaenta
2.5 mm longa.

Low compact cespitose herbaceous perennials forming a thick
spreading mat with few to several, spreading, woody caudex bran-
ches, these not densely covered with old leaves or petiole-bases,
arising from a slender, woody taproot; eaves basal, the leaf-
blade oblanceolate to elliptic, (5) 6-9 (10) mm long, 2.5-4 mm
wide, densely white-tomentose below, slightly less so and more
greenish above, glandular, the margin entire and plane, the pet-
iole short, 2-4 mm long, tomentose and glandular, the petiole-

1972 Reveal, New species & combinations in Eriogonum TTL

base narrowly triangular, 3-4 mm long, (1.5) 2-2.5 mm wide, thin-
ly tomentose without, glabrous within, membranous and papery,
light brown to tan; flowering stems scapose, (3) 5-7 cm long,
glandular, erect and very slender; inflorescences capitate, the
head 1-1.5 cm across; bracts 5-6, triangular, 1.3-1.6 mm long,
sparsely floccose and glandular without, glabrous within, conn-
ate at the base; peduncles lacking; <nvolucres congested, 5-7
per head, membranaceous, turbinate-campanulate, 2.5-3 mm long

and wide, glabrous without except for scattered glands near the
teeth, or sparsely floccose and glandular near the teeth and a-
long the ribs, sparsely pubescent within along the teeth, other-
wise glabrous, the 5-6 acute teeth I-1.5 mm long, the bractlet
linear, 1-1.5 mm long, fringed with elongated, white to hyaline
cells and marginal glands, the pedicel] 2.5-4 mm long, glabrous;
flowers pale yellow with a greenish midrib and base, becoming
tinged with pink in some at maturity, 2.5-3 (3.5) mm long, glab-
rous within and without except for scattered glands along the
midribs within, the tepals essentially similar, oblanceolate, I-
1.4 mm wide, the apex emarginate especially in fruit, united less
than 1/5 the length of the flower; Stamens slightly exserted, 3-4
mm long, the filament sparsely pilose basally, the anther pale
yellow, 0.5-0.6 mm long, oblong; achenes brown, 2.5 mm long, the
globose base tapering to a long, 3-angled beak.

TYPE: Among granite rocks near the top of a ridge south of
Aneroid Lake near the source of the East Fork of the Wallowa Riv-
er, Wallowa Mountains, Wallowa Co., Oregon, T.4S., R.45E., sec.
21, at 8500 feet elevation, 5 August 1966, Holmgren & Reveal 2957.
Holotype: US! Isotypes: BRY, NY, UTC! -- Distributed under the
name Exiogonum chrysops Rydb.

Exiogonwn Scopulorwn has been included previously under the
definition of E. chrysops Rydb., a low, desert range species of
southeastern Oregon. The exact nature of this species became ob-
vious recently while preparing the description of E. prociduwn
Reveal (in press) which is found in extreme northeastern Cal i f-
ornia and adjacent south-central Oregon. Once the true nature of
E. chrysops was established, two additional elements came to light
as well. These include E. Scopulonrwn, as defined here, and a sec-
ond species, yet to be described, from central Idaho eastward to
the Bitterroot Range of the Idaho-Montana state line region. The
plants from these high mountain ranges are currently being studied
and the results will be presented in 1973.

From Erxtogonum chrysops, E. Scopulorwm may be quickly dis-
tinguished by its glandular scapes and pale yellowish flowers.
The scapes of E. chtysops are floccose to tomentose and the flo-
wers are a bright yellow. In some respects, E. Scopulorwn is
similar to E. AoSsenSe Nels. & Kenn. of the Sierra Nevada, but the
involucre is totally different and the flower color and structure
is not the same.

172

PHYTOLOGIA You s'23, nosed

The series of species to which Ertogonum argophylum and E.
Scopulorum belongs is still under study as more isolated species
are being discovered; however, a key to the known elements is

presented below.
vol ucral

tube.

All of these entities have a membranaceous in-

A. Flowers yellow to pale yellow, not white or rose.

B.

BB.

Scapes glandular; leaves (5) 6-9 (10) mm long, 2.5-4 mm
wide, tomentose and glandular on both surfaces; flowers
pale yellow, 2.5-3 (3.5) mm long, the tepals oblanceol-
ate; involucres sparsely floccose and glandular or mere-
ly glabrous below and slightly pubescent near the teeth,
2.5-3 mm long; Wallowa Mountains, 8500 feet elevation.
Pee ogee tee Guo couse asinieke cae" «nie eich E. Scopulorun

Scapes floccose.

C.

CC.

Flowers bright yellow, 2.5-3 mm long; leaves I-2 cm
long, white-tomentose and eglandular on both surfac-
es, mostly spatulate, the petiole 2-5 mm long; invol-
ucres 3-5 per head, 2.5-3 mm long; low desert ranges
in southeastern Oregon mostly below 7000 feet eleva-

tion

nie Wee Wt Ae PE yh ROA ey TP E. chrysops

Flowers pale yellow; plants of Nevada.

D.

DD.

Flowers pale yellow, becoming tinged with redd-
ish=brown at maturity in some; leaves densely

and evenly tomentose on both surfaces, egl andul-
ar, the blade 4-8 mm long, 2-5 mm wide, obl anceo-
late to elliptic, the petiole 0.5-1 (1.5) mm long
and tomentose; involucres 5-7 per head, 2-2.5 mm
long; crusted mineralized sand, Sulphur Hot Spr-
ings, Ruby Valley, Elko Co., Nevada, at about
6050 feet elevation.........06. E. argophyllun

Flowers greenish-yellow to pale yellow, becoming
tinged with red at maturity, 3-3.5 mm long; lea-
ves densely tomentose below, less so and greenish
above, eglandular, the blade 5-10 (15) mm long,
(2) 3-6 mm wide, oblanceolate to elliptic, the
petiole 4-12 (15) mm long; involucres mostly 3-5
per head, 3-3.5 mm long; limestone slopes and
ridges, Ruby Mountains and East Humboldt Range,
Elko Co., Nevada, from 9000 to 10,400 feet elev-
EN OUS Cre ee slab Sue uae ious cats E. ringtt

AA. Flowers white to rose or red, not yellowish.

B.

Flowers glandular, not glabrous; plants above 10,000 feet
elevation.

C.

Leaves I=2 cm long, densely white-tomentose below,
less so and white above; scapes 3-8 cm long, glandu-

1972 Reveal, New species & combinations in Eriogonum 173

lar, not floccose; inflorescences containing 5-7 in-
volucres; pedicels glandular at the tip; granitic
sand and rocky outcrops, White Mountains, east-cent-
ral California and adjacent west-central Nevada, from
10,000 to 13,000 feet elevation.... E. gracilipes

CC. Leaves 3-10 mm long, densely white-tomentose and
greenish below, less so and greenish above; scapes up
to 3 cm long, floccose and glandular; inflorescences
containing 2-4 involucres; pedicels glabrous except
for a few scattered glands at the base; | imestone and
quartzite sand and rocky outcrops, Snake Range, east-
central Nevada, from 10,000 to 12,000 feet elevation.
on neta ie antes. ahi Ba, E. holmgrentt

BB. Flowers glabrous, white or cream; leaves mostly spatulate
and I-2 am long, densely white-tomentose on both surfaces;
scapes floccose; inflorescences containing 2-5 involucres;
clay slopes and hillsides, Grant and Powell cos., Montana,
southwestwardly into Lemhi, Clark and Custer cos., Idaho,
below 7000 feet elevation......-eeeeees E. mancwmn

Eriogonum tumulosum (Barneby) Reveal, stat. nov.

Eniogonum villiflonum A. Gray var. tumukosum Barneby, Leaf!l.
WB 8: l55incl9A9,

Low pulvinate cespitose herbaceous perennials with several
hundred rosettes of leaves, forming a rounded, hummock~1like mound
of matted caudex branches and leaves 1-4 dm across, the upper por-
tion of the caudex branches clothed with old persistent leaves
and petiole-bases, the lower portion bearing long strips of shag-
gy brown to reddish-brown bark, from a stout, woody gnarled tap-
root; Leaves clustered, persistent, forming tuft-like rosettes,
the leaf-blade oblanceolate to elliptic, 3-4 mm long, 0.7-1 mm
wide, white- or greenish- silky-tomentose on both surfaces, the
margin entire and plane to slightly thickened, the petiole short,
0.4-0.7 mm long, silky-villous, the petiole-base elongate, 1.2-
1.8 mm long, 0.8-1 mm wide, thinly villous without, glabrous with-
in; flowering Stems scapose, 1-9 mm long, villous, usually obscur-
ed by the leaves, erect; inflorescences capitate, the head 5-8 mm
across; bracts 4-5, semifoliaceous, linear-lanceolate to lanceo-
late, 2.5-3.5 mm long, 0.3-0.4 mm wide, villous without, glabrous
within, connate at the very base; peduncles lacking; involucres
solitary, campanulate, 2-4 mm long, (4) 5-8 mm wide, villous with-
out, glabrous within, the 7-10 lanceolate teeth 1.6-2.2 mm long,
the bractlets few, oblanceolate, 2-2.5 mm long, fringed with long
white cells, the pedicel short, 0.3-0.5 mm long, sparsely pubesc-
ent with scattered villous hairs; {2ower%s white with a greenish
or reddish margin, maturing rose or rustic, 3-4 mm long, densely
pilose without except for the upper part of the tepals, sparsely
pilose within, the tepals essentially similar; oblong to oblong~

17h Pi 2 PO) beOhGok A Vol s\23, now:

oblanceolate, 0.8-1 mm wide, united at least 1/2 the length of the
flower; Stamens slightly exserted, 3-4 mm long, the filament glab-
rous, the anther reddish-yellow, 0.60.8 mm long, oblong; achenes
light brown, glabrous, 1.8-2 mm long, the globose base tapering

to a short, 3-angled beak.

TYPE: Sandstone ledges and rockspavement on Red Plateau,
southwest of Woodside, Emery Co., Utah, at 5200 to 5300 feet ele-
vation, I3 June 1947, Ripkey & Barneby 8678. Holotype: CAS! Iso-
types: GH, NY, UTC!

DISTRIBUTION: Dry sandstone ledges and gravelly clay or
rocky places of eastern Utah in Duchesne and Emery cos. Flower=
ing from May to July.

The seemingly close relationship between Extogonum villiflor-
um and E. tumulosum is offset by their very different habit of
growth and geographical distribution. Unlike E. tumulosum which
forms large mounds of several hundred closely associated rosettes
of leaves with short, erect scapes topped with a capitate head of
involucres, E. vahliflonrum is a small, low, prostrate plant with
its ten to twenty loosely associated rosettes bearing a long,
prostrate flowering stem topped by a compact umbellate-cyme of
involucres. In the field, E. viklisloruwm is very easy to miss,
as in most cases it is difficult to distinguish from the array of
small rocks it so frequently grows with. Extogonum tumulosum, on
the other hand, is very obvious, and the very old, large mounds
of dead leaves and detritus often resemble miniature pack-rat
nests in the more open places among pinyon-juniper. Their geo-
graphical distribution is also distinct. Enxtogonun villiflorum
occurs from White Pine and Lincoln cos., Nevada eastward into
western Utah from Beaver and Sevier cos. south to (apparently)
Kane Co. where the type was collected but the plant not discover-
ed since. Extogonum tumulosum is found to the east of this ran-
ge, across the Wasatch Mountains - a physical barrier that reach-
es elevations of more than 10,000 feet. Its range is from the
benches around Duchesne eastward to near Myton in Duchesne Co.,
and then on the north end of the San Rafael Swells in Emery Co.
This type of distribution pattern is seen in other groups, not-
able in Adtragalus, Penstemon and Townsendia.

The three species which belong to this group can be distin-
guished as follows:

A. Flowers white, becoming rustic or red in age, 3-4.5 mm long;
involucres 6-8-lobed, 5-|2-f lowered.

B. Plants loosely cespitose at full development and composed
of 10-20 rosettes; flowering stems 2-5 (8) cm long, pro-
strate; inflorescences compact cymose-umbel late with sev-
eral evident rays; gravelly and clay soils of western
Utah in Beaver, Millard, Sevier and (apparently) Kane cos.

1972 Reveal, New species & combinations in Eriogonum 175

and in Lincoln and White Pine cos. of adjacent eastern
MONA ati «0cls.n siaieace th Miss otreivs was alee os E. uilliflorum

BB. Plants densely pulvinate and forming hemispheric mounds
up to 4 dm across at maturity, with several hundred ro-
settes; flowering stems |-9 mm long, erect; inflorescen-
ces capitate or essentially so; gravelly and clay soils
in eastern Utah from Duchesne Co. south to Emery Co.....
died Mstak Munsee p-0 pn ae mle an oA Se E. tunuloswm

AA. Flowers yellow, |.8-2.3 mm long; involucres 4-lobed, sol it-
ary, sessile in the rosettes, 3-4-flowered; plants densely
pulvinate and compact, 0.2-0.8 cm high; rare, endemic to red
calcareous gravel ridges, Paunsagunt Plateau in Red Canyon
and near Widtsoe, Garfield Co., Utah....... E. anretioides

Eriogonum contiguum (Reveal) Reveal, stat. nov.

Erxcdogonum inglatun Torr. & Frém. var. contiguum Reveal,
AU SOME ZZ 15) 970%

Erect herbaceous annuals (0.3) 0.5-3 dm high arising from a
slender, woody taproot; 2eaves basal, the leaf-blade mostly round
to round-oblong, (0.3) 0.5-1 (1.4) cm long and wide, hirsute on
both surfaces, greenish or yellowish, with longer and more numer-
ous hairs along the margin and midvein, usually slightly less
pubescent above than below, the margin entire and plane, the apex
round, the base usually round, occasionally slightly cordate, the
petiole hirsute, 0.4-1.5 cm long, the petiole-base triangular,
0.5-2 mm long, 0.5-1.5 mm wide, hirsute and minutely glandular
without, glabrous within; flowering Stems erect with |I-few stems,
slender, glandular at the base, otherwise glabrous, dark green,
up to 5 cm long; inflorescences densely paniculated-cymes, (0.2)
0.5-1.8 dm long, mostly trichotomously (occasionally more) bran-
ched at the first node, di- or trichotomous above, the secondary
branches lacking or few in number at the first node, the branches
slender and dark green, glandular at the nodes and along the low-
er portion of the branch, otherwise glabrous; bracts scale-like,
ternate, triangular, 0.3-1.4 mm long, glandular without, glabrous
within except for ciliated hairs at the apex and along the margin
of the connate bract, connate at the base; peduncles capillary, =
erect and straight, 0.3-1.2 (2) cm long, glandular at the node,
glabrous above, dark green to reddish; involucres turbinate, 1-1.3
mm long, 0.6-1 mm wide, glabrous within and without, the 5 (rare-
ly 4) acute teeth 0.2-0.4 mm long, the bractlet lacking, the ped-
icel 1-1.6 mm long, glabrous; {lowers golden-yellow with dark red
to reddish-brown midribs and bases, usually becoming reddish at
maturity, I1-1.3 (1.6) mm long, densely and conspicuously short-
hirsute without with coarse, whitish-yellow, curved hairs, glab-
rous within except for minute glands along the midribs, the tepals
essentially similar, lanceolate, united about 1/3 the length of
the flower; Stamens slightly exserted, 0.9-1.2 mm long, the fila-

176 P Heyer Or Leo Ger A&A Vol s"23,, "no, a

ments glabrous, the anther yellow, 0.3 mm long, oval; achenes
light brown, 1,5-1.8 (2) mm long, glabrous, the globose base tap-
ering to a long, 3-angled beak.

TYPE: Large playas southeast of the junction of Nevada 10
and U.S. Highway 95, east of Ash Meadows, T.I7S., R.5IE., sec.
21, at 2250 feet elevation, Nye Co., Nevada, 3 June 1969, Reveak
& Matthews 2157. Holotype: US! Isotypes: BRY, NY, UTC!

DISTRIBUTION: Dry desert flats and lower slopes in south-
western Nye Co., Nevada from Ash Meadows south to Pahrump Valley,
hence westward into eastern Inyo Co., California from north of
Death Valley Junction southward to near Tecopa. Flowering from
April to late June.

The elevation of this recently published variety of Eatogon-
un inffatum Torr. & Frem. to the species ranks comes after an
additional two years of field work. My early conclusion was bas-
ed on the supposed strength of the 5-tooth involucre which is so
typical of E. inflatum and the fact that annuals are a part of
the species (ie., var. fuscfowune of Utah and Colorado). Field
work in southern Nevada and adjacent California conducted in the
springs of 1970 and 1971 concentrated in those areas where E. in-
flatum, E. contiguum and E. trichopes Torr. could be found grow-
ing together. In these places, all three could be distinguished.
Naturally, the line of demarcation between E. sngélatum and E.
contiguum was easy to draw, but that between E. contiguum and E.
tnrichopes was much more difficult. For the most part, the number
of teeth on the involucre held, but as more and more plants were
studied, it became obvious that an occasional involucre on E.
contiguum would be four-toothed although the majority of involuc-
res on the same plant had five-teeth. The same held true for E.
trichopes, with a few plants too having four and five teothed in-
volucres on the same plant.

In the field studies, one aspect was confirmed. In general,
Exriogonum contiguum is glandular whereas E. trichopes is glabrous
throughout. Also, the branches of the first species are usually
reddish while those of the second species are usually yellowish-
green - a feature that can quickly separate the two entities in
the field even from a distance. It seems clear from the above
investigations, that it is impossible to hold E. contiguum within
E. 4nélatum, and the only recourse is to elevate it to the speci-
fic rank.

The differences between the various species associated with
Ercogonum inglatum can be separated as follows:

A. Involucres 5-lobed; plants annual or perennial with an open
inflorescence, the lower nodes with 3-5 branches, the upper
ones di- or trichotomous; flowers yellow, I-3 mm long; central

1972

AA.

Reveal, New species & combinations in Eriogonum 177

Baja California northward into the western United States.

B.

BB.

Plants glabrous and glaucous, annual or perennial, up to

10 dm high; involucres mostly I-I|.5 mm long; flowers |.5
-3 mm long; stems often inflated....... E. inflatum
Plants glandular, strictly annual, up to 3 dm high; in-
volucres I-1.3 mm long; flowers I-I|.3 (1.6) mm long;
SaeOES Hie LITT ATOR ses cate Gas see denies E. contiguum

Involucres 4-lobed; plants with verticillated whorls at each
node, sometimes densely so.

B.

BB.

Peduncles 3-15 mm long; leaves pilose to hirsute on both
surfaces.

C. Plants perennial, 2-10 (20) dm high with the inflor-
escence composed of verticillated cymes of 6-30 rad-
iating, rigid, secondary branchlets; leaves pilose

and crispate-undulate; central Baja California......
Bape Poi ata bane aga er ala ae & E. pilosum

CC. Plants annual, |1-4.5 (6) dm high with the infloresc-
ence composed of paniculated cymes with 5-20 radiat-
ing secondary branchlets at the lower nodes; leaves
hirsute and entire or merely crenulate; northern Baja
California and Sonora, Mexico, into the western Unit-
BU SHATAS a ie.« a's & ain ewe tie, we lalele hota «'s E. trichopes

Peduncles lacking or 1-4 mm long; leaves glabrous or
short-hirsute on both surfaces.

C. Plants perennial; peduncles !-4 mm long; flowers yel-
lowish but becoming reddish, densely hirsute without;
leaves glabrous, ciliated; involucres |.2-1.7 mm
long; west-central Baja California. E. scakane

CC. Plants annual; peduncles lacking; flowers white to
pink, becoming reddish, strigillose without; leaves
short hirsute, not ciliate; involucres 0.4-0.7 mm
long; Cedros Island and Vizcaino Desert, Baja Calif-
CEN lay. ab teks ALR ae eee E. intricatwn

ACKNOWLEDGMENTS

This paper was originally submitted to Dr. Lloyd Shinners
for consideration in SIDA. His untimely death delayed publicat-
ion to a point that the new names would not appear until 1973,
too late for their inclusion in the forth coming volume of the
INTERMOUNTAIN FLORA: THE VASCULAR PLANTS OF THE INTERMOUNTAIN
WEST by Cronquist, Holmgren, Holmgren and Reveal. | am grateful
to Dr. Shinners for reviewing the original manuscript and making
comments on it.

Field studies were support by National Science Foundation

178 P i, WeTeONOrGrd A Vol. 23, no. 1

grants to Arthur Cronquist of the New York Botanical Garden for
the Intermountain Flora Project for all species except the last.
Field work for studies in Nevada and California during the spring
seasons of 1970 and 1971 were fully supported by Contract No. AT
(04-1) Gen-12 between the University of California and the Divis-
ion of Biology and Medicine, United States Atomic Energy Commiss-
ion; | wish to take this opportunity to thank Dr. Janice C. Beat-
ley, Associate Research Ecologist, for her aid with this work.
Additional herbarium and field studies conducted in I97I and 1972
were largely supported by my National Science Foundation Grant,
GB-22645 for studies on Extogonum and the Intermountain Flora.

Lastly | wish to acknowledge the aid of Drs. Noel and Patri-
cia Holmgren of the New York Botanical Gardern for gathering mat-
erial for me in 1969 and 1970, and Mr. Conrad V. Morton of the
Smithsonian Institution for assistance with the Latin descriptions
and diagnoses.

LITERATURE CITED

BARNEBY, R.C. 1949. Eriogonum villiflorum and its near relatives
in the Great Basin. Leagl. W. Bot. 5: 151-154.

CRONQUIST, A., A.H. HOLMGREN, N.H. HOLMGREN & J. L. REVEAL. In
press. INTERMOUNTAIN FLORA: THE VASCULAR PLANTS OF THE
INTERMOUNTAIN WEST. Volume 1. Inttoduction and Vascular
Crytogams. New York.

JONES, M.E. 1903. Eriogonum. Contr. W. Bot. 11: 4-18.

REVEAL, J.L. In press. Two new species of Eriogonum (Pol ygonac-
eae) from California and adjacent states. Akiso.

PARMELIA JAMESII, AN UNUSUAL SPECIES IN SECTION IMBRICARIA
(LICHENES) FROM AUSTRALIA AND NEW ZEALAND

Mason E. Hale, Jr.
Smithsonian Institution, Washington, D.C. 20560

Parmelia jamesii, sp. nov.

Thallus adnatus, corticola, mollis, cinereo-albus, 5-10 cm
latus, lobis sublinearibus, 1.5-3.0 mm latis, margine sparse cili-
atis, superne planus, nitidus vel sparse opacus pruinosusque,
aetate rugosus, apicem versus leviter albo-reticulatus, modice
isidiatus, isidiis cylindricis, tenuibus, erectis, usque ad 0.3 mm
altis, subtus niger, modice rhizinosus, rhizinis simplicibus vel
pauce squarrose furcatis. Apothecia ignota.

Chemistry: Atranorin in the cortex, fumarprotocetraric acid
and protocetraric acid (trace) in the medulla (identified with thin-
layer chromatography).

Holotype: On Nothofagus by the coast, Wellington, North Island,
New Zealand, collected by P. W. James, NZ2118, January 1963 (US;
isotypes in BM, TNS).

Additional specimen examined: South side of Monga Mountain,
Braidwood Distr., N.S.W., Australia, Weber & McVean L-51532 (COLO,
US).

The distinguishing features of this species are the isidia,
unusual development of rhizines, and chemistry, this being one of
the very few species in subgenus Parmelia with fumarprotocetraric
acid. Superficially the species seems to belong in section Hypo-
trachyna because of the sublinear narrow lobes, much as in P. im-
bricatula, but the rhizines are not dichotomously branched. While
the rhizines are mostly simple, some have distinct short side
branches typical of squarrosely branched species in section Parme-
lia, where, in fact, I had tentatively placed the species, in spite
of the fairly obvious marginal cilia. I concluded that P. jamesii
is best assigned to section Imbricaria on the basis of the fine
structure of the surface of the cortex. There are numerous pores
at X500 under the scanning electron microscope such as one would
expect to find in section Imbricaria. No pores are produced in
section Parmelia.

ey

NOVELTIES FROM BRAZIL, PERU, VENEZUELA, AND THAILAND

Harold N. Moldenke

CITHAREXYLUM FLEXUOSUM var. GLABERRIMUM Moldenke, var. nov.

Haec varietas a forma typica speciei foliis utrinque glaber-
rimis nitidisque differt.

This variety differs from the typical form of the species in
having both surfaces of its leaf-blades completely glabrous and
shiny.

The type of the variety was collected by C. Earle Smith, Jr.,
and Jacinto Blas (no. 4905) along the trail to the cave across
the Rio Santo from Mancos, Callejon de Huaylas, Ancash, Peru, on
April 9, 1970, and is deposited in the United States National
Herbarium at Washington. The collectors describe the plant as a
closely branched shrub to 2 meters tall, the fruit being shiny
and black.

CLERODENDRUM SCHMIDTII var. MACROPHYLLUM Moldenke, var. nov.

Haec varietas a forma typica speciei laminis foliorum 25—28
em. longis 13--1).5 cm. latis et petiolis 8--13 cm. longis re-
cedit.

This variety differs from the typical form of the species in
having its petioles 8--13 cm. long and the leaf-blades 25--28
em. long and 13--1).5 cm. wide,

The type of the variety was collected by Ch. Charoenphol, Kai
Larsen, and E, Warncke (no. 5022) at Ko Rang Yai, 20 km. south
of Ko Chang, in southeastern Thailand, 1198' N., 102°23' E., on
November 19, 1970, and is deposited in the Herbarium Jutlandicum
of the Botanisk Institut, Aarhus Universitet, Aarhus, Denmark.
The collectors describe the calyx as "whitish green with a little
red" and the corollas as white.

LANTANA ARMATA var. VELUTINA Moldenke, var. nov.

Haec varietas a forma typica speciei laminis foliorum subtus
petiolisque pedunculisque ramulisque sarmentisque ubique densis—
sime breviterque velutinoso—pubescentibus recedit.

This variety differs from the typical form of the species in
the lower surface of its leaf-blades, its petioles, its pedunc-—
les, branchlets, and twigs very densely velutinous—pubescent
with very short and soft hairs, not at all hirsute nor hispidu-
lous.

The type of the variety was collected by Santiago Lépez—
Palacios (no. 2616) at El Rincén, near Urefia, Tachira, Venezuela,
on December 8, 1971, and is deposited in my personal herbarium
at Plainfield, New Jersey. The collector notes "Arbusto aparen-
temente inerme, hojas ovales a oval-lanceoladas, membran4ceas, un
poco 4speras por la haz cuando secas, velutinosas por el envés.
Pedtinculos por lo general menores que las hojas. Brdcteas pequefi-

180

1972 Moldenke, Novelties 181

as, lineal lanceoladas. Flores amarillas que cambian a anaran-
jado."

PAEPALANTHUS DICHOTOMUS var. BRASILIENSIS Moldenke, var. nov.

Haec varietas a forma typica speciei foliis 3--5 mm. longis 1
mm. latis glabris nitidisque ad apicem acutissimis et pedunculis
abbreviatis };--10 mm. longis recedit.

This variety differs from the typical form of the species in
having its rosulate leaves only 3—5 mm. long and 1 m, wide,
thick in texture, stiffly erect, glabrous and shiny, very acute
at the apex, and the peduncles much abbreviated, only )--10 m.
long. —

The type of the variety was collected by Gert Hatschbach (no.
27425) along the road near Salto da Divisa, in the municipality

of Diamantina, Minas Gerais, Brazil, at 1300 meters altitude, on
September 7, 1971, and is deposited in my personal herbarium at
Plainfield, New Jersey.

ADDITIONAL NOTES ON THE GENUS VERBENA. X

Harold N. Moldenke

VERBENA [Dorst.] L.

Additional & emended bibliography: Gattinger, Med. Pl. Tenn.
63--6,. 1894; Reiche & Phil., Fl. Chil. 5: 271-298, 300, 302,
30h, & 463. 1910; Graves, Eames, Bissell, Andrews, Harger, &
Weatherby, Bull. Conn. Geol. & Nat. Hist. Surv. : (Cat. Flow.
Pl.) 330--332. 1910; Harger, Bull. Conn. Geol. & Nat. Hist. Surv.
48: 74. 1930; Gathercoal, Checklist Nat. & Introd. Drug Pl. [22].
1942; Beston, Herbs & Earth 60--62, 131, & 133. 1950; Goossens,
Suid-Afrik. Blom. Pl. 185. 1953; Fogg, Weeds Lamm & Gard. 11. 1956;
Haramaki, Weed Abstr. 1\: 256. 1965; Ferrdndiz, Guia Med. Veg. 388-
389. 1967; Tawada, Okinawa Seibutsugakki [Biol. Mag. Okinawa] ) (6):
36. 1968; McCarthy & Morrow, Proc. 2th N. Cent. Weed Control Conf.
64. 1969; Soukup, Raymondiana 3: 26, 57, & 91. 1970; Graf, Exot. Pl.
Man., ed. 1, 320, 378, 10, & 411. 1970; Bright of America, Summers-
ville, W. Va., 25.GC.70 Petunias & Verbenas. 1970; Mayall, Weed
Abstr. 20: 321. 1971; Moldenke, Fifth Summ. 1: 6, 10, 1i—25, 27,
30--5,, 57—66, 7h--78, 81, 82, 84--86, 89, 91, 92, 98, 99, 101,
101-106, 109, 111, 120, 128, 137, 138, 143, 14h, 177—179, 18h,
187--190, 192—19, 200--211, 213--215, 222, 231, 23h, 238, 2l1,
246, 248, 252, 255, 257, 262, 264—267, 269-271, 278, 279, 281,
284, 290, 292, 29h, 298, 303, 306, 308, 311--313, 318, 328, 337,
341, 343, 34h, 349--353, 369--373, 375, 376, 389--391, 396, 397,
4OO, 402, 421, 468, h71, & 473 (1971) and 2: 491, 492, 520--523,
525527, 531, 534, 548, 558, 559, 568, 569, 575, 593, 59h, 618—
621, 645, 648--710, 736, 738, 741, 72, 7hh, 752, 766, 767, 769,

182 PHY DyOrhOoe: bs Vol. 23, n004.

773, 77h, 776, 781--785, 787, 780, 791--79, 912--922, 967-—-970,
& 973. 1971; Moldenke, Phytologia 22: 56--501. 506--508, 510, &
B12. 19726

VERBENA AMBROSIFOLIA Rydb.
Additional bibliography: Moldenke, Phytologia 22: 59-62,
71, 473, 485, & 497. 1972.

VERBENA ARAUCANA R. A. Phil.
Additional & emended bibliography: Reiche & Phil., Fl. Chil. 5:
289 & 290. 1910; Moldenke, Phytologia 22: 63 & 6h. 1972.

VERBENA ARISTIGERA S. Moore

Additional bibliography: Moldenke, Phytologia 22: )63—l6).
1972.

The corollas of this plant are described as "lilac" on Krapo-
vickas & Cristobal 16015. har:

Additional citations: ARGENTINA: Misiones: Krapovickas & Cris-
tébal 16015 (Ft). ees

VERBENA ATACAMENSIS Reiche
Additional & emended bibliography: Reiche & Phil., Fl. Chil. 5:
289 & 291--292. 1910; Moldenke, Phytologia 22: 6. 1972.

VERBENA BERTERII (Meisn.) Schau.
Additional bibliography: Reiche & Phil., Fl. Chil. 5: 287, 289,
293, & 294. 1910; Moldenke, Phytologia 22: 66. 1972.

VERBENA BONARIENSIS L.
Additional & emended bibliography: Reiche & Phil., Fl. Chil. 5:
283 & 284. 1910; Moldenke, Phytologia 22: 474-79. 1972.

VERBENA BRASILIENSIS Vell.

Additional bibliography: Rickett, Wild Fls. U. S. 3 (2): [367],
pl. 111 (1969) and 5 (2): [455], pl. 152. 1971; Moldenke, Phytolo-
gia 22: 478 & 488--\90. 1972.

Additional illustrations: Rickett, Wild Fls. U. S. 3 (2): [367],
pl. 111 (in color) [as V. litoralis] (1969) and 5 (2): [55], pl.
152 (in color) [as V. litoralis]. 1971.

Rickett (1969, 1971), in the color illustration which he uses
twice, purports to depict V. bonariensis L. and V. litoralis
H.B.K. -- the front plant in the photograph is obviously V. bonar-
iensis, but the rear plant is certainly not V. litoralis! A far
more likely identification for it is V. brasiliensis.

VERBENA CABRERAE Moldenke

Additional synonymy: Glandularia cabrerae (Mold.) Tronc., in
herb.

Additional bibliography: Moldenke, Phytologia 22: 90. 1972.

Krapovickas and his associates found this plant growing on
"barrancas de arroyo" and describe the corollas as "violet".

1972 Moldenke, Notes on Verbena 183

Additional citations: BOLIVIA: Tarija: Krapovickas, Mroginski,
& Fern4ndez 19272 (Ft).

VERBENA CANADENSIS (L.) Britton

Additional & emended bivliography: Graves, Eames, Bissell, An-
drews, Harger, & Weatherby, Bull. Conn, Geol. & Nat. Hist. Surv.
ly: [Cat. Flow. Pl.] 331-332. 1910; Winge, Proc. Linn. Soc. Lond.
150: 236. 1938; Moldenke, Phytologia 22: 92-98. 1972.

VERBENA CAROLINA L.

Additional synonymy: Verbena caroliniana Anderss. apud B. L.
Robinson, Proc. Am. Acad. 38: 197, in syn. 1902.

Additional & emended bibliography: Crantz, Inst. Rei Herb. 1:
573. 1766; [Retz.], Nom. Bot. 11. 1772; J. F. Gmel. in L., Syst.
Meus, OC. 13, pr. 1, 2: 2 (1789) and ed. 13, prs 2, 2: k2. 1796;
Pers., Sp. Pl. 3: 317. 1819; Hook., Trans. Linn. Soc. Lond. Bot.
20: 195. 1847; A. Wood, Class-book, [ed. 42], pr. 1, 537 (1861)
[ed. 2], pr. 2, 537 (1863), [ed. 2], pr. 3, 537 (1865), [ed. 2],
pr. 4, 537 (1867), [ed. 42}, pr. 5, 537 (1868), [ed. 2], pr. 6,
537 (1869), and [ed. 42], pr. 7, 537. 1870; A. Wood, Am. Bot. &
Flor., ed. 1, pr. 1, 236 (1870), ed. 1, pr. 2, 236 (1871), and ed.
1, pr. 3, 236. 1872; A. Wood, Class—book, [ed. 2], pr. 8, 537.
1872; A. Wood, Am. Bot. & Flor., ed. 1, pr. h, 236 (1873), ed. 1,
pr. 5, 236 (1874), and ed. 1, pr. 6, 236. 1875; A. Wood, Class-
book, [ed. 42], pr. 9, 537 (1876) and [ed. 2], pr. 10, 537. 1881;
O. R. Willis in A. Wood, Am. Bot. & Flor., ed. 2, 236. 1889;
Solered., Syst. Anat. Dicot. 713. 1899; B. L. Robinson, Proc. Am.
Acad. 38: 196--197. 1902; D. H. Scott in Solered., Syst. Anat.
Dicot., transl. Boodle & Fritsch, 1: 631. 1908; Lowe, Miss. State
Geol. Surv. Bull. 17: 236. 1921; Savage, Cat. Linn. Herb. Lond.
4. 19115; Howell & McClintock in Kearney & Peebles, Ariz. Fl., ed.
2, 726 & 727. 1960; J. F. Macbr., Field Mus. Publ. Bot. 13 (5):
615, 618, & 62h. 1960; Hocking, Excerpt. Bot. A.6: 91 (1963) and
A.7: 206 & 55. 1964; Rzedowski & McVaugh, Contrib. Univ. Mich.
Herb. 9: 65 & 107. 1966; W. C. Grim, Recog. Flow. Wild Pl. 228 &
229. 1968; Moldenke, Biol. Abstr. 9: 3252. 1968; Moldenke, Phy-
tologia 16: 186 & 30--31. 1968; Moldenke, Résumé Suppl. 16: 2
& 28. 1968; Hocking, Excerpt. Bot. A.13: 570 & 571 (1968) and A.
1h: 206. 1969; Moldenke, Biol. Abstr. 50: 418. 1969; D'Arcy, Tax-
on 19: 556. 1970; Gibson, Fieldiana Bot. 2) (9): 230--232, fig.
46. 1970; Wiggins & Porter, Fl. Galdp. Isls. 997. 1971; Moldenke
in Wiggins & Porter, Fl. Gal4p. Isls. 508. 1971; Moldenke, Fifth
Summ. 1: 51, 63, 75, 81, 8h, 85, & 370 (1971) and 2: 654, 660,
661, 673, 681, 690, 701, 706, 709, 787, 793, 913, & 973. 1971;
Moldenke, Phytologia 22: 501--502. 1972.

Additional illustrations: Gibson, Fieldiana Bot. 2h (9): 231,
fig. 46. 1970.

Recent collectors have found this plant growing along streams,
along paths in open disturbed secondgrowth thickets, and along
dry open or dry grassy roadsides, on railroad embankments and
grassy valley floors, in matorral, deciduous tropical woods, dense

18) P Hove Te, BeOrG?e DIA Vol. 23, HOsvs

forests, small barrancas, maize fields, disturbed oak woods, pine-
woods, pine-fir woods, and pine-oak woods, often in rocky soil.

It has been collected in fruit in March (in addition to the months
previously reported). Additional common names reported for it

are "chichavac" and "dori", Pennington reports that it is an ex-
cellent pasturage plant and that a medicinal tea is made from it
in Sonora to cure headaches.

The corollas are described as "lilac" on Flores Crespo 293,
"violet" on Franco R. s.n. [20.VIII.1967], "pale-violet" on Roe,
Roe, & Mori 678 8, "blue" on Breedlove 1))30a, Jiménez Re able “arid
Ss. Sieisoee z 89, “Wlight-blue" on Rowell 3061, "pale-vlue" on Flores
Crespo 45 and Breedlove & Kawahara 16887, and "purple" on Bautista
S.n. [13/VII1/1967], Farias 83, Martinez-Calderén 1765, R. Runyon
88h, and J. Rzedowski 1135, 1226, 2028), & 20988.

~ Breedlove & Kawahara found th ae plant growing | on the wooded
banks of streams with Alnus sp., Quercus spp., Padus serotina, and
Pinus engelmanni. In Guanajuato McVaugh found it "on steep rocky
(rhyolitic) mountainsides, weedy, not much seen", but in Tamauli-
pas it is reported by Runyon as "common on tops of mountains in
fertile soils". Rzédowski encountered it in "charcos cercanos a
la carretera", Gonz4lez Quintero in "ladera caliza con vegetacién
de Cephalocereus", and Jiménez R. on "ladera con vegetacién de
matorral xeréfilo",

Gibson (1970) states that "This has been reported from Guate-
mala as V. ehrenbergiana Schauer". Macbride (1960) cites Pennell
14770 from Lima and Soukup 3182 from Junin, Peru, but these are
probably misidentifications. The "Verbena Carolina L." of Lowe
(1921) is Stylodon carneus (Medic.) Moldenke, as is also the
"Verbena caroliniana, Carolina verbena" Boseieed by Grimm (1968).

~ Savage ( (1915) Tee as the type of V. carolina sheet 17 under
genus 35, Verbena, where it is identified as 'Verbena "carolin."!
with the comment "[cf. Jacq. list 1761. n. 8. det. Jacq: an
carolina]",

Franco R. s.n. [20.VIII.1967], Garcia Romero s.n. (Tie vires
1968], and Y M4rquez sn. [23/VII/1962] are mixtures 2s with V. men-
thaefolia Benth., while Pennington 215 is a mixture with V Ve pi pine-
torum Moldenke.

Charles Darwin, in the beginning of October, 1835, on the
famous voyage of the "Beagle", collected two specimens of vervain
on James Island in the Galapagos group. One of these was identi-
fied by Hooker on the label as "Verbena polystachya H.B.K. var.?"
Later (1847) he published this as "V. polystachya H.B.K. var.
foliis incisis segmentis grossé serratis" and made this comment:
"Sprengel unites this species with V. urticaefolia, to which our
plant bears a close affinity; but the margins of the leaves in the
present are far more deeply cut, and the segments again coarsely
serrated. They may very possibly be mere variations of one spe-
cies." Andersson (1859) misquoted Hooker's varietal description

1972 Moldenke, Notes on Verbena 185

and reduced it to "V. caroliniana Linn." Robinson (1902) and
Stewart (1911) adopted the name, V. carolina L., for the plant of
Darwin. There is no evidence that any of these later workers ac-
tually examined Darwin's specimen, although an unknowm hand
changed the determination on it to "V. urticifolia L. ?" and then
to "V. carolina L." Examination of the original specimen has
shown that it represents neither of the suggested species, but is
actually V. sedula var, darwinii Moldenke.

The Hodge 6237, he bw as Ve carolina, is actually V.
Tyson, Dwyer, & Blum 4300 are V V. litoralis He B.K.

Additional citations: ARIZONA: Santa Cruz Co.: Mason, Drouet,
MacEwan, & Price 1808 (Mi). MEXICO: Chiapas: Breedlove *1h30a
(Mi). Federal District: Aguirre Z. 121 (Ip); Carrillo 2 (Ip);
J. Espinosa s.n. [X.1954] (Ip); Farfas | R. 89 (Ip); Gutiérrez &
Molina s.n. [1.X.1967] (Ip); Islas F. s.n. [23/111/1962] (Ip);
Jiménez R. 151 (Ip); S. Lépez 89 (1 89 (Mi) ; M4rquez s.n. [23/VII/
1962], in part (Ip); Rojano 18 (Ip), 83 (Ip); J. Rzedowski 1135
(Ip), 1226 (Ip). Guanajuato: R. McVaugh 2206 (Mi). Guerrero:
Rowell 3061 (Mi). Hidalgo: Gonz4lez Quintero 1207 (Ip). Jalis-
co: Diaz Lu Luna 209 (Mi); Harker & Mellowes 35 (Ip, Ip, Mi). México:
Armenta 28 8B (Ac); | Franco R. s.n Sen. [20.VI1I .1967], in part (Ip);
Gonzdlez Quintero 1178 (Ip); Pineda R. 359 (Rf); Ryesky 101 (Ws),
102 (Ws); J. Rzedowski 2028) (RE); | Salinas M. 85 (Ac); Villegas
D. D. 76 (Ip). Michoacdn: Bautista Sone [13/VII1/1967] (Ip); Garcfa
Romero sen. [11.VIII.1968], in part rt (Ip); Hinton 13022 Sober
1748). Morelos: Flores Crespo 45 (Rf), 293 (Ac); Guerrero O.
a.n. [14/IX/1962] (Ip); H. Hern4ndez ~ Hern4ndez Sa, [12/VII/1965] (Ac).
Nuevo Leén: Barkley & Alanfs s 1012 (1 (Go, R: Rf). Oaxaca: J. Rzedow-
ski 20988 (Ac); C. L. Smith 22h (I (Ip). Puebla: Gonz4lez Quintero
180 (Ip). Sinaloa: Breedlove & Kawahara 16887 (Rf). Sonora:
Pennington 215 215, in part (Au--26),202). Tamaulipas: R. Runyon 88)
(Au—26820)). Veracruz: Martinez Calderén 1765 (Rf). GUATEMALA:
Quezaltenango: Roe, Roe, & Mori 678 (Rf). HONDURAS: Copén: F. A.
Barkley 0267 (RF). EL SALVADOR: San Salvador: C. Garcia 135
(Ca—128))17) .

VERBENA CAROLINA f. ALBIFLORA Moldenke

Additional bibliography: Hocking, Excerpt. Bot. A.13: 570 &
571. 1968; Moldenke, Résumé Suppl. 16: 2. 1968; Moldenke, Phyto-
logia 16: 88. 1968 ; "Moldenke, Biol. Abstr. 9: "3052, 1968; Mol-
denke, Fifth Sum, 1: 75, 81, 84, & 85 (1971) and 2: 67h, "688, &
913. i971.

Recent collectors have encountered this plant in villages and
at the edge of deciduous woods, flowering and fruiting in May.
Material has been misidentified and distributed in some herbaria

186 P HYP. O L0G DA Vol. 23, no. 1

as V. officinalis L.
Additional citations: MEXICO: Veracruz: Gutiérrez R. 342 (Rf).
GUATEMALA: El Quiché: E. Contreras 5247 (W--2558708) .

VERBENA CATHARINAE Moldenke

Synonymy: Verbena catharinensis Moldenke, Fifth Summ. 2: 661,
in syn. 19 [is

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 571.
1965; Moldenke, Phytologia 11: 51--52. 1965; Moldenke, Fifth
Summ. 1: 177 (1971) and 2: 661 & 913. 1971.

Hatschbach describes this plant as creeping and rooting, the
corollas violet or intensely lilac in color, blooming in October,
Additional citations: BRAZIL: Paran4: Hatschbach 1968 (Mi),

226) (Mi, N). iy) LS Sea

VERBENA CHACENSIS Moldenke
Additional bibliography: Moldenke, Phytologia 10: 101. 196);
Moldenke, Fifth Summ. 1: 187 (1971) and 2: 913. 1971.

VERBENA CHEITMANIANA Moldenke ;

Synonymy: Glandularia cheitmaniana Schnack & Rubens, Bol. Soc.
Argent. Bot. 13: 205, hyponym. 1970.

Additional bibliography: Moldenke, Phytologia 13: 188. 1966;
Schnack & Rubens, Bol. Soc. Argent. Bot. 13: 205. 1970; Moldenke,
Fifth Summ. 1: 200 & 370 (1971) and 2: 913. 1971.

The Ritchie specimen cited below is a mixture with V. rigida
Spreng. It is accompanied by a longhand label which I am unable
to decipher, but which appears to be only a diagnostic descrip-
tion of the plants. There does not seem to be any indication
that the specimens were taken from wild plants, so I assume that
they were cultivated where collected.

Additional citations: CULTIVATED: India: C. Ritchie ii in
part (Ed). ‘F a

VERBENA CHILENSIS Moldenke, Known Geogr. Distrib. Verbenac., ed.
By (ops Be) ee
Additional bibliography: Moldenke, Phytologia 16: 88. 1968;
Moldenke, Fifth Summ, 1: 200 (1971) and 2: 671, 776, & 913. 1971.
Bartlett found this plant growing on a mountain ridge above
the barrel cactus zone.
Additional citations: ARGENTINA: Mendoza: H. H. Bartlett 1910

(Au--195085, N). Neuquen: O'Donell 2008 (N).

VERBENA CILIATA Benth,

Additional synonymy: Verbena cilliata Benth. ex Moldenke, Ré-
sumé Suppl. 18: 14, in syn. 1969. Verbina ciliata Benth. ex Mol-
denke, Fifth Summ. 2: 708, in syn. 1971.

Additional & emended bibliography: Rydb., Fl. Rocky Mtns., ed.
2, pr. 1, 739 & 7hO (1922) and ed. 2, pr. 2, 739 & 7hO. 195h;
Howell & McClintock in Kearney & Peebles, Ariz. Fl., ed. 2, 725&
727. 1960; Martin & Barkley, Seed Ident. Man. 19). 1961; Turrill

1972 Moldenke, Notes on Verbena 187

in Curtis, Bot. Mag. 17: pl. 409. 1963; Hocking, Excerpt. Bot.
A.6: 91. 1963; Marroquin, Cuad. Inst. Invest. Cient. 1: 30 & 56.
1968; Moldenke, Phytologia 16: 186 & 196. 1968; Moldenke, Résumé
Suppl. ere (1968) and 18: 1). 1969; Gilberson & McHenry, Univ.
Et Agr. Exp. Sta. Tech. Bull. 186: 7 & 39. 1969; Hafez & You-

» Physiol. Pl. 22: 332. 1969; Kapp, How to Know Pollen 10h,
138° 231, & 249, fig. 208 & 276. 1969; Rydb., Fl. Rocky Mtns., ed.
25: Pleo 3, 739 & "70. 1969; Rickett, Wild Fls. U. S. 3 (2): [361]
& ” 362, pl. 109 (1969) and yy: 539, (sh), & 799, pl. 176. 1970;
Jarrett, Ariz. Highways 7 (8): (11) & 38. 1971; Moldenke in Cor
rell & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex. Res. Found.
Bot. 6:] 1317 & 1328, 1970; Correll & Johnston, Man. Vasc. Pl.
Tex. [Contrib. Tex. Res. Wound? Bot. 6:] 1799, "1876, & 1877. 1970;
Moldenke, Fifth Sumn, 1: 39, 42, 53, 58, 61, 63, 6s; 75,16; O23
& 370 (1971) and 2: 659, 663, 667, 708, 767, & 913. 1971; Molden
ke, Phytologia 22: 61, 62, 473, & 97. 1972.

"additional illustrations: Rickett, Wild Fls. U. S. 3 (2): [361],
pl. 109 (in color) (1969) and h: [sul], pl. 176 (in color). 1970;
Jarrett, Ariz. Highways 47 (8): [11] (in color). 1971.

The seeds of this plant are said by Martin & Barkley (1961) to
be compressed-oblong and slightly rounded-triangular in cross-
section, making them somewhat 2-sided, the rounded back bearing
lengthwise ridges toward the lower end and a network of cross-
ridges below, the inner face of the two planes meeting in a low
ridge, the surface generally covered with whitish papillae, the
margin bordered by a narrow flange, the attachment-scar whitish
and surrounded by a cuplike extension, the entire seed 1.5——-3 m.
long, with no endosperm,

Recent collectors have found this species growing on dry
plains and mesas, dry desert plains, flat dry plains, and low
banks near water, among large rocks, along dry roadsides, in xero-
phytic matorral, "pastizales", "zacatal", matorral with Opuntia,
maize fields, and Quercus-Cupressus woods, in openings in pine
forests, subalpine meadows, Prosopis-Larrea scrub, dry level
rocky grasslands, open stony roadside pastures, and matorral
roadsides, in sandy, volcanic, or rocky limestone soil. In ad-
dition to the months previously recorded, it has been found in
anthesis in Jamary.

The Andersons found V. ciliata "locally common in small barran-
cas with vegetation of shrubs, cacti, and a few small trees, with
limestone rocks, in Puebla and among shrubby vegetation in arro-
yos, abundant by roads, in Guanajuato; Garcfa Saucedo found it in
xerophilous matorral with Opuntia and Agave in Hidalgo; while
Stuessy encountered only "one clump about out l) feet in diameter in
Acacia grassland in Chihuahua. Cruz Cisneros found it on "ladera
andes{tica con vegetacién de pastizal con Quercus, Opuntia y Leg-
uminosae", "ladera andes{tica con vegetacién xeréfita", and "la-
dera basAltica con vegetacién de matorral de Acacia y Opuntia".

The corollas are described as having been "lilac" on Asteinza

sn. (5.VII.1966], Iffiguez 7), and Villegas D. 388, "blue" on Pi-

188 P Heels LeOrGr vk Vol. 23, now 2d

neda R. 38, "violet" on Latotre s.n. [12 May 1968], "lavender-
purple" cmDietents 35753 "lavender" on Anderson & Anderson 5048
and Anderson & Laskowski 3548, "blue-purple" on Stuessy peu
"reddish" on McGregor 16626, "pinkish-purple" on Vilas 39, "red-
dish-violet" on Vilas 29, 29, "lavender, turning rose-pink in press"
on Anderson & Anderson 531k, and imum on Dominguez & McCart
8270, Galicia a; Garcia Saucedo 2607, Hidalgo & Anda s.n, [27.IV.
1967], Nevling & Gémez—Pompa 1075, Po Powell, Scudday, & Sikes 13h,
J. Rzedowski 3051, 3428, 20301, 1, 20543, 22205a, & 2 & 23895, and Ven-
tura A. 1545.

Gilberson & McHenry (1969) report that this plant is attacked
by both the pycnia and aecia stages of the fungus, Puccinia
aristidae Tracy, in Yavapai County, Arizona. Marroquin (1968)
cites Alanfs 12), 258, & 278 (FCB 290, 291, & 2492) and Guti-
érrez Lobatos ca (FCB 1353) from Nuevo Leén, Mexico.

~~ Mutis deploy “cited below, bears a printed aber reading "PLANTS
OF COLOMBIA", but the Etim was undoubtedly collected in Mexi-
co. J. Rzedowski 22205a is a mixture with V. menthaefolia Benth.
The color illustration purporting to represent V. ciliata in the
Rickett publications (1969, 1970) seems definitely to be not this
species.

faterial of V. ciliata has been misidentified and distributed
in herbaria under the names V. ambrosaefolia Rydb. and V. micro-
phylla H.B.K. On the other hand, the J. H. Ehlers 8367, distribu-
ted as V. ciliata, is actually ve ambrosifolia ja Rydb.; D Demaree
41126, Hinton & at ee and Mears & Mears ae are Ve anbrosi-

= fet, SS

Lag. & Rodr.; Ramirez & CArdenas 15) is Ve ciliata var. longiden-
tata Perry; Beasley & Finzel 851 is Vv. ciliata var. pubera
(Greene) Perry; Atwood 1730, Cue Le Hitchcock 2550 & 2561), and
Munz, Johnson, & Harwood Teh a are ev. gooddingii Briq.; ; Pinkava,
Lewis, “Noble, & . Lehto 112h9 aS) Wis gooddingii var. nepetifolia
Tidestr.; Pr Pruitt j 210: and Be B. C. Tharp 49-1118 are V. pumila Rydb.;
and Barkley, Pax Paxson, & Webster 2493, Rebolledo Vélez sen. [20.
VIII .1967], and J. | Je Soule nrsL al 1009 a are V. teucriifolia Mart. & Gal.
Additional citations: MEXICO: Chihuahua: Stuessy 959 (Au--
257767), 965 (Au--257701), 101) (Au--257670). Coahuila: Johnson
& Johnson 1649 (Ws) ; Latotre s.n. [12 May 1968] (Au--2650)5);
Powell, Scudday, & Sikes 113 l) (Au--257010). Durango: Anderson &
Taskowsicl 358 (Mi); Joh Johnson & Johnson 1712 (Ws). Federal Dis-_
trict: Bopp p 21h (Ip). Guanajuato: Anderson & Anderson 50,8 (Mi).
Hidalgo: : Garcfa Saucedo 2607 (Rf); Gonzdlez Quintero 3L9 (A (Au-—
21298); J. J. Rzedowski 16927 (Ip), 20543 (Ac), 23895 (RE); S. Sdn-
chez 37 (Ac). Jalisco: J. V. A. Dieterle 3575 (Mi) (Mi); R. L. Mc

1972 Moldenke, Notes on Verbena 189

Gregor 16626 (N). México: Asteinza s.n. [5.VII.1966] (Ip); Cruz
Cisneros 99 (Mi), 52h (Ip), 562 (Mi); E E. R. Garcfa 60 (Rf); Hi- Hi-
dalgo y Anda Bene’ (2Te 27-1V.1967) ] (Ip); Iffiguez 7 (Ac); | Me Padilla
117 (Ac); Pi Pineda R. 348 (Rf); Rebolledo do Vélez s SMe (20 .VIII .1967]
(Ip); Je Rzedowski 15678 (Ip), 20301 (Ip), 22 22205a, in part (Ip),
22206a (Ip); Vargas N. 9 (Rf); "Villegas D. D. 368 368 (Ip), 388 (Ip).
Nuevo Leén: Dominguez z &. EcCart 8270 (Au—222192) ; Le Gilbert 63
(Au--252),09) ; H. Herndndez s.n SNe ~[21/V/1965] (Ip). Oaxaca: yaca: Vilas
29 (Ws), 39 (Ws). Puebla: Anderson & Anderson 531) (Mi); Guerra
Me 16 they Long & Burch 3321 (N); Manning & Manning 53719 (Au- (Au--
231068) ; Ventura A. 15L5 (Wi). San Luis / Luis Potosf: J. | Je Rzedowski
3051 (Ip), 3428 (Ip). Tamaulipas: Serna 34 (Lk). Tlaxcala: Gal-
icia 3 (Ip--226h, Ip). Veracruz: Nevling & Gémez-Pompa 1075
(G). State undetermined: Mutis 1919 9 (W—1562721) ; Schnée s.n.
{[VII--XII] (Mi).

VERBENA CILIATA var. LONGIDENTATA Perry

Additional synonymy: Verbena ciliata longidentata Perry apud
Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex. Res.

Found. Bot. 6:] 1876. 1970.

Additional bibliography: Moldenke, Phytologia 16: 88. 1968;
Kapp, How Know Pollen 104, 135, & 231%: fig. 208 & 276. 1969; Mol-
denke in Correll & Johnston, Man, Vasc. Pl. Tex. (Contrib. Tex,
Res. Found. Bot. 6:] 1217 & "1325. 1970; Correll & Johnston, Man.
Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:] 1876. 1970;
Moldenke, Fifth Summ, 1: 58, 62, & 75 (1971) and 2: 663 & 913.
1971; Moldenke, Phytologia 22: 162, 473, & 97. 1972.

Raditional i}lustrations: Kapp, How Know Pollen 10) & 135, fig.
208 & 276. 1969.

Runyon describes this plant as an annual, 1 foot tall, its
leaves petiolate or subsessile, cuneate-lanceolate in outline,
deeply incised or lobed and dissected, the bark green, the roots
fibrous, the flowers odorless, and the fruit a "cylindric capsule"
{it is actually a schizocarp]. It has been collected as altitudes
from sealevel to 2,00 feet. In addition to the months previously
reported, it has been collected in anthesis and fruit in January
and September. The corollas are described as "blue" on R. Runyon
297, "violet—purple" on R. Runyon 2361, "rose-purple" on mR, Run- Run-

= pe SS acter

and R. ear 1576 & 1577. Recent collectors have found it Boe
ing in black heavy soil, sandy soil over limestone, and sand along
roadsides. Hutchins found it "in calcareous clay loam of barditch"
and his specimen is accompanied by a photograph of the plant in
situ. Cory found it to be "infrequent in grassland of roadsides"
in San Patricio County, Texas. Runyon avers that it blooms from
February to April in Cameron County, where it is "a very common
verbena covering large patches and producing flowers in abundance,
forming large patches in early spring", although for his no. 295
he notes "occasional in open ground". ie

190 P BiYeTyO LyO.GetsA Vol. 23, now. 1

Cumbie 53 bears a notation that it is a voucher for anatomical
studies; it was misidentified as V. bipinnatifida Nutt., so the
anatomical studies, if they have been published, are probably in-
correctly said to refer to that species.

Material of V. ciliata var. longidentata has also been misiden-

tified and distributed in some herbaria under the names V. ambrosi-
folia eglandulosa Perry and V. cloveri var. lilacina Moldenke. On
the other hand, the Demaree 7685, distributed as V. ciliata var.
longidentata, seems to be var. r. pubera (Greene) Perry instead.

Additional citations: TEXAS: Bexar Co.: McCullough 6 (N). Cam-
eron Co.: C. L. Lundell 10656 (N); R. Runyon 1576 (Au-—269653),
1577 (Au--26965), 2361 (Au--268823), 2495 (Au--268726, Au-—
269652, N, N), 2497 (Au--268720). Fayette Co.: Ripple 51-580 (Au—
226865). Garza Co.: B. Hutchins 319 (Lk). Kent Co.: Cumbie 53
(Lk). Kinney Co.: Strother 263 (Au—23 8209). San Patricio Co.:
Cory 51256 (Mi). Webb Co.: “Barrera & Laurel 59 (Lk). Wichita
Co.: Whitehouse 9513 (N). Zapata Coe: Novoa & Cantu & Cantu 18a (Au--
245164, Ip). Zavala Co.: Ramirez & Cardenas a3 ~ (Au—2521)) .
MEXICO: Coahuila: G. A. Voss ss 186 (Sd=-63759)

VERBENA CILIATA var. PUBERA (Greene) Perry

Additional bibliography: Howell & McClintock in Kearney &
Peebles, Ariz. Fl., ed. 2, 727. 1960; Moldenke, Phytologia 16: 88—
89. 1968; Moldenke in Correll & Johnston, Man, Vasc. Pl. Tex.
[Contrib. Tex. Res. Found. Bot. 6:] 1317 & 1325. 1970; Correll &
Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:]
1876. 1970; Moldenke, Fifth Sum. 1: 51, 58, 62, 63, & 65 (1971)
and «221,522,663 0690s 6913 197k.

Recent collectors have found this plant growing in rocky sandy
loam, shallow alkaline soil, gravelly sand of Acacia-Yucca-grass
comminities, and along railroad tracks. Howell & Mc McClintock
(1960) describe the plant as "more prostrate and compact and more
revolute leaf margins" and maintain that it is found "throughout
the range of the species" in Arizona.

The M. Winter él, distributed as V. ciliata var. pubera, is ac-

)
tually Vv. pumila Rydb.
{

—--—-—— ~~

Additional citations: TEXAS: Bailey Co.: Rosson 506c (Lk).
Garza Co.: B, Jensen 8 (Au--2l801). Jeff Davis Co.: Small & Wher-
ry 12072 (Ld). Lubbock Co.: Demaree 7685 (Lk). ARIZONA: Santa
Cruz Co.: Beasley & Finzel 851 (N).

VERBENA CLAVATA Ruiz & Pav. |

Additional synonymy: Verbena clvaara Rufz & Pav. apud Pers.,
Sp. Pl. 3: 346, sphalm. 1819.

Additional bibliography: Pers., Sp. Pl. 3: 346. 1819; Steud.,
Nom. Bot. Phan., ed. 1, 873. 1821; J. F. Macbr., Field Mus. Publ. .
Bot. 13 (5): 613, 615, "617-618, 621, & 625. 1960; Moldenke, Phy- |
tologia 16: 49. 1968; Moldenke, "Résumé Suppl. 16: 28. 1968; *yWol-

1972 Moldenke, Notes on Verbena 191

denke, Fifth Summ. 1: 143 & 192 (1971) and 2: 621, 658, 663, 668,
701, & 913. 1971.

Macbride (1960) keeps V. calcicola Walp. separate from V. cla-
vata Ruiz & Pav., with V. - gynobasis Wedd. as a synonym of the
former. In his discussion of V. calcicola he says "Included by
Schauer in V. clavata R. & P. and conceivably it may be an eco-
logical condition". He cites for it Field Mus. neg. 17409, as
well as Gay s.n. from Cuzco, Meyen s.n. and Weberbauer 1455 from
Arequipa, and Weberbauer Tél, and | and Weddell sn. from Moquegua. For
V. clavata he cites Ferreyra yra 8031, Goodspeed ed 918k, and Sandeman
L611 from Ancash, Eyerdam erdam 25159, | Goodspeed E Exped. 15719, Isern Isern
277, and Mexia oLi.70 from Areq Arequipa, Isern 2478 f from Moquegua,
and | Metcalf f 3037h fro from Tacna, and says 3 that it occurs also in
Chile. Ir regard | the ie and Mexia collections as var. cas-
mensis Moldenke.

Macbride also attempts to keep V. fissa Hayek as a separate
entity, citing for it the following collections: PERU: Ancash:
Raimondi s.n. [Hualas]; Weberbauer 2768 (type). Arequipa: Fer-
reyra 5554; Guenther & Buchtien 133 & 133a; I. M. Johnston 3560;
Weberbauer 389. Tacna: Metcalf 3037) Talso cited by him as Vs
clavata]; Weberbauer 7385. He comments: "Near V. thymoides
Cham. but more depressed, more densely pubescent, larger flow-
ers; as V. calcicola Walp., cited as a synonym in Index Kewensis,
may also p prove to be a variant of V. clavata R. & P. as suggested
by Johnston. About 3 dm. tall, flowers white or pink, on middle
and lower green slopes (Johnston)...F. M. Neg. 1716." He at-
tempts to separate the three supposed species as follows in his
key:
eaves closely canescent strigillose-hispidulous.....V. calcicola
Leaves more or less canescent hirsute-villous-strigose.V. clavata
Leaves lightly hispidulous, greenish.........ssssseeeeeeeVe fissa
' Reece citations: PERU: Arequipa: Vargas Calderén 18089

Ac

VERBENA CLAVATA f, ALBIFLORA Moldenke

Additional bibliography: J. F. Macbr., Field Mus. Publ. Bot.
13 (5): 618. 1960; Moldenke, Phytologia 11: 53. 1965; Moldenke,
Fifth Sum, 1: 143 (1971) and 2: 913. 1971.

VERBENA CLAVATA var. CASMENSIS Moldenke

Additional bibliography: J. F. Macbr., Field Mus. Publ. Bot.

13 (5): 618. 1960; Moldenke, Phytologia 11: 53. 1965; Moldenke,
Fifth Sum. 1: 143 (1971) and 2: 913. 1971.

Macbride (1960) says of this variety: "Var. casmensis....has
scarcely revolute leaves mostly drying blackish, about 3 m. wide,
with short unevenly spreading twisted trichomes both sides; flow-
ers lilac, fragrant. Leaves resemble those of V. trifida; spread-
ing pubescence differs from both plants."

192 P Pee ie@ LaOsG sigA Vol. 23, noOsel

XVERBENA CLEMENSORUM Moldenke

Synonymy: Verbena clemensorum Moldenke apud Howell, Wasmann
Journ. Biol. 10: 377. 1952. Verbena officinalis L. x V. robusta
Greene ex Moldenke, Résumé 371, in syn. 1959. Verbena robusta
Greene x V. officinalis L. ex Moldenke, Résumé 373, in syn. 1959.

Additional bibliography: Moldenke, Phytologia 11: 453—l5).
1965; Munz, Suppl. Calif. Fl. 101. 1968; Moldenke, Fifth Summ. 1:
65 (1971) and 2: 686, 693, & 913. 1971.

Munz (1968) says nv. Clemensorum Moldenke described as possible
hybrid between V. officinalis and V. robusta, from Jackson, Amador
Co. Coarse herb with glabrous stems, stiff ovate incised ioe 2.5
8 cm. long; infl. spicate, compound, "+ puberulent, elongate; corol-
la 2 mm. wide."

VERBENA CLOVERAE Moldenke

Additional bibliography: Hocking, Excerpt. Bot. A.6: 91. 1963;
Moldenke, Phytologia 16: 186. 1968; Moldenke, Résumé Suppl. 16:

2. 1968; Rickett, Wild Fls. U. S. 3 (2): BEE 1969; Moldenke in
Correll & Johnston, Man. Vasc. Pl. ane’ [Contrib. Tex, Res. Found.
Bot.wos) 1315, 1316, & 1321. 1970; Correll & Johnston, Man, Vasc.
Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6: ] 1876 & 1877. 19705
Moldenke, Fifth Summ. 1: 58 & 75 (1971) and 2: 663, 685, 77h,

913. 1971.

Recent collectors have found this plant growing in sandy soil,
sandy silt, red sand, or light brown sandy loam, on clay and
gravel hills, and along roadsides. Wood reports that it "covers
large areas fin Starr County, Texas], common, very similar to no.
692 but flowers twice as large and very saan. Drawe found it _
"abundant on sandy loam soil in heavily grazed pastures" in Hidal-
zo County and Atwood in Opuntia-mesquite communities in Dimmit
County, while Runyon describes it as having "leaves pinnately
cleft, rugose, petiole winged, flowers with no odor, blooming from
April to May.....2 low herb apparently with perennial roots". Pla-
deck states that it has "perennial rootstocks". The corolla color
is given as "blue" on Drawe 296, R. Runyon 2611, and Stokes 19, as
"brilliant blue" on Pladeck s Son. [May Sy 1940 oie and as "lavender
to blue" on A. D. Wood 693. “R. Runyon 1869 is a mixture with V.
plicata Greene. _ A a

Material of V. cloverae has been misidentified and distributed
in some herbaria as V. neomexicana var. hirtella Perry and V.
scabra Vahl. On the other hand, the Novoa & Cantu 18a, distributed
as V. cloverae, is actually V. eee var. mi onetientsen Perry.

Additional sieationa: Brooks Co.: Pladeck sen. [May 5, 190]
(Ws). Dimmit Co.: Atwood 2035 (N). Edwards Co.: H. R. Reed 7
(Mi). Hidalgo Co.: Drawe 296 (Lk); R. Runyon 2611 ~ (Au--268731) ,
4869, in part (Au--269729). Jim Hogg Co.: R. Bruno lj (Lk); Sosa
332 (Lk). Kleberg Co.: F. B. Jones 2977 (Lk). La Salle Co.: Al-
varez, Guajardo, Salazar, | & -McCart 7614 (Lk); F. A. Barkley 171078
(Ip); A Castro 55 (Lk). Starr Co.: + Ramos & wurilio 52 (Au); Re

1972 Moldenke, Notes on Verbena 193

Runyon 4,885 (Au--266150); A. D. Wood 693 (Au--2),7078, Au). Webb
Co.: Co.: Abrigo eon. [April 12, , 1963] (Au--219722) ; McCart 7289 (Mi);
R. Le “MeGregor 1676) (N); wt Stokes 19 (Lk). Zapata Co.: Flores
& & Powell 112 (Lk); M. Gonzalez 5 (Au--2),5139, Ip). Zavala Co.:
Guerra, erra, Garcia, & Garcia Fo) (Au--2)))879) «

VERBENA CLOVERAE f. ALBA Lundell

Synonymy: Verbena cloverae alba Lundell apud Correll & Johnston,
Man. Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:] 1876. 1970.

Additional bibliography: Moldenke, Phytologia 16: 9. 1968;
Moldenke, Résumé Suppl. 16: 2. 1968; Moldenke in Correll & Johns-
ton, Man. Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:] 1315,
1316, & 1321. 1970; Correll & Johnston, Man. Vasc. Pl. Tex. [Con—
trib. Tex. Res. Found. Bot. 6:] 1876. 1970; Moldenke, Fifth Sum.
1: 58 (1971) and 2: 663 & 913. 1971.

This form has been found in flower and fruit in May.

Additional citations: TEXAS: Dimmit Co.: Tharp & York 27 (ip).

VERBENA COCCINEA Raf.
Additional bibliography: Moldenke, Phytologia 10: 102—103.
1964; Moldenke, Fifth Summ. 1: 66 (1971) and 2: 793 & 913. 1971.

VERBENA COCHABAMBENSIS Moldenke
Additional bibliography: Moldenke, he aes 16 NOs 1960.
Moldenke, Fifth Summ. 1: 18) (1971) and 2 : 68h & 913. 1971.

VERBENA COMONDUENSIS Moldenke

Bibliography: Moldenke, Phytologia 18: 3)3--3l4. 1969; Moldenke,
Biol. Abstr. 50: 1298. 1969; Hocking, Excerpt. Bot. A “18: Wy.
1971; Moldenke, Fifth Summ. 1: 75 (1971) and 2: 913. 1971.

My wife, my son, and I visited the type locality of this species
last year and find that it makes a very conspicuous show as it
grows in masses in wet sandy depressions in roadside flats for sev-
eral miles in the type area.

Citations: MEXICO: Baja California: Moldenke & Moldenke 2922 (Z—

ype), 25107 (Ac, Ft, Rf, Z).

xVERBENA CONATA Moldenke

Synonymy: Verbena halei Small x V. officinalis L. ex Moldenke,
Résumé 365, in syn. 1959. Verbena officinalis L. x V. halei Small
ex Moldenke, Résumé 371, in syn. 1959. Fe

Additional bibliography: Hocking, Excerpt. Bot. A.1l: 430. 1959;
Moldenke, Phytologia 11: 45). 1965; Moldenke, Fifth Summ, 1: 370
(1971) and 2: 649, 672, 686, & 913. 1971.

VERBENA CONCEPCIONIS Moldenke
Additional bibliography: Moldenke, Phytologia 9: 201. 1963; Mol-
denke, Fifth Summ, 1: 192 (1971) and 2: 913 5019TLs

XVERBENA CORRUPTA Moldenke
Additional synonymy: Verbena peruviana (L.) Britton x V. phlogi-

194 P By TO b0°Gy EA Vol. 23, no. 1

flora Cham. ex Moldenke, Résumé 372, in syn. 1959. Verbena
phlogiflora Cham. x V. peruviana (L.) Britton ex Moldenke, denke, Résumé

372, in syn. 1959.

Additional bibliography: Moldenke, Phytologia 16: 89 & 93.
1968; Moldenke, Résumé Suppl. 16: 28. 1968; Moldenke, Fifth Sum.
1: 370 (1971) and 2: 522, 689, & 913. 1971.

It appears that the cultivars known as "Defiance" (Hurst;
Vaughan's; Waller-Franklin; Withamfogg), "Defiance Re-selected"
(Watkins & Simpson), "Défiance" (Haage & Schmidt), and "Défiance
Purpurea" (Haage & Schmidt) all belong to xV. corrupta rather
than to xV. hybrida Voss as previously reported by me. "Defiance"
is described as spreading, 12—-15 inches tall, blooming in early
July (in the U. S. A.), the flowers 1/2 inch wide, rich-scarlet
or red with a small creamy-white "eye", Haage & Schmidt describe
"Défiance" as "feuer scharlach" [fiery-scarlet] and "Défiance Pur-
purea" as "purpurroth" [purple-red] in color of corolla,

VERBENA CORYMBOSA Ruiz & Pav.

Additional & emended bibliography: Pers., Sp. Pl. 3: 37. 1819;
Steud., Nom. Bot. Phan., ed. 1, 873. 1821; Reiche & Phil., Fl.
Chil. 5: 283 & 285. 1910; Angely, Fl. Anal. Paran., ed. 1, 571.
rae Yotaro, Gard. Pl. World 1: 131. 1965; Moldenke, Phytologia

: hg--50. 1968; Coats, Pl. Hunters 376. 1969; Heusser, Pollen &
ne Chile 62 & 78. 1971; Moldenke, Fifth Sum, 1: 143, tity
189, 192, & 370 (1971) and 2: 652, 688, 736, & 914. 1971.

The corollas of this plant are described as "blue-violet" on
Burkart 19986. Heusser (1971) says that the pollen has "a ten-
dency to occur chiefly as a pericolpate or pericolporate type
with apertures associated with the contact edges of the faces;
79 u", based on a collection by J. Gallardo (SGO 66315), no
locality of collection designated. He gives the distribution of
the species in Chile as "Atacama--Valdivia" and compares its pol-
len grains with those of Junellia uniflora (R. A. Phil.) Moldenke,

which are syncolporate. He does not specify if he does or does
not regard this as sufficiently important to be a generic charac-
ter.

Coats (1969) reminds us that Elliott & Comber, during their
second Chilean trip in 1929--1930, collected for eventual intro-
duction into the horticultural trade the "hardy Verbena corymbosa".

Additional citations: BRAZIL: Santa Catarina: Smith & Klein 13460

(N). URUGUAY: Burkart 19986 (N, W--2595165).

XVERBENA COVASII Moldenke

Additional synonymy: Verbena santiaguensis (Covas & Schnack)
Moldenke x V. tenuisecta Briq. ex Moldenke, Résumé 373, in syn.
1959. Verbena tenuisecta Briq. x Vv. ganisaciansis (Covas &
Schnack) Moldenke, Résumé 376, in syn. 1959.

Additional & emended bibliography: J. A. Clark, Card Ind. Gen.
Sp. Var. issue 191. 1945; Schnack & Covas, Revist. Argent. Agron.
12: 22) & 228. 1945; Moldenke, Phytologia. 11: 45). 1965; Moldenke,

1972 Moldenke, Notes on Verbena 195
Fifth Summ. 1: 370 (1971) and 2: 522, 69h, 700, & 91h. 1971.

VERBENA CRITHMIFOLIA Gill. & Hook.

Additional synonymy: Verbena crithmithfolia Gill. & Hook. apud
Beetle, Bot. Rev. 9: 670, sphall sphalm. 1943. Glondularia erithnifolia
Gill. & Hook. ex Moldenke, Fifth Summ. 2: 523, in syn. 1971.

Additional bibliography: Autran, Trab. Mus. Farmac. Fac. Cienc.

Méd. Buenos Aires 13: 33. 1907; Frenguelli & Cabrera, Rev. Mus. La
Plata, n. ser., sec. ofic., 1938: 77. 1939; J. A. Clark, Card Ind.
Gen, Sp. Var. issue 183. 19h; Troncoso in BUtcher, Hjerting, &
Rahn, Dansk Bot. Arkiv. 22 (15: 109. 1963; Troncoso in Cabrera,
Fl. Prov. Buenos Aires 5: 137. 1965; Moldenke, Phytologia 16: 186.
1968; Moldenke, Résumé Suppl. 16: 28. 1968; Moldenke, Fifth Sum,
1: 137, 192, & 200 (1971) and 2: 521, 523, 53h, 66h, "665, 668, &
914. 1971.

Collectors have found this plant growing in dry alkaline soil
and in riverbeds, fruiting in April, September, and November (in
addition to the months previously reported) . Semper 117 bears a
label stating that the plant was "rare" where collected (in Men-
doza, Argentina), although this same collector reports it as "a=
bundant" in other parts of the same province on other labels. The
corollas are said to have been "lilac" on Semper 117, "pinkish-
white" on Semper 585 & 616, and "lilac-white" on Semper 65.

Troncoso (1965) comments that "Todas las citas de V. crithmi-
folia......dadas para la Provincia [Buenos Aires] deben referirse
a esta especie [V. hookeriana] ."

The Eyerdam, Beetle, & Grondona 233, distributed as V. crith-
mifolia, is actually V v. hookeriana (Covas & Schnack) Moldenke.

Tdiitional citations: ARGENTINA: Mendoza: Lourteig 89 [Herb.
Inst. Miguel Lillo 113935] (N), 925 (Herb. Inst. Miguel Lillo
114040] (N); Semper 7 (N, S), 22 (N), 65 (N), 117 (N), 439 (N),
585 (N), 616 (N). Rfo Negro: O'Donell 18) (N).

XVERBENA CROOKSHANKSI Moldenke

Synonymy: Verbena bracteosa x hastata Rydb., Fl. Rocky Mtns.,
ed. 2, pr. 1, 740. 1922. Verbena bracteata Lag. & Rodr. x V.
hastata L. ex Moldenke, Fifth Sum, 2: 656, in syn. 1971. Verbena
hastata L. x V. belieteata Lag. & Rodr. ex Moldenke, Fifth Summ, 2:
673, in syn. 1971.

Bibliography: Rydb., Fl. Rocky Mtns., ed. 1, 70 (1917), ed. 2,
pr. 1, 70 (1922), ed. 2, pr. 2, 70 (195k) and ed. 2, pr. 3, 71,0.
1969; Moldenke, Fifth Summ, 2: 656, 673, sit, & 967. 1971.

Rydberg (1917) describes this hybrid as "Resembling most V.
bracteosa in habit, but stouter, more erect, with broad, laciniate
rather stout dissected leaves and shorter bracts. Neb.——Colo."

VERBENA CUNEIFOLIA Ruiz & Pav.
Additional bibliography: Pers., Sp. Pl. 3: 346. 1819; Steud.,
Nom. Bot. Phan., ed. 1, 873. 1821; Reiche & Phil., Fl. Chil. 5:

196 P Here O Ose eek Vol. 23, nd,

287. 1910; J. F. Macbr., Field Mus. Publ. Bot. 13 (5): 61h & 619.
1960; VMoldenke, Phytologia 16: 50. 1968; Moldenke, Fifth Sumn, 1:
143 & 192 (1971) and 2: 665, 701, & 91. 1971.

Vargas Calderén found this species in flower and fruit in Octo-
ber. Macbride (1960) cites Field Mus. neg. 1711 and the follow-
ing specimens: PERU: Ancash: Raimondi s.n. [Pomabamba, Huaraz],
Weberbauer 3060. Cuzco: Herrera 1409. Huancavelica: Raimondi s.
n. [Tayacaya]._ Hudnuco: Macbride & “& Featherstone 1210. Junin:
Soukup B53T & 3970, Stork 10920. San Martin: Ruiz & Pavon s.n.
[Moyobamba], type. iio i of

Additional citations: PERU: Ancash: Vargas Calderén 10320 (Ac).

VERBENA CURTISII Moldenke
Additional bibliography: Moldenke, Phytologia 10: 103. 196;
Moldenke, Fifth Summ. 1: 23 & 370 (1971) and 2: 793 & 914. 1971.

XVERBENA DEAMIT Moldenke

Additional synonymy: Verbena bracteata Lag. & Rodr. x V. stricta
Vent. ex Moldenke, Résumé 359, in syn. 1959. Verbena stricta
Vent. x V. bracteata Lag. & Rodr. ex Moldenke, Résumé 375, in syn.
1959. ae

Additional bibliography: Rydb., Fl. Rocky Mtns., ed. 2, pr. 1,
740 (1922) and ed. 2, pr. 2, 740. 195h; Hitche., Cronq., & Ownbey,
Vasc. Pl. Pacif. Northwest l: 2h). 1959; Moldenke, Phytologia 16:
50. 1968; Moldenke, Résumé Suppl. 16: 1, 2, & 28. "1968; Rydb., Fl.
Rocky Mtns ., ed, 2: pr. 3, 70. 1969; Moldenke, Fifth Summ Le. 56
toils Og ls, 43, 5, oie & 53 (1971) and 2: 656, 657, 666, 697,
é98, & “OU. 1971.

Fell & Fuller found this hybrid growing along roadsides in Win-
nebago County, Illinois. Patterson notes that in Henderson County
of the same state he found some plants procumbent, some ascending,
and some erect -- which is to be expected of the hybrid, consider—
ing its parentage.

Additional citations: ILLINOIS: Henderson Co.: H. N. Patterson
s.n. [Oquawka, July] (Pa, Pa, Pa). Winnebago Co.: Fell & Fuller
56-301 (Ws).

VERBENA DELICATULA Mart.
Additional bibliography: Moldenke, Phytologia 10: 104. 196);
Moldenke, Fifth Summ, 1: 75 (1971) and 2: 665 & 91h. 1971.

VERBENA DELTICOLA Small

Additional bibliography: Hocking, Excerpt. Bot. A.6: 91. 1963;
Moldenke, Phytologia 16: 186. 1968; Moldenke, Résumé Suppl. 16: 2.
1968; Rickett, Wilders. U.S. 3 (2): 36h. 1969; Moldenke in Cor-
rell & Johnston, Man. Vasc. Pl. Tex. (Contrib. Tex. Res. Found.
Bot. 6:] 1316, Anya & 1322--1323. 1970; Correll & Johnston, Man.
Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:] 1799, 186, &
1876. 1970; Moldenke, Phytologia 22: 97. 1971; Moldenke, Fifth
Summ. I: 58 & 75 (1971) and 2: 665 & 914. 1971.

[to be continued]

BOOK REVIEWS

Alma L. Moldenke

"FUNDAMENTALS OF ECOLOGY" 3rd Edition by Eugene P. Odum, xiv &
57h pp., illus., W. B. Saunders Company, London W. C. 1,
Toronto 7 & Philadelphia, Pennsylvania 19105. 1971. £5.00
U.K., $12.10 Canada, $11.75 U.S.

Each of the editions of this text in its time surpassed and
now surpasses in quality of its presentation, content, clarity of
language, instructional value and accuracy of its many diagrams
and photographs, and principles developed -- all others on the
market by wide, wide margins.

I remember coming upon the first edition new in 1953 long after
my college days when there were no ecology courses and being liter-
ally thrilled reading this text from cover to cover.

This particular edition has been extensively revised and has
been emended with much new illustrative material and up-to-date
bibliography. It has been so organized by stated chapters that it
can be used by three different groups of students: (1) non-biology
majors in "man and environment” or "human ecology" courses, (2)
those in the standard or advanced undergraduate courses in ecology
with field and laboratory work, and (3) postgraduates whose courses
deal with principles, environments and ecological technology.

When such an outstanding text by such an outstanding ecologist
and teacher is kept up-to-date and available, why do some college
professors still order all those mediocre wore for their classes?

"FOREST TREES OF THE PACIFIC SLOPE" by George B. Sudworth, xv &
455 pp., illus., facsimile edition of the 1908 Forest Service
of the U. S. D. A. edition. Dover Publications, Inc., New
York, N. Y. 10014. 1967. $5.00 paperback,

In Volume 22, Number 5, of this journal this book was most ap-
preciatively reviewed, but with the present publishing house some-
how inadvertently omitted.

"A FIFTH SUMMARY OF THE VERBENACEAE, AVICENNIACEAE, STILBACEAE,
DICRASTYLIDACEAE, SYMPHOREMACEAE, NYCTANTHACEAE, AND ERIOCAUL
ACEAE OF THE WORLD AS TO VALID TAXA, GEOGRAPHIC DISTRIBUTION,
AND SYNONYMY" by Harold N. Moldenke, 2 vols., 9,2 Pp., Sean
Brumfield, Printers. Orders to 303 Parkside Road, Plainfield,
New Jersey 07060. 1971. $25.00 plus 80 cents postage.

This is more a report than a review of a detailed work that sun—
marizeé over four decades of taxonomic, field and herbarium studies
in these groups. Over five thousand specific and infraspecific taxa
and hybrids are recognized in over a hundred accepted genera. A di-

197

198 Pe YO Bor Gre Vol. 23, m0. 2

chotomous key distinguishes the genera and 2)8 subgroups. Over
15,000 synonyms and rejected names are listed alphabetically. Ac-
tually 20,753 scientific names are accounted for.

Since these families have worldwide distribution, the geograph-
ic distribution ultimately mentions all countries in the world,
with the larger ones subdivided into their states, departments, etc.

Among the cultivated members almost 700 species, varieties and
hybrids are considered.

Fossil forms are represented by 37 species with their geologic
horizons and locations noted.

Because of format and indexing it is easy to locate any species
or geographic area.

Surely this reference work should be indispensable to any in-
stitution undertaking botanical research.

"MAN AND THE ECOSPHERE — Readings from 'Scientific American'" with
commentaries by Paul R. Ehrlich, John P. Holdren & Richard W.
Holm, 307 pp., illus. W. A. Freeman & Co., San Francisco,
California 9410). 1971. $11.00 clothbound, $5.75 paperbound.

These papers have been chosen by an outstanding trio of scien-
tists long and acutely aware of our global ecological situations
and prospects from a 15-year span of this excellent journal. The
27 papers are organized under the following topics: the ecosphere
and preindustrial man; limits rarely perceived as to human popu-
lation, water supply, depleted lateritic soils, food; dimensions
of intervention as in DDT use, thermal pollution; on management
and buying time as desert reclamation; use of fusion powers urban-
ization.

There are many effective diagrams and photographs. Cross ref-
erences, biographical notes, bibliographies and a general index
are given. The only defects noted were the misspelling of Pleisto-
cene on p. 7 and defective color keys to diagrams on pp. 89 and
221%,

This valuable material should (must) reach many eyes (and
minds) in all our higher schools and throughout our literate pub-
lic. It makes excellent supplementary or "required" reading for
general biology and ecology courses along with the companion
volume on "The Biosphere".

"EXOTIC PLANT MANUAL — Fascinating Plants to Live with — Their
Requirements, Propagation and Use" by Alfred Byrd Graf, 80
pp., illus., Roehrs Company, East Rutherford, New Jersey
OO 7See 191s seetebO .

For those who know and use that marvellous work known as
"Exotica" all that is necessary for an evaluation of the present
book under review is to label it a "Baby Exotica" and announce that
it contains over ),200 photographs, with 170 in true color, and
has similar but condensed text on the origin, history, geography,
arrangement, planting care and pest control of exotic plants. The
"Baby" was surely not born like normal mortals but, like Athena,

ee. wee eee eee ee eee

1972 Moldenke, Book reviews 199

sprung full-formed from the head of Zeus. It contains so much
closely compacted information, yet the author's personal en-
thusiasm shines through,

For those who do not know "Exotica" — they should. And this
book is a reasonably priced introduction to it.

The competent plant-loving author has spent a lifetime in
publication and horticulture with trips all over the world.

The taxonomic material is treated with considerable accuracy.
For members of the Verbenaceae the following changes are sugges-
ted:

Verbena tenera var. maonetti for V. peruviana on p,. 410.

Verbena hybrida (1873) for V. x hortensis (1909) as the older
name on p. }11.

Clerodendrum umbellatum var. speciosum for C. speciosum on p.
492 -—- scarcely, if at all, different from the wild C. umbellatun.

Clerodendrum philippinum for C. fragrans pleniflorum -- an un-
fortunate recently discovered name-change apparently forced on us
by the International Rules -- on p. 92.

"AN ISLAND CALLED CALIFORNIA — An Ecological Introduction to its
Natural Communities" by Elna S. Bakker, xiv & 357 pp., illus.,
University of California Press, London, Los Angeles, New York
and Berkeley, California 94720. 1971. $10.00.

The title comes not only historically from Montalvo's grotesque
description in the 1500's "on the right hand of the Indies there is
an island called California, very near the Terrestrial Paradise"
but also ecologically, topographically (San Andreas fault) and
climatically from its differences with the rest of the continent
in species and natural communities. "Isolated by sea, mountain
range, and desert, this area has developed in its own way and at
its own pace; evolutionary history here has woven numerous dis-
tinctive patterns of interaction between life form and the land.
Like the fire pines and cypresses, many such patterns are tag
ends, remnants of much larger biological designs. Others, like
chaparral and coniferous forest, run repeatedly through the warp
and woof of natural California. Such are the patterns of survival
through a rainless summer, on a serpentine slope, in a Sierra can-
yon....-.that are incredibly intricate, multiple, and unfortunate-
ly irreplaceable, if obliterated by man's heedlessness or apathy."

The text describes accurately and attractively the natural com-
munities encountered on this book-tour from the San Francisco Bay
area eastward across the High Sierras to the state's border and
then south through the deserts. There are attractive black and
white photographs and some excellent ecological diagrams. At the
end there are a finely chosen topical bibliography, useful but
not complete (for example, wocas on p. 201) lists of the scientif-
ic names of the plants and animals mentioned, and an index. The
book looks very neat, yet a few words were misspelled: absence on
p. 71, descendants on p. 97, and chapter on p. 209.

The University of California Press deserves much appreciation

200 PH YenyOel OsGrrlA Vol. 23, maace

for its many popular and/or scientific contributions of high
caliber rolling off its presses year after year.

"WORLD VEGETATION TYPES" edited by S. R. Lyre, 26) pp., illus.,
Columbia University Press, New York, N. Y. 10025. 1971.
$12.50.

This book, small in size but far from small in its scientific
value or purchase price, contains a representative selection of
writings which deal with examples of those main types of wild
vegetation which covered most of the land areas of the earth un-
til relatively recent times. They range in original publication
time from 1918 to 1969, from a wide variety of journals and
books, and from 2l, recognized writers from several countries and
times. They all make valuable reading and all are provided with
their own bibliographies.

At the close of the book there is no index but there is a
selected bibliography topically arranged as follows: tropical
rain forest, tropical seasonal forest and tropical savanna, tropi-
cal montane vegetation, mid-latitude forests, sclerophyllous
shrublands, mid-latitude grasslands and grassland-forest ecotones,
mid-latitude desert and semi-desert, tundra and tundra-forest
ecotones, and regional accounts. The thinking is strongly in-
fluenced by the Clementsian "life-form" and "climatic climax"
concepts.

There are some fine black/white photographs of several of
these world vegetation types.

"MICROORGANISMS AND MAN" by Orville Wyss & Curtis Eklund, ix &
389 pp., illus., John Wiley & Sons, Inc., London, Sydney,
Toronto & New York, N. Y. 10016. 1971.

"This textbook is a [well structured and really interesting]
guide for an elementary course in microbiology....The students
who complete this course will acquire a sympathetic view of sci-
ence and scientists and will learn that procedures for securing
scientific information are within their own capabilities." The
text is effectively illustrated with many black/white photo-
graphs and diagrams. The glossary and index are both useful.

The book stresses the various types of organisms covered in
microbiology, antigens and antibodies, infectious diseases of
man, water and food and industrial microorganisms,

PHYTOLOGIA

Designed to expedite botanical publication

CONTENTS

EDDEMA, C., Sclerocarpus uniserialis (Compositae) in Texas

ENN Sod 2 YW die es ok Whine tae ts os Gan ye ores! ow ne 201

Mi 01 DENKE, H. N., Notes on new and noteworthy plants. LV ....... 210
JLDENKE, H. N., Additional notes on the genus Verbena. XI ...... 212

r ‘ , H. H., The taxonomy of Zea mays (Gramineae) ............ 248
. 250°

SEE A.J, BOOK TEVIEWS °c). 2 ooi cs oss ee eh as ee

LIBRARY

APR 20 1977

NEW YORK

ee ee ees eee eee es sis =
=i = ~~ -

Published by Harold N. Moldenke and Alma L. Moldenke

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

Price of this number, $1; per volume, $7.50, in advance,
or $8, at close of volume

April, 1972 No.

SCLEROCARPUS UNISERIALIS (COMPOSITAE) IN TEXAS AND MEXICO

Charles Feddema
Forest Service Herbarium, Fort Collins, Colorado 80521.

The genus Sclerocarpus (Compositae-Heliantheae) consists
of yellow-rayed sunflowers common locally in southeastern

Texas and in much of Mexico and Central America. Of the
eight species known, one occurs widely in tropical Africa
whereas the others are American. Although one species is
perennial the rest are somewhat weedy annuals most commonly
encountered along roadsides and field margins or in grain-
fields and pastures. Superficially, Sclerocarpus resembles
several other genera of the Heliantheae. It is unique,
however, in that the receptacular bracts completely surround
the disk achenes, and at maturity those near the margin of
the disk become thick and hard, closely surround the achene,
and develop tubercles or swellings on the surface. This
peculiar fruit type, consisting of the achene and its closely
investing bract, is here designated a sclerocarp.

The genus was proposed by J. N. Jacquin (1781) who pub-
lished a series of plates including one entitled Sclerocarpus
africanus. It represented the African species later validly
published under that name. A second species was discovered
by Thomas Drummond at San Felipe, Austin County, Texas, and
described by Hooker (1837) as Gymnopsis uniserialis. This
was later transferred to the proper genus by Hemsley (1881).
Correll and Johnston (1970) recognize S. uniserialis as the
only species in Texas, and indicate that it is widespread in
Mexico. A number of populations closely related to S.
uniserialis do occur in Mexico, and recent studies on their
morphological variation and geographic range have been the
basis of the taxonomic revision of the group presented here.

One population from Mexico was described as S. uniserialis
var. papposus Greenman. This differs in so many characters
and to such a degree from the Texas plants, however, that it
must be regarded as a distinct species. The remaining popu-
lations are more closely related, morphologically, to the
Texas plants and must be considered as sub-specific elements
of S. uniserialis. Thus circumscribed, S. uniserialis is
the most variable species in the genus and the most wide-
ranging one in America. It consists of three major popu-
lations and a number of minor ones, all apparently having
the same chromosome number and varying degrees of isolation
from each other. These major populations are here treated
as varieties and may be distinguished from each other and
from S. papposus by the following key.

201

202 PHY T.O°L01G, 2 A Vol. 23, no. 2

KEY TO THE TAXA

A. Involucral bracts inconspicuous, acicular-involute,
mostly less than 3 mm. long; receptacle white-pubescent,
scarcely enlarged from the peduncle; pappus of marginal
disk achenes 1.5-4 mm. long; anthers conspicuously
exserted at anthesis..............eseee- 1. S. papposus

A. Involucral bracts evident, mostly 4-30 mm. long;
receptacle glabrous, conspicuously enlarged from the
peduncle; pappus of marginal disk achenes less than
1.5 mm. long; anthers mostly retained in the corolla
at Tanehesd's) spore ave e eile! s)«isinis)slafo).s\elle) «ieee =1-)~ 2. S. uniserialis

B. Receptacular bracts green in bud or red only on
apical margin; mature marginal sclerocarps ribbed
or with low, uniform tubercles; involucral bracts
mostly more than 5 mm. long; plants mostly more
than 5 dm. tall.

C. Mature marginal achenes elliptic-fusifom;
mature marginal sclerocarps cylindrical-conoid,
the body mostly low-tubercled, gradually
reduced to the beak, not or scarcely bent......
poodedsDooOU GOOD He OUdOS ...2a. var. uniserialis

C. Mature marginal achenes obliquely obovate in
lateral outline; mature marginal sclerocarps
somewhat laterally compressed, the body smooth
or ribbed, the beak strongly bent from it......
suononoonooanouDCdOooboGSaC 2b. var. frutescens

B. Receptacular bracts red or purplish in bud; mature
marginal sclerocarps swollen about the middle with
large complex tubercles; involucral bracts mostly
less than 5 mm. long; plants mostly less than 5 dm.
(ANUS Goan o00 shelolololelelsielelalelsle/eitiele(2 Cll nicl G/mebUb ied Gelis lls

1972 Feddema, Sclerocarpus 203
TAXONOMY
1. SCLEROCARPUS PAPPOSUS (Greenm.) Feddema, comb. nov.

Sclerocarpus uniserialis var. papposus Greenman, Field
Mus. Publ. Bot. 2:346. 1912. Type: MEXICO: GUERRERO:
Rio Balsas, 26 Aug 1910. C. R. Orcutt 4208 (Holotype
F! Isotype MO!).

Erect, mostly strigose annuals to 1 m. tall, the branches
long and slender, opposite below, alternate above; leaves
5-12 (20) cm. long, blades ovate-trullate to deltoid,
scabrous to strigose above, strigose beneath, petiole
mostly one-fourth to one-third as long as the blade; heads
few to several, obconoid, 10-20 mm. high, on long, often
curved, peduncles; receptacle green, strigose, conical,
1.5-3.5 mm. high, little wider than the peduncle; involucre
inconspicuous, bracts 3-7, green, mostly less than 3 mm.
long, involute-acicular in age; ray flowers 3-7, neutral,
ligule yellow-orange, mostly broadly ovate, 6-12 mm. long,
tube greenish, subequal to the ligule; ray achenes green,
fleshy throughout when living, persistent on the receptacle;
disk flowers (4) 6-12 (15), arcuate, ascending, corolla
8-14 mm. long, greenish or yellow below, reddish near the
lobes, these reddish at the base with the apex recurved,
bristly without, pappilate within; anthers exserted at
anthesis; immature receptacular bracts thin, green, arcuate,
tubular, enclosing the corolla nearly to the lobe; mature
sclerocarps to 13 mm. long, cylindric-conoid narrowing
gradually to the apex of the achene, more sharply so to the
beak or enlarged again about the pappus, the surface with
rows of low tubercles; disk achenes fusiform-arcuate, 7-11
mm. long including the pappus, this a conspicuous green or
yellow crown of irregular fused bristles to 4 mm. long.

Guerrero, at elevations of approximately 500 to 2,000
meters, flowering August to October.

This species is most closely related to S. uniserialis
from which it differs chiefly in its very long pappus, its
longer sclerocarps, its very small pubescent receptacle,
short involucral bracts, and reduced number of disk flowers.
It is a rare species known only from the type collection
and two others.

Additional specimens examined: GUERRERO: North of Rio
Balsas, Dist. Adama, Temisco, top of Barranca Limo, 30 Oct
1939, Ynez Mexia 8707 (F, GH, MO, NY, US); Taxco Viejo,

18 Sept 1937, R. Q. Abott 402 (GH).

20 PH eT, Ob Osa A Vol. 23, sure

2. SCLEROCARPUS UNISERIALIS (Hook.) Benth. & Hook. f. ex
Hemsley, Biol. Centr. Am. Bot. 2:164. 1881.

Erect, mostly strigose annual, to 2 m. tall, branches
long, ascending, opposite below, alternate above, pubes-
cence sometimes somewhat spreading; leaves 3-6 (20) cm.
long, the cauline large, long-petioled, soon withering,
those of the branches smaller, petiole mostly one-fourth
to one-third as long as the blade; blade ovate-trullate to
deltoid, scabrous or with ascending hairs above, strigose
beneath, margin dentate, apex mostly acute or acuminate,
base mostly cuneate; heads few to numerous, hemispheric to
cylindric, 7-25 mm. high, solitary, terminating the long-
pedunculate branch-ends; receptacle whitish, glabrous,
conical, to 10 mm. long; involucre uniseriate, campanulate
in bud, spreading or reflexed in age, bracts 5-9 (12) green,
mostly 5-10 (15) mm. long, linear-elliptic or oblanceolate,
often oblique; ray flowers 5-9, neutral, ligules yellow-
orange, ovate-oblong to suborbicular, 6-20 (30) mm. long,
the apex shallow-toothed, tube one-fifth to one-half as
long as the ligule; ray achenes greenish, fleshy throughout,
linear-arcuate when living, shriveled and persistent on
the receptacle when dry; disk flowers (5) 15-40 (50),
spreading to erect, corolla 6-13 mm. long, gradually
enlarged from base to throat, yellow or reddish or with
zones of both colors, lobes 2-5 mm. long, recurved,
strigose or bristly at the apex without, long papillate
near the middle within; anthers mostly retained in the
corolla tube; receptacular bracts green or purplish in bud,
enclosing the corolla to about the middle, the mature bract
(sclerocarp) 3-10.5 mm. long, beaked, green, brownish,
purple, or mottled, the surface sparsely strigose, low-
ribbed or tubercled, the body often somewhat laterally
compressed; marginal disk achenes 3-6 mm. long including
the pappus, black, often somewhat laterally compressed,
fusiform-arcuate or obliquely obovate in lateral outline;
Ppappus a crown of basally fused bristles sometimes reduced
to form one or more scales, mostly less than 1 mm. long.

The total range of S. uniserialis includes an elongated
area from southeastern Texas to northwestern Guatemala.
The three varieties form a graded series, morphologically,
from north to south in respect to the height of the plants,
the length of the involucral bracts, the form of the mature
receptacular bract (sclerocarp) and other characters.
Plants from Texas are 1-2 m. tall with involucral bracts
1-3 cm. long and straight sclerocarps with small tubercles

1972 Feddema, Sclerocarpus 205

or none. Plants from Guerrero are typically less than 1/2
m. tall, have involucral bracts less than 1/2 cm. long,
sclerocarps which are strongly bent and have large tuber-
culate swellings about the middle. Plants from eastern
Mexico are intermediate in these characters.

2a. SCLEROCARPUS UNISERIALIS (Hook.) Benth. & Hook. f. ex
Hemsl. var. UNISERIALIS

Gymnopsis uniserialis Hook., Icones Plantarum, Vol.
2.t.145. 1837. Type: TEXAS: Austin Co., San Felipe,
Drummond 135 "bis ...Texas II" (Holotype K!;
isotypes GH! K! NY!).

Aldama uniserialis (Hook.) A. Gray, Journ. Boston Soc.
Nat. Bist. 672285 1850:

Sclerocarpus major Small, Fl. Southeastern U.S. 1250.
1903. Type: TEXAS: Bexar Co., Comale Creek, 1847-
1848, Lindheimer 432 (Holotype NY!; isotypes F! GH!
MO! NY! US!).

Mostly 1-2 (2.5) m. tall, usually much-branched; leaves
3-6 (17) cm. long, blade ovate-trullate or rhombic, with
3-8 (16) irregular teeth per side; heads few to many, 10-25
mm. high on peduncles to 15 cm. long; receptacle 3-5 (9)
mm. long; involucre often strongly reflexed in age, bracts
mostly 5-10 (15) mm. long; ray flowers mostly 5-9, 19-20
(35) mm. long, the ligule mostly narrowly ovate; marginal
disk flowers inserted nearly perpendicular to the surface
of the receptacle; disk corollas yellow or brownish-red;
receptacular bracts green or reddish on the distal margin
in bud; mature marginal sclerocarps 6-10.5 mm. long, nearly
horizontal on the receptacle, the body 1.5-3 mm. diam,
long-conoid, narrowing gradually to the beak; the body with
a few long longitudinal ribs, sometimes with small rounded
tubercles on the ribs or scattered on the beak; mature
marginal achenes 4-6 mm. long, narrowly fusiform, slightly
oblique, pappus mostly 0.2-0.5 mm. long, usually lengthened
adaxially and abaxially, and reduced laterally.

Chromosome number: N = 12 (Feddema 1468, MICH).

The Texas populations of S. uniserialis are well isolated
from those in Mexico. Although the species ranges nearly

to the Rio Grande at Brownsville, the nearest coastal col-
lections known from Mexico are from southern Tamaulipas.
Inland, the nearest Mexican plants of the species have been

found near Monterrey. In Texas the species ranges from

206 PH YoTVOsLOrGi T A Vol. 23, no. 2

Fort Bend County, along the coastal plain to Cameron
County, up the Rio Grande to Webb County, and its western
limits run through Medina and Bexar Counties to Williamson
County. Variety uniserialis is very variable genetically
and in response to environmental conditions. J. K. Small
described S. major which he differentiated from S.
uniserialis on the basis of the larger heads and involucral
bracts of the former. Plants of this kind occur widely.
The first head to develop terminates the main stem and is
usually the largest. It may be especially so early in the
season if water is plentiful.

Two distinct color forms of var. uniserialis occur. In
most plants the disk flowers, like the rays, are yellow.
Most plants from Duval, Kenedy, Brooks, Jim Hogg, Starr,
and Willacy Counties, however, have reddish or purplish-
brown disk flowers.

2b. SCLEROCARPUS UNISERIALIS (Hook.) Benth. & Hook. f. var.
FRUTESCENS (Brandg.) Feddema, comb. nov.

Sclerocarpus frutescens Brandg., Univ. Calif. Publ.
Bot. 4:281. 1912. Type: SAN LUIS POTOSI: between
Bagre and Minas de San Rafael, Jul 1911, C. A.
Purpus 5157 (Holotype UC; isotypes F! GH! MO!).

Long-lived annual, mostly 3-10 dm. tall, sometimes
persisting with thickened stem and root and rooting at
lower stem and branch nodes; leaves mostly 3-8 cm. long;
blade mostly narrowly ovate-trullate, sometimes strigose-
pilose beneath, the pubescence sometimes yellowish, margin
rarely entire, mostly with 4-6 teeth per side; heads few
to several, 8-19 mm. high; receptacle (2) 3-5 (9) mm. long;
involucre often reflexed at maturity, bracts mostly 4-8
(13) mm. long; ray flowers mostly 5-7, 9-15 (23) mm. long,
the ligule narrowly ovate; disk flowers spreading or
ascending, yellow or with a purplish area above the middle,
the lobes sometimes with a band of purplish hairs within;
receptacular bracts green; mature marginal sclerocarps
5-8 (9) mm. long, with broad body and narrow beak about as
long and strongly bent; body of the sclerocarp few-ribbed,
1-2 mm. wide at base, widening above, then narrowing sharply
to tne tubular beak, rarely with numerous small uniform
tubercles; mature marginal achenes 3-4.6 mm. long, consid-
erably oblique in lateral outline, sharply narrowed to the
pappus.

Chromosome number: N = 12 (Feddema 1502, MICH).

1972 Feddema, Sclerocarpus 207

Southeastern Coahuila, Nuevo Leon, southern Tamaulipas,
San Luis Potosi, Hidalgo, Guanajuato, Mexico, Veracruz,
eastern Oaxaca, Campeche, Yucatan, Quintana Roo, Chiapas,
and northeastern Guatemala, at elevations from sea level to
2700 meters, usually flowering from March to September but
sometimes persisting into the second season.

This variety is distinguishable from var. uniserialis by
its shorter, bent sclerocarps and from var. rubridiscus by
the longer involucral bract, green receptacular bracts in
bud and sclerocarps which are merely ribbed or with small
uniform tubercles.

2c. SCLEROCARPUS UNISERIALIS (Hook.) Benth. & Hook. f. ex
Hemsl. var. RUBRIDISCUS Feddema, var. nov.

Plerumque brevis, 2-6 (15) dm. alta; folia (2) 3-4 (9)
cm. longa; lamina anguste trullata (2) 3-4 (7) cm. longa;
bracteae involucri lineares vel oblanceolatae, circiter
2.5-5 mm. longae, 1-2 mm. latae; receptaculum glabrum;
marginis receptaculi paleae maturae valde curvae, 6-8 mn.
longae, in medio valde turgido-tuberculatae, corpore
rostrum plerumque paullo longiore; pappus 0.3-0.9 mm.
longus; achaenia 4-5 mm. longa.

Plants typically 2-5 dm. tall, the lower branches long,
the upper 1-3 short; leaves mostly 2-5 cm. long; blade 1-4
(8) toothed, rarely entire or with short basal lobes or
rounded apex; heads 9-20 mm. high; receptacle mostly 5-7
(10) mm. long; involucre appressed to the sclerocarps or
slightly spreading in age, bracts mostly 2.5-5 mm. long;
ray flowers mostly 5-6, ligule ovate or suborbicular, 8-14
mm. long; disk flowers oriented nearly vertically on the
receptacle, the corolla 6-12 mm. long, the lobes reddish
below, yellow apically with purplish hairs within near the
middle; receptacular bracts reddish-purple in bud; mature
marginal sclerocarps 5-9 mm. long, with a broad body
subequal to the strongly bent narrow beak; body of the
sclerocarp 3-4 mm. long, enlarged above with irregular
rounded swellings, narrowed sharply to the tubular beak;
marginal achenes 4-5 mm. long, somewhat laterally
compressed, obliquely obovate in lateral outline.

Chromosome number: N = 12 (Feddema 1710, MICH).

Guerrero, Morelos, western Oaxaca, at elevations of 600
to 2,000 meters, flowering July to November.

208 Po Yas sOyGed A Vol. 23, no. 2

Type: GUERRERO: 6 mi S. of Chilpancingo, 13 Aug 1961,
Feddema 1650 (Holotype MICH).

This variety is distinct in its typical form which occurs
in a narrow area of Guerrero from near Iguala to Chilpancingo.
It is distinguished by its low stature, the purplish recep-
tacular bracts in bud, the short involucral bracts, and the
swollen-tuberculate mature sclerocarps.

In addition to the above major populations of Sclerocarpus
uniserialis, there are a number of others related to var.
rubridiscus but having various combinations of characters
making them somewhat intermediate between this and the other
varieties. Most plants from near Taxco and from southern
Morelos are taller than the typical Guerrero variety (var.
rubridiscus) but those in Michoacan appear intermediate
between it and var. uniserialis from Texas. In an elongated
area from southern Morelos and Puebla through the Valley of
Oaxaca to the Isthmus of Tehuantepec, many populations are
found which have some characteristics of var. rubridiscus.
These plants constitute a relatively uniform population over
a large area and are intermediate in form between var.
rubridiscus and var. frutescens.

Chromosome counts for the three varieties of S.
uniserialis and for intermediates between var. rubridiscus
and var. uniserialis and between var. rubridiscus and var.
frutescens indicate that N = 12 for all populations. No
counts have been made for S. papposus but other species
were counted at N = 11, 14, and 18. The existence of
intermediates between the principal populations of S.
uniserialis, and their similar morphology and the same
chromosome numbers suggests that they are closely related
and interfertile. The variation and distribution of the
various populations of S. uniserialis and S. papposus
Suggest a classic pattern of speciation in which an
ancestral population, relatively homogeneous and inter-
breeding, by isolation and migration, develops variant
populations. Variation has progressed rapidly and to a
high degree in S. papposus. S. uniserialis var.
uniserialis, apparently not at present interbreeding with
other populations, includes considerable variation within
itself. Var. frutescens and var. rubridiscus, well
differentiated in parts of their ranges, have intermediates
between them. These may be relict populations from a
period when differentiation was less advanced or may be
the result of more recent hybridization and introgression
after differentiation and a degree of isolation.

1972 Feddema, Sclerocarpus 209
ACKNOWLEDGMENTS

I wish to express appreciation to Dr. Rogers McVaugh
who assisted materially in this study and to Dr. F. J.
Hermann and Dr. William A. Weber who read the manuscript.

REFERENCES
CORRELL, D. S. AND M. C. JOHNSTON. 1970. Manual of
the Vascular Plants of Texas. Texas Research Foundation.

Renner, Texas.

HEMSLEY, W. B. in GODMAN AND SAVIN. Biologica Centrali-
americana. Botany 1: 364.

HOOKER, W. J. 1837. Icones Plantarum. Vol. 2: t.145.

JACQUIN, J. N. 1781. MIcones Plantarum Rariorum. Vol.
etek 6’

NOTES ON NEW AND NOTEWORTHY PLANTS. LV

Harold N. Moldenke

BOUCHEA SPATHULATA var. LONGIFLORA Moldenke, var. nov.

Haec varietas a forma typica speciei tubo corollae 3--l cm.
longis recedit.

This variety differs from the typical form of the species in
having its corolla-tubes 3--) cm. long.

The type of the variety was collected by Ivan Murray Johnston
(no. 872) on rocky flats and ridges in the vicinity of Aguaje del
Pajarito, canyon at the west end of the Sierra de la Fragua, 2-3
km. north of Puerto Colorado, in western Coahuila, Mexico, between
September 1 and 3, 191, and is deposited in the herbarium of the
University of Texas at Austin, Texas. The collector describes the
plant as a common bush, 2--3 feet tall, in the type locality, with
lilac or bluish corollas.

CALLICARPA ACUMINATA var. ARGUTEDENTATA Moldenke, var. nov.

Haec varietas a forma typica speciei marginibus laminarum
foliorum patente arguteque irregulariterque dentatis recedit.

This variety differs from the typical form of the species in
having the leaf-blades irregularly sharp-dentate along the margins
with spreading, not appressed, teeth.

The type of the variety was collected by Marshall C. Johnston
(no. 5799) along a small creek in thorn forest on limestone slopes
16 miles east of Casas in the northern foothills of the Sierra de
Tamaulipas, Tamaulipas, Mexico, on September 28, 1960, and is de-
posited in the herbarium of the University of Texas at Austin,
Texas. The collector notes that the plant was "infrequent" at the
type locality.

CALLICARPA ACUMINATA var. PRINGLEI (Briq.) Moldenke, comb. nov.
Callicarpa pringlei Briq., Bull. Herb. Boiss., ser. 1, : 345—
36. 1896.

KALAHARIA UNCINATA f. RUBRA Moldenke, f. nov.

Haec forma a forma typica speciei corollis rubris recedit.

This form differs from the typical form of the species in
having red (rather than yellow) corollas.

The type of the form was collected by Robert J. Rodin (no.
2710) in open bush-veld with Acacia, Zizyphus, Grewia, and other
low trees, near Otjiwarongo on the road to Outjo, Namibia, on De-
cember 2, 19,7, and is deposited in the United States National
Herbarium at Washington. The collector describes the plant as
an herb 1 to 11/2 feet tall with curved thorns and red flowers.

LANTANA VELUTINA f. MACROPHYLLA Moldenke, f. nov.
Haec forma a forma typica speciei laminis foliorum usque ad
210

1972 Moldenke, New and noteworthy plants 215;

6.5 cm. longis et ) cm latis recedit.

This form differs from the typical form of the species in hav-
ing its leaf-blades to 6.5 cm. long and l, cm. wide.

The type of the form was collected by Cyrus Longworth Lundell
and Amelia A. Lundell (no. 12320) on a mountainside at kn. 157 of
the Acapulco Highway near Taxco, Guerrero, Mexico, on August l,
1943, and is deposited in the Lundell Herbarium at the University
of Texas at Austin, Texas. The collectors describe the plant as
a low shrub, the corollas white with a yellow eye, and the fruits
white.

PAEPALANTHUS FULGIDUS var. ZULOAGENSIS Moldenke, var. nov.

Haec varietas a forma typica speciei foliis parvioribus recedit.
This variety differs from the typical form of the species in
its shorter leaves, which are only to 15 mm. long and about 2 mm.

wide.

The type of the variety was collected by Julian Alfred Steyer-
mark (no. 103839) in a swampy savanna, at 2300 m. altitude, on the
Planicie de Zuloaga, Rio Titirico, Amazonas, Brazil, between
October 10 and 15, 1970, and is deposited in the Britton Herbarium
at the New York Botanical Garden. The collector notes that the
plant grew in dense clumps, the leaves being rich-green on both
surfaces, coriaceous-stiff, and the flower-heads whitish.

VERBENA GUARANITICA (Troncoso) Moldenke, comb. nov.
Glandularia guaranitica Troncoso, Darwiniana 16: 618--621, fig.

3. 1971.

VITEX SCHUNKEI Moldenke, sp. nov.

Arbor, foliis 5-foliolatis, foliolis chartaceis oblong-elongatis
usque ad 22 cm. longis juventute adpresso cinereis maturitate
glabratis nitidisque margine obscure denticulatis; inflorescentiis
cymosis, cymis parvis, pedunculis 2.5--3 cm. longis minutissime
puberulis, pedicellis usque ad 2 mm. longis adpresse puberulis,
calyce campanulato 1 mm. longo 2 mm. lato adpresso-puberulente,
corolla hypocrateriforme, tubo ca. 5 mm. longo.

Tree, about 8 m. tall; branchlets and twigs rather slender,
gray, glabrate; leaves decussate-opposite, 5-foliolate, long-
petiolate; petioles ca 12 cm. long on mature leaves, angular-
furrowed, very minutely and obscurely appressed-puberulent or sub=
glabrate; leaflets 5, chartaceous when mature, unequal, 10.5--22
em. long when mature and ],--6 cm. wide, elongate-oblong, long-
acuminate at the apex and base (the base simulating a winged peti-
olule), glabrous and shiny above, glabrate or subglabrate beneath
(the puberulence extremely minute and obscure, discernible only
under a lens), more densely and visibly appressed-puberulent with
ashy-gray hairs beneath when very immature; inflorescence cymose,
abundant, crowdec on the youngest twigs; peduncles slender, 2.5--3
em. long, flattened and nigrescent in drying, very minutely puber-
ulent; cymes small, few-flowered, about 1.5 cm. long and wide;
pedicels filiform, to 2 m, long, appressed-puberulent; calyx

212 PIR (50; 00.G Th Vol. 23, no. 2

campanulate, about 1 mm. long and 2 mm. wide, appressed—puberu-
lent; corolla hypocrateriform, its tube slender, cylindric, about
2 mm. long, densely appressed-puberulent on the outside, more
conspicuously so upwards, the limb about 8 m. wide, densely
puberulent on both surfaces.

The type of this species was collected by José Schunke V. (no.
908) in a low forest in Neshuya, at 250 m. altitude, Loreto, Peru,
on October 2, 1965, and is deposited in the Britton Herbariun at
the New York Botanical Garden. The collector describes the color
of the corollas as "2.5P5/8".

ADDITIONAL NOTES ON THE GENUS VERBENA. XI

Harold N,. Moldenke

VERBENA [Dorst.] L.

Additional synonymy: Veraena Lam. ex Tawada, Biol. Mag. Okin-
awa [Okinawa Seibutsugakki] ] (6): 36, sphalm. 1967.

Additional & emended bibliography: Matth., Disc. Valgr., ed.
1, 2: 399 & 1107 (1585) and ed. 2, 2: 1107. 1586; Schrtder,
Pharm. Med.—Chem., ed. 2, : 167--168. 169; Micheli, Cat. Plant.
Hort, Caes. Florent., ed. 1, 98 & 182. 178; Lanfossi, Giorn.
Fis. Chim. Stor. Nat. Med. & Art., ser. 2, 10: 48. 1827; Micheli,
Cat. Plant. Hort. Caes., Florent., ed. 2, 98 & 182. 1831; Jan,
Elench, Pl. 1. 1831; Moris, Fl. Sard. 3: 342343. 1859; P. M.
Rodriguez, Pl. Med. Parag. 1915; Tischler, Tabul. Biol. 4: 2h.
1927; Furusato, Bot. & Zool. Theoret. & Appl. Tokyo 8: 1303--
1311. 1940; Covas & Schnack, Revist. Argent. Agron. lj: 229 & 231,
fig. 32. 1947; Darlington & Wylie, Chrom. Atl., pr. 1, 322--323
& 518. 1956; Schnack, Fehleisen, & Cocucci, Revist. Argent. Agron.
2h: 129--135. 1957; Cabrera, Revist. Invest. Agric. 11: 332 & 398.
1957; Rattenbury, Madroflo 15: 50--51. 1959; Schnack, Fehleisen, &
-Cocucci, Revist. Fac. Agron. La Plata 35: {7]--56, fig. 1--3.
1959; Lems, Sarracenia 5: 79. 1960; Lewis & Oliv., Am. Journ. Bot.
8: 638—613, fig. 1—26. 1961; Martin & Barkley, Seed Ident. lian.
37, 58, 19y--195, & 221, fig. 260, pl. 23h--236 & 397. 1961; Rat-
tenbury, Madrofio 17: 116. 1963; Ruiz Leal, Revist. Facult. Cienc.
Agrar. Mendoza 11: 173 & 17h, fig. 8. 196; Banerji, Rec. Bot.
Surv. India 19: 75. 1965; Huynh, Schweiz. Naturforsch. Ges. Mém.
Soc. Helvet. Sci. Nat. 85: 1--178. 1965; Tawada, Biol. Mag. Okin-
awa [Okinawa Seibutsugakki] (6): 36--37. 1967; T. H. Everett,
Living Trees World 297. 1968; M. Martinez, Pl. Med. Mex., ed. 5,
3h0—-3h1, 505, & 579. 1969; Bolkh., Grif, Matvej., & Zakhar.,
Chrom. Numb. Flow. Pl. 715--717. 1969; Angely, Fl. Anal. Fitoge-
ogr. Est. S. Paulo, ed. 1, : 825, 838--8)0, & xix. 1970; Tron-
coso, Darwiniana 16: [613]—621, fig. 1--3. 1971; Vyas & Agarwal,
Phyton 28: 1-5. 1971; Anon., Biol. Abstr. 53 (1): B.A.S.I.C. S.

aa {2 Moldenke, Notes on Verbena 213

267. 1972; D. Burpee, Burpee Seeds 1972: 1, 28, 56, & 57, pl
3177 & L3hs. 1972; W. J. Park, Park's Flower Book 1972: 47. 1972;
Vyas & Agarwal, Biol. Abstr. 53: 499. 1972; Moldenke, Phytologia
23: 191--196. 1972.

Additional excluded species are:
Verbena foemina Brunf. = Senecio vulgaris L., Carduaceae
Verbena foemina Trag. = Sisymbrium officinale L., Brassicaceae
Verbena recta sive mas Fuchs = Sisymbrium officinale L., Brassi-

caceae

The Tawada reference in the bibliography above is sometimes
cited as "1968", but the date, December 25, 1967, appears both on
the title-page of the number and on the first page of Tawada's
paper. The Madrofio references (1959, 1960, 1962) are sometimes
cited as "Solbrig, Madrofio...." or "Darlington, Document. Chron.
Numb. Pl.....", but actually are in several continuations of Rat-
tenbury's original enumeration, with no new author or compiler
indicated. The Tischler (19275 reference is sometimes cited as
"Tischler, Pflanzl. Chrom....."

VERBENA ABRAMSI Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 65 (1971)
and 2: 69, 679, & 912. 1971; Moldenke, Phytologia 22: 58--59.

1972.

XVERBENA ADULTERINA Hausskn,

Additional bibliography: Moldenke, Fifth Summ. 1: 206 (1971)
and 2: 686, 699, 710, & 912. 1971; Moldenke, Phytologia 23: 59.
1972.

VERBENA ALATA Sweet
Additional synonymy: Verbena elata Jan, Elench, Pl. 1. 182),
Additional bibliography: Jan, Elench. Pl. 1. 182); Moldenke,
Fifth Sum. 1: 177, 189, & 369 (1971) and 2: 649 & 912. 1971;
Moldenke, Phytologia 22: 59. 1972.

VERBENA ALATA f. ALBA Moldenke
Additional bibliography: Moldenke, Fifth Sum. 1: 177 (1971)
and 2: 912. 1971; Moldenke, Phytologia 22: 59. 1972.

VERBENA AMBROSIFOLIA Rydb.

Additional bibliography: Bolkh., Grif, Matvej., & Zakhar.
Chrom. Numb. Flow. Pl. 716. 1969; Moldenke, Phytologia 23: 182,
188, & 190. 1972.

VERBENA AMBROSIFOLIA f. EGLANDULOSA Perry
Additional bibliography: Bolkh., Grif, Matvej., & Zakhar.,
Chrom. Numb. Flow. Pl. 716. 1969; Moldenke, Fifth Sum. 1: 50,
52, 57, 61, 63, & 7h (1971) and 2: 69, 650, 658, & 912. 1971;
Noldanke, Phytologia 22: 461--462 (1972) and ans "188 & 190.
972.

21h P By Ye TO B40). Ge IA Vol. 23, .ngene

VERBENA AMOENA Paxt.
Additional bibliography: Moldenke, Fifth Summ, 1: 7h & 399
(1971) and 2: 672 & 912. 1971; Moldenke, Phytologia 22: 462. 1972.

VERBENA ANDALGALENSIS Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 200 (1971)
and 2: 912. 1971; Moldenke, Phytologia 22: 62, 1972.

VERBENA ANDRIEUXII Schau.

Additional bibliography: Moldenke, Fifth Summ. 1: 7, 205, &
369 (1971) and 2: 651 & 912. 1971; Moldenke, Phytologia 22: i63.
1972.

VERBENA ARENARIA Moldenke
Additional bibliography: Moldenke, Fifth Summ, 1: 200 (1971)
and 2: 912. 1971; Moldenke, Phytologia 22: 63. 1972.

XVERBENA ARGENTINA Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 200 (1971)
and 2: 912. 1971; Moldenke, Phytologia 22: 163. 1972.

VERBENA AURANTIACA Speg.
Additional bibliography: Moldenke, Fifth Summ. 1: 192 & 200
(1971) and 2: 912. 1971; Moldenke, Phytologia 22: 63 & 6h. 1972.

VERBENA AURANTIACA var. GLABERRIMA Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 200 (1971)
and 2: 912. 1971; Moldenke, Phytologia 22: l6l--l65. 1972.

VERBENA AURANTIACA f. ROSEA Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 200 (1971)
and 2: 912. 1971; Moldenke, Phytologia 22: 65. 1972.

VERBENA AUSTRALIS Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 177 (1971)
and 2: 912. 1971; Moldenke, Phytologia 22: 65. 1972.

xVERBENA BAILEYANA Moldenke

Additional bibliography: Moldenke, Fifth Summ, 1: 369 (1971)
and 2: 672, 673, 686, 697, 700, 702, & 912. 1971; Moldenke, Phyto-
logia 22: f6e. 1972.

VERBENA BAJACALIFORNICA Moldenke
Additional bibliography: Moldenke, Fifth Summ, 1: 7 (1971)
and 2: 912. 1971; Moldenke, Phytologia 22: 65. 1972.

VERBENA BALANSAE Briq.

Additional bibliography: Moldenke, Fifth Summ. 1: 177, 187, &
200 (1971) and 2: 702 & 912. 1971; Moldenke, Phytologia 22: 65.
OT ea

Recent collectors have found this plant fruiting in September

1972 Moldenke, Notes on Verbena 215

and December, growing in "pastizal" and on high campos with Butia
yatay (Mart.) Becc. The corollas are described as "blue" on
Krapovickas, Cristébal, Arbo, Marufiak, Marufiak, & Irogoyen 17113
and "clear lilac" on Krapovickas & Cristébal 16028.

Additional citations: ARGENTINA: Corrientes: Krapovickas &
Cristébal 16028 (Rf); Krapovickas, Cristébal, Arbo, Marufiak, Mar-

ufiak, & Irogoyen 17113 (Ac). a

VERBENA BALLSII Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 200 (1971)
and 2: 912. 1971; Moldenke, Phytologia 22: 465. 1972.

VERBENA BANGIANA Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 18) (1971)
and 2: 912. 1971; Moldenke, Phytologia 22: 65. 1972.

VERBENA BARBATA Grah,.
Additional bibliography: Moldenke, Fifth Summ. 1: 7 (1971)
and 2: 653 & 912. 1971; Moldenke, Phytologia 22: 66. 1972.

xVERBENA BEALEI Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 369 (1971)
and 2: 67), 680, & 912. 1971; Moldenke, Phytologia 22: 66. 1972.

VERBENA BERTERII f. ALBIFLORA Moldenke
Additional bibliography: Moldenke, Fifth Sum, 1: 13 & 192
(1971) and 2: 653 & 912. 1971; Moldenke, Phytologia 22: 66. 1972.

XVERBENA BINGENENSIS Moldenke

Additional bibliography: Moldenke, Fifth Sum. 1: 6) (1971)
and 2: 656, 679, & 912. 1971; Moldenke, Phytologia 22: )66--)67.
when yae

VERBENA BIPINNATIFIDA Nutt.

Additional synonymy: Verbena bipinnatifolia Nutt. ex Bolkh.,
Bean? Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 716, sphaln.
1969.

Additional & emended bibliography: Lewis & Oliv., Am. Journ,
Bot. 48: [639]—61, fig. 6. 1961; Rattenbury, Madrofio 16: 267.
1962; Bolkh., Grif, Matvej., & Zakhar., Chrom. Numb. Flow. Pl.
716. 1969; Vyas & Agarwal, Phyton 28: 1--5. 1971; Moldenke, Fifth
Sum. 1: 27, 32, 33, 36, io, 43, k5, 47, 49, 51, 52, 57, 61, 63,
7h, & 369 (is715 and 2: 521, 652-65), 658, 665, 671, 690, 708,
& 912. 1971; Moldenke, Phytologia 22: 460, 461, 4o7--li7h, & 197
(1972) and 23: 188 & 190. 1972; Vyas & Agarwal, Biol. Abstr. 53:
499. 1972; Anon., Biol. Abstr. 53 (1): B.A.S.I.C. S.267. 1972.

Emended illustrations: Lewis & Oliv., Am. Journ. Bot. 8: 60,
fig. 6. 1961.

Solbrig (1962) reports the chromosome number for this species
as 2n = 30. In this he agrees with Lewis & Oliver (1961), but

216 P Bawa 0 LOvGut A Vol. 23, nOsec

disagrees with his own previous finding which may have been based
on a misidentification or on a specimen representing one of the

many forms which have at times been given taxonomic standing. If
the latter is the case, this would be most important information.

The liadrofio (1962) reference in the bibliography above is cited
as "Darlington, Document. Chrom. Numb. Pl. 1962" by Bolkhovskikh
and his associates (1969).

Vyas & Agarwal (1971) report on the effect of thiourea and as-
corbic acid on the seed germination of this species, both indi-
vidually and jointly. The seeds are positively photoblastic and
possess embryo dormancy. Both positively photoblastic and ascor-
bic acid at 200 ppm. concentration have a promoting effect. Thi-
ourea makes the seeds light-indifferent, while ascorbic acid causes
negative photoblastic response in these seeds. Two roles of as—
corbic acid have been observed: one is the synergism with thiourea
in stimulating germination and the other is the antagonism to the
stimulatory effect of thiourea in light. It is most probable
that the plant used by these authors was V. tenuisecta priq., not
V. bipinnatifida as they claim.

The corollas are described as having been "purple" on Perino &
Perino 29. =

Additional citations: OKLAHOMA: Roger Mills Co.: Perino & Per-
ino 29 (Au--302808). MEXICO: Chihuahua: Townsend & Barber 139
(Au--2920)8) .

VERBENA BIPINNATIFIDA var. LATILOBATA Perry

Additional bibliography: Moldenke, Fifth Summ. 1: 45, 53, 57,
61, 63, 7h, & 369 (1971) and 2: 65) & 912. 1971; Moldenke, Phyto-
logia 22: 173—h7h. 1972.

XVERBENA BLANCHARDI Moldenke

Additional bibliography: Moldenke, Fiith Sum, 1: 16, 21, 3h,
38, 3, 53, & 369 (1971) and 2: 651, 654, 672—67h, 695, & 912.
1971; Moldenke, Phytologia 22: 7h. 1972.

VERBENA BONARIENSIS L.

Additional & emended bibliography: Jan, Elench, Pl. 1. 182);
Noack, Biol. Zentralbl. 57: 383--388. 1937; Rosengurtt, Estud.
Prad., Nat. Urug. 3: 325 & 326. 1943; Covas & Schnack, Revist. Ar-
gent. Agron. 14: 229. 1947; Schnack, Fehleisen, & Cocucci, Revist.
Fac. Agron. La Plata 35: 9, [54], & 55, fig. 3. 1959; Lems, Sar—
racenia 5: 79. 1960; Bolkh., Grif, Matvej., & Zakhar., Chrom.
Numb. Flow. Pl. 716. 1969; Moldenke, Phytologia 23: 182. 1972.

Additional illustrations: Schnack, Fehleisen, & Cocucci, Re-
vist. Fac. Agron. La Plata 35: [5h], fig. 3. 1959.

Walker found this species in low waste places on Okinawa. Lems
(1960) records it from thickets with Arundo donax on Gomera and
Tenerife islands of the Canaries and cites Lems 2806. Schnack and
his associates (1959) report that V. bonariensis is apomictic in
its natural reproduction. =

1972 Moldenke, Notes on Verbena 217

Additional citations: ARGENTINA: Buenos Aires: Brizuela 159
(Au--12176)) , 156 (Au--121757), 1638 (Au—121758). RYUKYO ISLAND
ARCHIPELAGO: Okinawa: E. H. Walker 8133 (W--2619388) .

VERBENA BONARIENSIS f. ALBIFLORA Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 177 & 200
(1971) and 2: 912. 1971; Moldenke, Phytologia 22: 79. 1972.

VERBENA BONARIENSIS var. CONGLOMERATA Briq.

Additional bibliography: Moldenke, Fifth Summ. 1: 16, 25, 39,
65, 92. 177, 187, 189, 200, 350, & 369 (1971) and 2: 912. 1971;
Moldenke, Phytologia 22: 78 & 79. 1972.

VERBENA BRACTEATA Lag. & Rodr.

Additional & emended bibliography: Jan, Elench. Pl. 1. 182;
Reiche & Phil., Fl. Chil. 5: 296. 1910; Graves, Eames, Bissell,
Andrews, Harger, & Weatherby, Bull. Conn. Geol. & Nat. Hist. Surv.
14: (Cat. Flow. Pl.] 331. 1910; Noack, Biol. Zentralbl. 57: 383—
388. 1937; Rattenbury, Madrofio 15: 51 (1959) and 15: 220. 1960;
Bolkh., Grif, Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 716.
1969; Moldenke, Fifth Summ. 1: 15--19, 21--23, 27, 30, 32, 3h,
36-13, 45, 47, h9—51, 53, 57, 63--65, 75, 203205, 369, & 396
(1971) and 2: 656, 657, 659, 66), 672, 673, 677, 679, 691, 693,
697, 698, 705, 736, 766, 913, & 967. 1971; Moldenke, Phytologia
22: 61, 467, 479--488, 490, 499, & 501 (1972) and 23: 188, 195,
& 196. 1972.

The Madrofio (1959, 1960) references given in the bibliography
above are sometimes attributed to Solbrig, while Bolkhovskikh and
his associates (1969) cites them as "Darlington, Document. Chrom.
Numb. Pl....'' Actually they appear in the periodic continuation
of a series started by Rattenbury, with no more subsequent com-
piler mentioned.

The corollas of V. bracteata are described as "very pale-
violet, tube purplish" on Twisselmann 898. Twisselmann reports
the plant as "locally common in low areas flooded by May and June
storms in the Lower Sonoran grasslands" of Kern County, Califor-
nia. The R. MM. Stewart 1180, distributed as V. bracteata, is ac-
tually V. plicata Greene.

Additional citations: CALIFORNIA: Kern Co.: Twisselmann 89),8
(Au--297797) .

VERBENA BRACTEATA f. ALBIFLORA (Cockerell) Moldenke

Additional bibliography: Moldenke, Fifth Summ. 1: 51 & 64
(1971) and 2: 657 & 913. 1971; Moldenke, Phytologia 22: 88.
1972.

VERBENA BRASILIENSIS Vell.

Additional & emended bibliography: Lewis & Oliv., Am. Journ.
Bot. 8: [639]--6h1, fig. 22. 1961; Bolkh., Grif, lMatvej., & Zak-
har., Chrom. Numb. Flow. Pl. 716. 1969; Moldenke, Phytologia 23:
182. 1972.

218 Pi Yer OL OrGe ink Vol. 23, no. 2

Emended illustrations: Lewis & Oliv., Am. Journ. Bot. 8: 60,
si Jp ocd ke /o ale

Recent collectors describe this as an erect perennial herb, to
1.5 m, tall, growing in hedgerows and cultivated fields. Ellison
found it an "abundant population on roadbanks of train track in
disturbed habitat, full sun" in North Carolina. The corollas
were "lavender" on Ellison 817 and "purple" on Mexia 5275.

Additional citations: BRAZIL: Minas Gerais: Mexia 5275 (Au--
121763). SHo Paulo: I. S. Gottsberger 1)3 [h] (Ft).

VERBENA BRASILIENSIS var. SUBGLABRATA Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 192 (1971)
and 2: 913. 1971; Moldenke, Phytologia 22: 90. 1972.

VERBENA CALIFORNICA Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 65 & 370
(1971) and 2: 913. 1971; Moldenke, Phytologia 22: 490--91. 1972.

VERBENA CALLIANTHA Briq.

Additional bibliography: Moldenke, Fifth Summ. 1: 177, 18h,
187, & 200 (1971) and 2: 658, 662, 667, 683, & 913. 1971; Molden-
ke, Phytologia 22: 91. 1972.

VERBENA CALLIANTHA var. MICROSOMA Briq.
Additional bibliography: Moldenke, Fifth Summ. 1: 187 (1971)
and 2: 913. 1971; Moldenke, Phytologia 22: hOl. 29726

VERBENA CAMERONENSIS L. I. Davis

Additional & emended bibliography: Lewis & Oliv., Am. Journ.
Bot. 48: (639]--6h1, fig. 7. 1961; Bolkh., Grif, Matvej., &
Zakhar., Chrom. Numb. Flow. Pl. 716. 1969; Moldenke, Fifth Summ.
1: 57 & 75 (1971) and 2: 658, 682, & 913. 1971; Moldenke, Phyto-
logia 22: 491--92. 1972.

Emended illustrations: Lewis & Oliv., Am. Journ. Bot. 8: 60,
fags. (eye dole

Johnston and Davis collected this plant on the coastal low-
lands of Veracruz, Mexico, in 196 and comment that this was "ap-
parently the first record from Veracruz", but this statement is
obviously incorfect since the species was collected in that Mexi-
can state by Henri Guillaume Galeotti in 180 -- 106 years ear-
lier!

Additional citations: MEXICO: San Luis Potosf: Davis & Johns-
ton s.n. [13 April 1946] (Au--278272). Veracruz: Johnston & Da-

vis s.n. [20 April 196] (Au--278281).

VERBENA CAMPESTRIS Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 177 (1971)
and 2: 912. 1971; Moldenke, Phytologia 22: 92. 1972.

VERBENA CANADENSIS (L.) Britton
Additional & emended bibliography: Jan, Elench. Pl. 1. 182);

1972 Moldenke, Notes on Verbena 219

Noack, Biol. Zentralbl. 57: 383--388. 1937; Rattenbury, Madrofio
15: 51. 1959; Bolkh., Crif, Matvej., & Zakhar., Chrom. Numb. Flow.
Pl. 715 & 716. 1969; Moldenke, Phytologia 23: 183. 1972.

There is disagreement among cytologists as to the chromosome
number in this species. It is given as 30 by Junell (193k), Der-
men (1936), Noack (1937), and Solbrig (1959), but as 10 by Sharma
& Mukhopadhyay (1963). It would be most interesting to see if
herbarium vouchers were kept by these workers and then to ascertain
whether the last-mentioned report is not perhaps based on a mis-
identification. Verbena tenuisecta Briq., a commonly cultivated
and escaped species, is the one that is most often misidentified
in India.

The Madrofio reference cited in the bibliography above is often
credited to Solbrig, while Bolkhovskikh (1969) cites it as "Dar
lington, Document. Chrom. Numb. Pl...."

The corollas are described as having been "purple" on Perino &
Perino 377 and these collectors describe the plant as "common" in
Pushmataha County, Oklahoma.

The Kibler 26, distributed as V. canadensis, is actually V.

halei Small.
Additional citations: OKLAHOMA: Pushmataha Co.: Perino & Peri-
no 377 (Au--302806) .

VERBENA CANADENSIS f. CANDIDISSIMA (Haage & Schmidt) Palmer &
Steyerm.
Additional bibliography: Moldenke, Fifth Summ, 1: 3, 5, 58,
& 370 (1971) and 2: 653, 653, 659, 78h, & 913. 1971; Moldenke,
Phytologia 22: 498. 1972.

VERBENA CANESCENS H.B.K.

Additional bibliography: Lewis & Oliv., Am. Journ. Bot. ),8:
(639]-& 641. 1961; Bolkh., Grif, Matvej., & Zakhar., Chrom.
Numb. Flow. Pl. 716. 1969; Moldenke, Fifth Summ. 1: 51, 58, & 75
(1971) and 2: 659--661, 68), 694, 767, 769, & 913. 1971; Molden-
ke, Phytologia 22: 85 & hob—SOl. 1972.

Lewis & Oliver (1961) report the diploid chromosome number for
this species as 1). Recent collectors have found it growing in
matorral formation in Mexico. The corollas are described as
"purple" on Ventura A. 157h.

Additional citations: MEXICO: Oaxaca: Johnston & Davis s.n.
(27 July 1947] (Au--278287). Puebla: Ventura A. 157) (Au—
291,666) . fri toes

VERBENA CANESCENS f. ALBIFLORA Moldenke
Additional bibliography: Moldenke, Fifth Summ. 1: 75 (1971)
and 2: 913. 1971; Moldenke, Phytologia 22: 500. 1972.

VERBENA CANESCENS var. ROEMERIANA (Scheele) Perry
Additional & emended bibliography: Lewis & Oliv., Am. Journ.
Bot. 48: [639]--61, fig. 13. 1961; Moldenke, Fifth Summ. 1: 58,

220 PHY f @ L0°G T's Vol. 23, U0s1e

65, & 75 (1971) and 2: 659--661, 693, 702, & 913. 1971; Moldenke,
Phytologia 22: 485 & 499--501. 1972.

Emended illustrations: Lewis & Oliv., Am. Journ. Bot. 8: 60,
free 1356 190M,

Recent collectors describe this plant as a perennial and found
it growing on gravelly knolls. The corollas are described as
having been "blue" on Rollins & Tryon 58)1.

Additional citations: MEXICO: Nuevo Leén: Rollins & Tryon 58)1
(Au--300250) .

VERBENA CANIUENSIS Moldenke
Additional bibliography: Moldenke, Fifth Sum, 1: 177 (1971)
and 2: 660 & 913. 1971; Moldenke, Phytologia 22: 501. 1972.

VERBENA CAROLINA L.

Additional bibliography: Jan, Elench. Pl. 1. 182; Lewis &
Oliv., Am. Journ. Bot. 8: [639}—6)1, figs 1). 196); eBollkings
Grif, Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 716. 1969; Mol-
denke, Phytologia 23: 183--186. 1972.

mended illustrations: Lewis & Oliv., Am. Journ. Bot. 8: 6)0,
fig. 1h. 1961.

Runyon describes this plant as "common in waste places" in the
state of México. The corollas are said to have been "purple" on
R. Runyon 1360.

Additional citations: MEXICO: México: R. Runyon 1360 (Au--
290200). Oaxaca: Johnston & Davis s.n. [1 July 197] (Au-—-278268).

VERBENA CILIATA Benth.

Additional & emended bibliography: Lewis & Oliv., Am. Jqurn.
Bot. 8: [639]--641, fig. 8. 1961; Bolkh., Grif, Matvej., & Zak-
har., Chrom. Numb. Flow. Pl. 716. 1969; Moldenke, Phytologia 23:
186--190 & 192. 1972.

Emended illustrations: Lewis & Oliv., Am. Journ. Bot. )8: 60,
fies GO. 1L96l,

The J. Rzedowski L3hh, distributed as V. ciliata, is actually
Vv. elegans H.BoKe

Additional citations: MEXICO: Chihuahua: Pennington 42 (Au--
287710). Nuevo Leén: Kenoyer C C.u8 (Au--122037).

VERBENA CLOVERAE Moldenke

Additional & emended bibliography: Lewis & Oliv., Am. Journ.
Bot. 8: [639]--6h1, fig. 15. 1961; Bolkh., Grif, Matvej., & Zak-
har., Chrom. Numb. Flow. Pl. 716. 1969; Moldenke, Phytologia 23:
190 & 192--193. 1972.

VERBENA CORYMBOSA Ruiz & Pav.

Additional bibliography: Bolkh., Grif, Matvej., & Zakhar.,
Chrom. Numb. Flow. Pl. 716. 1969; Moldenke, Phytologia 23: 19).
MOV Ar

1972 Moldenke, Notes on Verbena 221

VERBENA CUMINGII Moldenke
Additional bibliography: Moldenke, Phytologia 13: 191. 1966;
Moldenke, Fifth Summ. 1: 192 (1971) and 2: 621 & 91). 1971.

VERBENA DELTICOLA Small

Additional & emended bibliography: Lewis & Oliv., Am. Journ.
Bot. 48: (639]--641, fig. 9 & 10. 1961; Bolkh., Grif, Matvej., &
Zakhar., Chrom. Numb. Flow. Pl. 716. 1969; Woldenke, Phytologia
32,196. 1972.

Emended illustrations: Lewis & Oliv., Am. Journ. Bot. 48: 6)0,
fig. 9 & 10. 1961.

Recent collectors have found this plant growing in roadside
ditches, in mountain meadows, and in Acacia-Prosopis communities,
often in dark soil. Runyon describes it as an "erect or ascen-
ding herb, in open ground, occasionally along highways, clay soil,
the flowers with no odor, blooming in April and May, the leaves
nearly trifid, ovate in outline, fruit a small capsule [actually
it is a schizocarp], bark green, roots fibrous" and "flowers in
terminal panicles [actually spikes], growing in sandy loam of
open ground and fields, the leaves ovate in outline, lobed, abun-
dant, forms colonies sometimes covering large areas."

Fleetwood reports that it is "common in dry clayey loam" in
Hidalgo County, Texas, while Ton has found it on shrubby and
steep heavily wooded slopes in Chiapas -- the latter a notable
southward extension of the known range of the species! Sikes &
Babcock report it as "abundant" in Sonora. Rickett (1969) avers
that it blossoms in Texas from February to August, growing "in
sandy or clay soil in open ground, often very profuse, in south-
ern Texas and northern ifexico". He describes the corollas as
ranging "from pink to magenta, lavender, and purple". Pennington
tells us that in Sonora the plant is boiled and salted.

The corolla is described as "light-blue" on Sikes & Babcock
161, "pink-purple" on C. L. Lundell 10680, "red" on R. Runyon
2587, and "purple" on Re, Runyon eben seet

~~ Material of Vv. delticola has been misidentified and distribu-
ted in herbaria as Vv. stricta Vent., while the Pladeck SNe [Lay
5, 1940], distributed as V. delticola, is actually V. cloverae
Moldenke and Pennington ar is xV. me Voss.

Additional citations: TEXAS: Cameron Co.: C C. L. Lundell 10680
(N); Re Runyon 2587 (Au--268721). Hidalgo Co.: Fleetwood 8157
(W--2518783) ; Re Runyon 2625 (Au--268719). MEXICO: Chiapas: T Ton
1863 (2), 2210 ) (Rf). Coahuila: E.G. Marsh 118), (Au--212082),
1592 (Au—-213301), 2147 (Au--213883). Hidalgo: V. H. Chase 7116
(MN); Johnston & Davis s.n. [17 April 196] (Au--2 278290). Ni Nuevo
Leén: Atwood 2059 (N) (N); Dia Diaz L. sen. [10/IX/1959] (Ip), s.n. [16.
1X.1959] (Ip (Ip); Le BiG Higgins 2683 83 (Mi); Johnston & Davis s.n. [7
April 19),6] (Au—-278279, Au—-278291) ; H. M. Parker 501 (iaea=
302752). Sonora: Pennington 312 (Au--26257); Sik Sikes 2s & Babcock

222 P YoT@) b/O"G ah Vol. 23, m0. 2

161 (Au--259161, Au--2845)0), Tamaulipas: Stanford, Lauber, &
Taylor 2100 (Se--1)9132).

VERBENA DELTICOLA f. LILACINA L. I. Davis

Additional synonymy: Verbena delticola lilacina L. I. Davis ex
Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex. Res. Found.
Bot. 6:] "1876. 1970.

Additional bibliography: Moldenke, Phytologia 9: 64—65. 1963;
Moldenke in Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex.
Res. Found. Bot. 6:] 1316 & 1323. 1970; Correll & Johnston, Man.
Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:2] 1876. 1970;
Moldenke, Fifth Summ. 1: 58 (1971) and 2: 665 & 91). 1971.

VERBENA DEMISSA loldenke
Additional bibliography: Moldenke, Phytologia 13: 27. 1966;
Moldenke, Fifth Summ. 1: 137 (1971) and 2: 91h. 1971.

This plant has been collected at altitudes of 2550 to 3700
meters, flowering and fruiting in April, July, and September. The
Rose, Pachano, & Rose 22851, cited below, was previously misiden-
tified and ed as V. “glabrata H.B.K. Material has also been
misidentified and distributed in some herbaria under the name V.
polystachya H.B.K. mw

Additional citations: ECUADOR: Azuay: Asplund 17801 (N); Rose,
Pachano, & Rose 22851 (N, wW--1022h8). Cafiar: J&tiva & Epling 260

(ie

XVERBENA DERMENI lioldenke

Additional bibliography: Moldenke, Phytologia 10: 10). 196);
Moldenke, Résumé Suppl. 17: 3. 1968; Moldenke, Fifth Summ. 1: 13,
18), 187, 200, & 370 (1971) and 2: 655, 665, 674, & 914. 1971.

Steinbach describes this plant as "locally common", with the
corollas "bright-violet, rose-purple at base outside". It has been
collected in fruit in February and April, in addition to the months
previously reported. Vargas Calderén 19,93 is a mixture with Vv.
parvula var. gigas Moldenke

Additional citations: PERU: Arequipa: Lépez Guillén 29 (Rf);
Vargas Calderén 1993, in part (Ac). Ayacucho: Soukup 5501 (Z).
BOLIVIA: Cochabamba: R. F. Steinbach 6) (N). ARGENTINA: Formosa:
I. Morel 614 (N).

VERBENA DISSECTA Willd.

Additional & emended synonymy: Verbena laciniata Kuntze ex liol-
denke, Suppl. List Invalid Names 9, in syn. 19h1 [not V. laciniata
Briq., 1960, nor (L.) Briq., 190, nor (Lam.) Briq., 1939, nor Rates
1833, nor Sessé & Moc., 1940]. Glandularia dissecta (Willd. ex
Spreng.) Schnack & Covas apud J. A. Clark, Card Ind. Gen. Sp. Var.
issue 183. 19h).

Additional & emended bibliography: Reiche & Phil., Fl. Chil. 5:
289 & 292. 1910; J. A. Clark, Card Ind. Gen. Sp. Var. issue 183.

1972 Moldenke, Notes on Verbena 223

1944; Cabrera, Man. Fl. Alred. Buenos Aires 396--398, fig. 148 f—
n. 1953; J. F. Macbr., Field Mus. Publ. Bot. 13 (5): 614, 619, &
629. 1960; Martinez-Crovetto, Bonplandia 1: 291 & 31h. 196); An-
gely, Fl. Anal. Paran., ed. 1, 571. 1965; Martfinez-Crovetto, Bon-
plandia 2: 19. 1965; Troncoso in Cabrera, Fl. Prov. Buenos Aires
5: 132 & 138. 1965; Meyer & Weyrauch, Inst. Mig. Lill. Misc. 23:
6h & 123. 1966; Martinez-Crovetto, Bonplandia 2: 130. 1967; Mol-
denke, Phytologia 16: 51. 1968; Solbrig, Passani, & Glass, Am.
Journ, Bot. 55: 1238. 1968; Bolkh., Grif, Matvej., & Zakhar.,
Chrom., Numb. Flow. Pl. 715 & 716. 1969; R. F. V. Cooper in Pas-
tore, Bol. Soc. Argent. Hort. 157: 12) & 125. 1969; Schnack & Ru-
bens, Bol. Soc. Argent. Bot. 13: 206. 1970; Moldenke, Fifth Summ,
1: 177, 18h, 189, 192, 200, & 370 (1971) and 2: 521, 621, 655,
662, 666, 667, 678, 700, 708, & 91). 1971.

Additional illustrations: Cabrera, lian. Fl..Alred. Buenos Aires
396, fig. 148 f--n. 1953; R. F. V. Cooper in Pastore, Bol. Soc.
Argent. Hort. 157: 12). 1969.

It should be pointed out here that the Verbena laciniata (H.B.K.)
Briq., referred to in the synonymy above, is a synonym of V. lac-
iniata (L.) Briq., while V. laciniata Raf. is probably xV. perri-
ana Moldenke.

Macbride (1960) credits V. dissecta to "Willd. ex Schauer in
DC. Prodr. 11: 552. 1847", but the name actually goes back to
Sprengel in L., Syst. Veg., ed. 16, 2: 750 (1825). He cites a
Meyen s.n. from Pisacoma, Puno, Peru, but I doubt very much that
V. dissecta occurs at all in Peru. Meyen's plant is probably V.
laciniata (L.) Briq., as liacbride himself suggests. He gives the
distribution of V. dissecta as "To Uruguay. Chile; Bolivia; Braz-
il". Troncoso (1965) gives its distribution as "Brasil meridio-
nal, Uruguay y Argentina. En praderas arenosas y en las barrancas
del Parand y del litoral bonaerense". She cites Burkart 3082 and
Rodrigo 2339 in the San Isidro herbarium,

Martinez-Crovetto (196), 1965) record the Chaco Amerind names
"imal4k labé (hoja azul)" and "uompé chak(o)lfp" for this species;
Cooper (1969) calls it "lilac verbena". The corollas are descri-
bed as "rose" on 3. L. Miller 152, 190, 203, & 209, "lilac" on
Collector undetermined 311 and Luna 132, 21, & 547, "purple" on
R. M. Aguilar 286, "bluish" on Herb. Inst. Miguel Lillo 981,
"blue" on Olea L, "blue-violet" on Semper 138, "reddish" on B. Le
Muller 165, "bluish-lilac" on Varela 0h, and "violet" on P. Gar-
cia s.n.

Additional citations: BRAZIL: Rio Grande do Sul: Palacios &
Cuezzo 2519 (N). ARGENTINA: Catamarca: Brizuela 628 (N), 1037
(N); B. L. Miller 152 (N), 165 (N), 190 (N); O'Donell s.n. [IV-}]
(N); Pierotti 11529 [Herb. Inst. Liguel Lillo 28312] (N), "H" [12/
TII/4k) (N), "H" [15/vV/44) (N), "h" [La Merced, 19/V/l] (N).
Chaco: Buratovich 69 (N). Cérdoba: Varela Oh (N); Villafaflie 120
(Se--130302), 642 (N). Formosa: I. Morel 4391 (N); Pierotti 6543

22h Peer On 01g a Vol. 23, mone

(N). La Rioja: B. L. Muller 203 (N), 209 (N). Mendoza: Semper
138 (N). Salta: R. M. Aguilar r 286 (N)3 L Luna 132 (N), 547 ~(N)5 1 Me
R. Malv4rez 179 (N), 302 (N); O'Donell 3094 (N). Santiago del Es-
tero: Balegno 0 103 (N); P P. Garcia s.n. [Herb. Inst. Miguel Lillo
71h] (N), sen. “Herb. Inst. Miguel Lillo 981) (N); Luna 2)1 (N);
T. Meyer 6838 (N); Pierotti "H" [11/IV/1944] (N), "h" h" (N)5 Viet-
Martinez sen. [Herb. Inst. Miguel Lillo 5] (N). Tucun4n: Collec-
tor undetermined 311 (N); M. R. Malvdrez 70 (N); Olea 4 (N), 190
(M); Rocha 3818 (Au--122073).

VERBENA DISSECTA f. ALBA Moldenke
Additional bibliography: Moldenke, Phytologia 16: 51. 1968;
Moldenke, Fifth Summ. 1: 200 & 370 (1971) and 2: 914. 1971.

XVERBENA DISSOLUTA Moldenke

Additional bibliography: Moldenke, Phytologia 11: 55. 1965;
Moldenke, Fifth Summ. 1: 370 (1971) and 2: S2l, 667, 678.nec0.
700, & crie 1971.

VERBENA DOMINGENSIS Urb.

Additional bibliography: Ciferri, Mycopathologia 7: 89. 195h;
Hansford, Sydowia Ann. liyc., ser. 2, Beih. 2: 68). 1961; Molden-
ke, Phytologia Tals NSE alles Moldenke, Fifth Summ. 1: 98 & 370
(1971) and 2: 666, 686, & 914. 1971.

Recent collectors describe this plant as slender, herbaceous,
erect, and 30--50 cm. tall, or sometimes decumbent, the rhizome
creeping. The corollas are described as having been "violet" on
Gastony, Jones, & Norris 363 and Liogier 12077 & 12197 and "blue"
on Holdridge 83h and Jiménez 4831. The species has been found
along trails through rocky p: pine ne forests; Liogier found it "in
open places near rivers.....in mostly secondgrowth vegetation"
and calls it "common in pine barrens in rocky mountains". It has
been collected at altitudes of 500 to 2710 meters. The cultiva-
ted material from California was grown from seeds of Liogier
12077 and 12197.

Ciferri (195) records the fungus, Meliola aciculosa var.
verbenae Cif., from this host plant on the basis of Ekman 3161
from Hispaniola.

Additional citations: HISPANIOLA: Dominican Republic: Gastony,
Jones, & Norris 363 (N, N); Jiménez 1,831 (N); Liogier 12077 (N,
N, N, 2), 12197 (N, Rf). Haiti: Holdridge 834 (Sd—-618 a
CULTIVATED: ): California: A. R. Moldenke 3395 ( (Ac, Z).

VERBENA DUSENII Moldenke

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 571.
1965; Moldenke, Phytologia 13: 19). 1966; Moldenke, Fifth Summ. 1:
WI CEOL) and. 2: 91). 1971.

Hatschbach 17637 is described by the collector as having had
violet flowers "dos brejos".

1972 Moldenke, Notes on Verbena 225

Additional citations: DRAZIL: Parand: Hatschbach 17637 (Ft, W—
253653)).

VERBENA EHRENBERGIANA Schau.

Additional & emended bibliography: Howell & McClintock in
Kearney & Peebles, Ariz. Fl., ed. 2, 726 & 727. 1960; Lewis &
Oliv., Am. Journ. Bot. 8: [639]. 1961; Hocking, Excerpt. Bot. A.
6: 91. 1963; Moldenke, Phytologia 16: 186--187. 1968; Bolkh.,
Grif, Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 716. 1969; Gib-
son, Fieldiana Bot. 2) (9): 232. 1970; Rickett, Wild Fls. U.S.

4 (3): S40 & 799. 1970; Moldenke, Fifth Summ. 1: 63 & 75 (1971)
and 2: 667 & 914. 1971; Moldenke, Phytologia 23: 18). 1972.

The only record of this species from the United States is a
sheet of Stalmach 198 in the herbarium of the University of Texas
labeled merely "Arizona". Gibson (1970) states that all Guate-
malan specimens so identified are actually V. carolina L.

Verbena ehrenbergiana has been found growing in association
with Cerastium, Llavea, Penstemon, and Romanschultzia in Hidalgo,
Mexico. Sullivan found it growing in humid Quercus-Liquidambar
forests on limestone karst topography in Tamaulipas, flowering in
June.

Material of this species has frequently been misidentified
and distributed in some herbaria under the names V. urticaefolia
L. or V. urticifolia L. On the other hand, the Mears 192, dis-
tributed as V. ehrenbergiana, is actually V. litoralis H.B.K.

Additional citations: MEXICO: Hidalgo: Mears 299a (Au——-25)970) .
Tamaulipas: J. R. Sullivan 09 (Au--298185), 475 (Au—298162),
554 (Au--298174). State undetermined: Schnée s.n. [Mexique, VI-
X) (Mi, N, S, W--25),6669).

VERBENA ELEGANS H.B.K.

Additional synonymy: Verbena elegans Humb. & Bonpl. ex Steud.,
Nom. Bot. Phan., ed. 1, 873. 1821. Glandularia elegans H.B.K.
ex Solbrig, Passani, & Glass, Am. Journ. Bot. 55: 1235. 1968.
Verbena elegans (H.B.K.) Raf. ex Moldenke, Résumé Suppl. 18: 1k,
in syn. 1969. Glandularia elegans Solbrig, Princ. & Meth. Pl.
Biosystem. 148. 1970.

Additional bibliography: Steud., Nom. Bot. Phan., ed. 1, 873.
1821; Lewis & Oliv., Am. Journ. Bot. 48: [639] & 641, fig. ll.
1961; Hocking, Excerpt. Bot. A.6: 91. 1963; Moldenke, Phytologia
16: 187. 1968; Solbrig, Passani, & Glass, Am. Journ. Bot. 55:
1235--1239. 1968; Bolkh., Grif, Matvej., & Zakhar., Chrom. Numb.
Flow. Pl. 716. 1969; Solbrig, Passani, & Glass, Biol. Abstr. 50:
4151. 1969; Synge, Suppl. Dict. Gard., ed. 2, 548. 1969; El-
Gazzar & Wats., New Phytol. 69: 83 & 485. 1970; Moldenke in Cor-
rell & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex. Res. Found.
Bot. 6:] 1316, 1317, & 1323. 1970; Correll & Johnston, Man. Vasc.
Pl. Tex. (Contrib. Tex. Res. Found. Bot. 6:] 1876. 1970; Solbrig,
Princ. & Meth. Pl. Biosystem. 18. 1970; Moldenke, Fifth Sum. 1:

226 P) HeXeT, O. LoOuGo Duck Vol. 23, mos 2

63, 75, 205, & 370 (1971) and 2: 521, 658, 667, 679, 684, & 91h.
1971; Moldenke, Phytologia 22: 61 & 71. 1972.

Recent collectors have encountered this plant on very wet
slopes, riolitic slopes with disturbed oak vegetation, coniferous
woods, in oak forests on wet mountainsides, on grassy slopes with
Quercus, and on limestone mountains, as well as in oak woods and
in Abies woods, at altitudes of 1000 to 3230 meters. Mears found
it growing in association with Cornus, Drymaria, Dudleya, Heuchera,
Lobelia, Maurandya, Piqueria, and Reseda, the Andersons refer to
it as a "common roadside herb", and Beaman found it to be "common
on open meadows in pine forests". It has been collected in fruit
in November, in addition to the months previously reported in
these notes. The vernacular name, "verbena", has been recorded
for it. A tea made from the leaves is taken for stomach disorders
in Chihuahua,

The corollas are said to have been "lavender" on Ton 1113,
"mauve" on Anderson & Anderson 658, "purple" on Diaz L Luna 11 and
J. Rzedowski 7971, and "brilliant crimson-lavender" on Beaman
270. Lewis & Oliver (1961) report the chromosome number as

= 30.

Solbrig and his associates (1968, 1969, 1970) have crossed
this species with material which they have identified as V. peru-
viana (L.) Britton, V. stellarioides Cham., and V. pulchella
Sweet, for all of which, as well as for the hybrids, they employ
the generic name Glandularia, although in some cases I cannot
find that the combinations have ever been proposed validly in ac-—
cord with the present very complicated International Rules of
Botanical Nomenclature. "Analysis of the behavior of chromosomes
at meiosis in the hybrid indicated that the chromosomes of G.
pulchella are similar to one of the genomes of G. elegans. ~ Non-
homologous pairing and presence of multivalents also indicated
that G. elegans is a segmented allohexaploid derived most likely
from So. Amer. ancestors." They found pure V, elegans seeds to
show 99 percent fertility, but the hybrid from less than 5 to
over 70 percent.

Material of V. elegans has been misidentified and distributed
in some herbaria under the names V. elegans var. asperata Perry
and V. prostrata R. Br. On the other hand. hand, the Detling 962 and
Je Rzedowski 21919, distributed as V. elegans, are re actually | Ve.
bipinnatifida lutt., while Johnston & Davis s.n. [7 April 196]
and H. M. Parker 501 are V. delticola Small and S. Sanderson 262
is V. elegans var. asperata Perry.

Additional citations: MEXICO: Chiapas: Ton 111) (N, W--
2556725), 1489 (N). Federal District: Huerta sen. (8/VIII/1962]
(Ip). Hidalgo: Anduaga A. 3 (N); M. R. Diaz ipiaz 27 (Ac); Gonzdlez
Quintero 1933 (Ac), 2167 (. (Ip, RS) 2710 (Ip), 2868, in part (Mi);
Hidalgo & Anda s.n. [25/VI/1967] (Mi, Mi, Mi); L. I. Davis sen.

1972 Moldenke, Notes on Verbena 227

[7 July 1944] (Au--278267); Mears 265a (Au--255309), 307a (Au--
256227); Medellin s.n. [7/VIII/1954] (Ip); J. Rzedowski 1 17002
(Au--2),3525); Vela G. 1397 (Ip). Michoac4n: Diaz Luna 11 1 (Ip).
Nuevo Leén: Beaman 270) 1, (Au--2),0747, W--257537h) 5 Johnston & Davis

sen. (21 April 19k6é] (Au--278283). San Luis Potosf: Anderson & &
Anderson 4658 (Mi); Johnston & Davis s.n. [16 June 197] (Au--—
278273, Au--27827h); J. Rzed Je Rzedowski 13), (Au--169899), 7971 (Ip).
Tamaulipas: M. B. Webster 138 (Au--241511).

VERBENA ELEGANS H.B.K. x V. PERUVIANA (L.) Britton ex Moldenke,
Fifth Summ. 1: 370 (1971) and 2: 91. 1971.

Synonymy: Glandularia elegans x peruviana Solbrig in Solbrig,
Passani, & Glass, Am. Journ. Bot. 55: 1236--1238. 1968.

Bibliography: *Solbrig, Passani, & Glass, Am. Journ. Bot. 55:
1236--1238. 1968; Solbrig, Passani, & Glass, Biol. Abstr. 50:
4151. 1969; Solbrig, Princ, & Meth. Pl. Biosystem. 148. 1970;
Moldenke, Fifth Summ, 1: 370 (1971) and 2: 91) & 970. 1971.

Solbrig and his associates (1968, 1969, 1970) speak of the
artificial crosses which they have made between these two spe-
cies, producing a tetraploid hybrid which showed less than 5
percent fertility and a segmental allooctoploid hybrid which
showed over 70 percent fertility. A binomial designation has
not yet been proposed for these hybrids.

VERBENA ELBGANS H.B.K. x V. PULCHELLA Sweet ex lioldenke, Fifth
Summ. 1: 370 (1971) and 2: 91) & 970. 1971.

Synonymy: Glandularia elegans x pulchella Solbrig in Solbrig,
Passani, & Glass, Am. Journ, Bot. 55: 1236--1238. 1968. Verbena
pulchella Sweet x V. elegans H.B.K. ex Moldenke, Fifth Sum. mm. Ll:

370 (1971) and 2: 919 & 970. 1971.

Bibliography: Solbrig, Passani, & Glass, Am. Journ. Bot. 55:
1236--1238. 1968; Solbrig, Passani, & Glass, Biol. Abstr. 50:
4151. 1969; Solbrig, Prine. & lieth. Pl. Biosysten. 148. 1970;
Moldenke, Fifth Summ. 1: 370 (1971) and 2: 91) & 970. 1971.

Solbrig and his associates (1968, 1969, 1970) speak of the
crosses which they have made between these two species, producing
a tetraploid hybrid showing less than 10 percent fertility. I am
not certain whether their concept of V. pulchella is the same as
mine or is that of some modern Argentine botanists, so it seems
best not to propose a binomial name for their hybrid until the
exact parentage is verified by examination of herbarium vouchers.

VERBENA ELEGANS H.B.K. x V. STELLARIOIDES Cham. ex Moldenke,
Fifth Summ. 1: 370 (1971) and 2: 914 & 970. 1971.
Synonymy: Glandularia elegans x stellaroides Solbrig in Sol-
brig, Passani, & Glass, Am. Journ. Bot. 55: 1236 & 1237. 1968.
Bibliography: Solbrig, Passani, & Glass, Am. Journ, Bot. 55:
1236 & 1237. 1968; Moldenke, Fifth Summ. 1: 370 (1971) and 2:
914 & 970. 1971.

228 P HYT.0 LeOsG 4 Vol. 23, NOs 2

Solbrig and his associates (1968) speak of the hybrid which
they have produced between these two species, but it seems best
not to propose a binomial designation for this hybrid until the
parentage can be verified by examination of herbarium vouchers.
It is possible that one of the parents may have been V. sessilis
(Cham.) Kuntze instead of V. stellarioides Cham., since the dif-
ferences between these two species are not always obvious.

VERBENA ELEGANS var. ASPERATA Perry

Additional synonymy: Verbena elegans var. asperate Perry ex
Moldenke, Résumé Suppl. 18: 14, in syn. 1969.

Additional & emended bibliography: Lewis & Oliv., Am. Journ.
Bot. 48: [639] & 641, fig. 11. 1961; Turrill in Curtis, Bot. Mag.
17: pl. 409. 1963; Hocking, Excerpt. Bot. A.6: 91. 1963; Mol-
denke, Phytologia 15: 492 & 493 (1968) and 16: 62. 1968; Synge,
Suppl. Dict. Gard., ed. 2, 58. 1969; Moldenke in Correll &
Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:]
1316, 1317, & 1323. 1970; Correll & Johnston, Man. Vasc. Pl. Tex.
[Contrib. Tex. Res. Found. Bot. 6:] 1876 & 1877. 1970; Moldenke,
Fifth Sum. 1: 58, 75, 20h, 205, & 370 (1971) and 2: 667 & 914.
1971; Moldenke, Phytologia 22: 61. 1972.

Additional & emended illustrations: Lewis & Oliv., Am. Journ.
Bot. 8: 651, fig. 11. 1961; Turrill in Curtis, Bot. Mag. 17k:
pl. 409 [as V. wrightii] (in color). 1963.

Recent collectors have encountered this variety along road-
sides, by streams, and in both Quercus and Abies woods. Breedlove
found it on slopes with Arbutus, Pinus, and Quercus, while with
Kawahara he encountered it on wooded banks of streams with Alnus
sp., Padus serotina, Pinus engelmanni, and Quercus spp. Rzedowski
reports it from "ladera margosa con vegetacién de matorral secun-
dario abierto", "ladera caliza con vegetacién de zacatal", "ladera
caliza con vegetacién de encinar", "ladera riolitica con vegeta-
cién de bosque de pino-encino", and "matorral de Cordia boissieri'!
It has been collected at altitudes of 710 to 3000 meters, and, in
addition to the months previously reported by me, in flower and
fruit in November.

The corollas are described as "lavender" on Breedlove 15558,
"deep-rose" on Breedlove & Kawahara 16890, and "purple" on Breed-
love 1)18 and J. Rzedowski 2)),93 & 282. Lot describes the
plant as herbaceous, to 0 cm. tall, and notes "flor lila".

The color plate presented by Turrill (1963) as illustrative
of V. wrightii A. Gray is very obviously V. elegans var. asperata
instead. a

Material of V. elegans var. asperata has been misidentified
and distributed in some herbaria under the name V. prostrata R.
Br. On the other hand, the Mendiola s.n. [Saltillo Torreén],
distributed as V. elegans var. asperata, is really V. ambrosifolia
Rydb., while J. Rzedowski 7971 is typical V. elegans H.B.K.

Additional & emended citations: MEXICO: Chiapas: Breedlove

1972 Moldenke, Notes on Verbena 229

Ups18 (N, Ws). Hidalgo: Ch4évez 0. s.n. [.VIII.63] (Ip); Gonzd-
lez lez Quintero 2868, in part (Ip), 3252 (1 (Rf), 3288 (Ac), son. i Agee
V1I.1964) (Ip). PR Luis Potosf{: J. Rzedowski i 6389 (Ip), 10243
(Ip), 24493 (Ip), 24676 (Ac), 2u8h2 (Ac), 21865 (Ip); S. Sander-
son 262 2 (Au--29763h) . Sinaloa: Breedlove 15558 (Rf); Breedlove
& & Kawahara 16890 (Rf). Sonora: Tucker 3478 (Se (Se--6079); R. hi.
Turner 59-50 0 (Sd--65667). Tamaulipas: Stanford, Lauber, & Taylor
2486 (Se--1),9113) ; M. B. Webster 155 (Au--2)1557). Veracruz: Lot

29 (G). GERMANY: Gizantoro s.n. [9.66] (Lu).

XVERBENA ENGELMANNII Moldenke

Additional & emended synonymy: Verbena hastata y oblongifolia
Nutt. ex A. Wood, Class-book, ed. 2 pel, lileue 1847. Verbena
hastato-urticifolia Bebb, in herb.

Additional & emended ppriaecsnar A. Wood, Class—book, ed. 2,
eetenit2 (107), 6d. 2, pr. 2, 112 (188), ed. 10, pr. 1; ti2
(18485, ed. 10, pr. 2, li2 (1849), ed. 10, pr. 3, 412 (1850), ed
17, 412 (1851), ed, 23, 12 (1851), ed. 29, 12 (1853), ed. 35,
Bies(405))) edsdil, pr. 1, hi2 (1855), and ed. I; pr. 2, 12.
1856; Poindexter, Trans. Kans. Acad. Sci. 65: 09, 10, 16, &
Th Ty *1962; Radford, Ahles, & Bell, Guide Vasc. Fl. Carol. 282.
196); Moldenke, Phytologia 16: 2° & 90. 1968; Moldenke, Résumé
Suppl. 16: ficial 1968; F. C. Seymour, Fl. New Engl. 156. 1969;
Domville & Dunbar, John Burroughs Nat. Hist. Soc. Bull. 8: of.
1970; Moldenke, rifth Summ, 1: 15--23, 34, 36--O, 43, 45, 53, &
370 (1971) and’ 2: 649, 667, 673--675, "688, 697, 70h, 705, 783, &
91). 1971.

Wood's (1847) description of Nuttall's V. hastata y oblongi-
folia is "Lvs. lance-ovate or lance-oblong, sharply serrate;
spikes filiform, loosely paniculate; fls. smaller. -- Penn. to
Ia. and Mo. I have frequently observed this tall (l--6 f.)
variety, and many others, on the sandy plains of Indiana. They
appear to be hybrids between V. hastata and V. urticaefolial"

He regards V. paniculata Lam. as a synonyn, but most workers
regard Lamarck's name as applying to the non-hastate leaved
form of V. hastata L.

Morley found the hybrid "scattered in creek bottoms" in Kansas,
while Wharton found it "associated with V. hastata in waste
ground" in Kentucky. Domville & Dunbar (1970) report it "rare"
in Ulster County, New York, where it blooms in the "summer" and
is simply called "vervain". Radford, Ahles, & Bell (196) de-
scribe it as a "rare hybrid" between V. scabra Vahl and V. urti-
cifolia L., but in this they are in error, since it is actually
a hybrid between V. hastata L. and V. urticifolia L. and occurs
commonly in areas not inhabited by V. scabra. They record it
from Ashe County, North Carolina. Seymour (1969) found it at
Leicester (Addison County) and Pownal (Bennington County), Ver-
mont. The corollas on Wharton 5633a are described as having been
"blue", “se Eo

230 P eee OG ak Vol. 23, no. 2

The H. R. Bennett 2707, distributed as this hybrid, seems to
be typical V. hastata L. Three color photographs of this hybrid
in situ (A. R. Moldenke 1261 & 1278) are preserved in my personal

herbarium.

Additional citations: NEW JERSEY: Somerset Co.: Moldenke & Mol-
denke 25635 (Ps--67). ILLINOIS: Cass Co.: Geyer sen. [Beards-
town, July 182] (Ws--isotype). Tazewell Co.: V. H. H. Chase 3230
(Se-—201)860) « Winnebago Co.: M. S. Bebb s.n. (Fountaindale, , 1871]
(Pa). KENTUCKY: Madison Co.: Wharton 5633a (Mi). KANSAS: Republic
Co.: Morley 820 (N). MISSOURI: St. Louis: Engelmann s.n. [St.
Louis, Aug. “Aug. 182] (Ws), sen. [Prairies, 12 August 1877] (Pa).

XVERBENA ENGELMANNII f. ALBIFLORA Moldenke

Bibliography: Moldenke, Résumé Suppl. 3: 2. 1962; Moldenke,
Phytologia 9: 97. 1963; Moldenke, Fifth Summ. 1: 18 (1971) and 2:
91h ° 1971 e

VERBENA EPHEDROIDES Cham,

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 571.
1965; Moldenke, Phytologia 16: 52. 1968; Moldenke, Fifth Summ. 1:
177, 189, & 200 (1971) and 2: 678, 705, 706, & 91h. 1971.

XVERBENA FABRICATA Moldenke
Additional bibliography: Moldenke, Phytologia 11: 57. 1965;
Moldenke, Fifth Sum, 1: 370 (1971) and 2: 67h, 705, & 914. 1971.

VERBENA FASCICULATA Benth.

Additional bibliography: J. F. Macbr., Field Ifus. Publ. Bot.
13 (52 O63, 61k, 620, 625, .&.629. aaeO Moldenke, Phytologia 1:
281 & 282. 1967; *Moldenke, "Fifth Summ. 1: 143 (1971) and 2: 682,
686, & 914. 1971.

According to Macbride (1960), the type of V. fasciculata was
collected by Richard Brinsley Hinds at "Huamantango", which he
says is "probably Huamantamba near Canta, Lima, which is in the
scope of Cuming's brief visit". He cites only Cuming 95) "fide
Bentham" from "On the coast" of Peru. He attempts to keep V.
fasciculata Benth. and V. matthewsii Briq. apart as separate
species, keying them as follows:

Leaves "3-divided nearly to the base, rarely in part entire or
DISURCALSY ae Pedra Gils Celioits, weet Katte Sey 6 -V. fasciculata
Leaves "more or less trifid but obviously cuneate-based or petio-
lately narrowed, the segments variously serrate and pinnate
or dissected, rather entire" ........ . V. matthewsii
As reported by me previously, Edgar Milne-Redhead has compared
the types of both species at Kew and reports that they cannot be
distinguished from each other,

In his discussion of what he calls V. matthewsii Macbride says
"Neighbor of V. Berterii (Meisn.) Schauer and V. clavata R. & P.
with procumbent branches, the former too with shorter bracts, the

1972 Moldenke, Notes on Verbena 231

latter with 3-divided leaves, the segments pseudoverticillate
(author). — Type was distributed as V. laciniata (L.) Briq. and
doubtfully a distinct species but as here listed leaf-segments
broader, in type seen acute but Lima and Hudnuco specimens with
more obtuse segments could be placed here. Species name here
spelled with two t's, as it was sometimes spelled by the collec-
tor. F.M. Neg. 24691." He cites the following Peruvian speci-
mens: Ancash: Macbride 2539. Ayacucho: Ferreyra 5470. Cajamarca:
Stork & Horton 10032. Junin: Soukup 3537. Lima: “Mathews 498,
Stork & Vargas 931.

Verbena fasciculata has been collected at altitudes of 00 to
3900 meters. Recent collectors have found it flowering in Octo-
ber and November and fruiting in October.

Additional citations: PERU: Ancash: Vargas Calderén 10276
(Ac). Arequipa: Vargas Calderén 1826 (Ac).

XVERBENA FECUNDA Moldenke

Additional bibliography: Hocking, Excerpt. Bot. A.1l: 30.
1959; Moldenke, Phytologia 1): 282. 1967; Moldenke, Fifth Sum,
1: 370 (1971) and 2: 658, 782, & 91h. 1971.

XVERBENA FERAX Moldenke

Additional bibliography: Hocking, Excerpt. Bot. A.l: 30.
1959; Moldenke, Phytologia 14: 283. 1967; Moldenke, Fifth Sum.
1: 370 (1971) and 2: 658, 692, & 914. 1971.

VERBENA FERREYRAE Moldenke
Additional bibliography: J. F. Macbr., Field Mus. Publ. Bot.

13 (5): 613, 620-621, & 624. 1960; Moldenke, Phytologia 11: 457—

458. 1965; Mloldenke, Fifth Summ. 1: 143 (1971) and 2: 91h. 1971.

VERBENA FILICAULIS Schau.

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 571.
1965; Moldenke, Phytologia 16: 52-53. 1968; Moldenke, Fifth
Summ, 1: 177 & 200 (1971) and 2: 690 & 91). 1971.

Hatschbach describes this plant as decumbent, growing in acid
turf-covered soil. It has been collected in anthesis in April
and October, in addition to the months previously reported. The
corollas are described as having been "violet" on Hatschbach
14192 & 20082. t eee

Additional citations: BRAZIL: Parand: Hatschbach 1192 (W—
2564173), 20082 (N), 22719 (N). Santa Catarina: Smith & Klein
13390 (N). 2

VERBENA FLAVA Gill. & Hook. in Hook., Bot. liisc. 1: 170. 1829.
Additional synonyny: Verbena flava var, flava Troncoso in
Bocher, Hjerting, & Rahn, Dansk Dansk. Bot. Arkiv 22 (1): 109. 1963;

Additional & emended bibliography: Reiche & Phil., Fl. Chil.
5: 295 & 463. 1910; Hauman, Anal. Soc. Cien. Argent. 86: 150.
1918; J. A. Clark, Card Ind. Gen. Sp. Var. issue 183. 19);

232 Pee TO eit OG ore Vol. 23, 10sec

Troncoso in Boécher, Hjerting, & Rahn, Dansk Bot. Arkiv 22 (1):
109. 1963; foldenke, Phytologia 16: See 1968; Moldenke, Résumé
Suppl. 16: 6. 1968; *Bolkh., Grif, Matvej., & Zakhar., Chrom. Numb.
Flow. Pl. 715 & 717. 1969; Schnack & Rubens, Bol. Soc. Argent.
Bot. 13: 206. 1970; Moldenke, Fifth Summ. 1: 200 (1971) and 2:
521, 668, & 91h. 1971.

Bucher, Hjerting, & Rahn (1963) collected this plant at 2150
meters altitude (no. 2138).

Additional citations: ARGENTINA: Neuquen: E. Ammann 113 (N).

VERBENA FLAVA var. ANGUSTILOBA Speg.
Additional bibliography: Moldenke, Phytologia 9: 122--123.
1963; Moldenke, Fifth Summ. 1: 200 (1971) and 2: 668 & 91h. 1971.

VERBENA FLAVA var. LATILOBA Speg.

Additional bibliography: Hauman, Anal. Soc. Cienc. Argent. 86:
150. 1918; Troncoso in Bocher, Hjerting, & Rahn, Dansk Bot. Arkiv
22): 109. 1963; Moldenke, Phytologia Cero ae "1963; Moldenke,
Fifth Summ. 1: 200 (1971) and 2: 668 & oll ripheakeiy alae

Troncoso (1963) states that this variety was collected by
Hicken at 2000 meters altitude in the Diamente area.

VERBENA FLAVA var. LATILOBA f. ABORTIVA Hicken
Additional bibliography: Moldenke, Phytologia 9: 123. 1963;
Moldenke, Fifth Summ. 1: 200 (1971) and 2: 91). 1971.

VERBENA GALAPAGOSENSIS Moldenke

Additional bibliography: Moldenke, Phytologia 11: 58 (1965)
and 16: 32. 1968; Wiggins & Porter, "F1. Galap.. Isls ..997/<. 17a.
Moldenke in Wiggins & penne Fil. Galdp. Isls. 503--505 & 509.
1971; Moldenke, Fifth Summ, 1: 138 (1971) and 2: 91). 1971.

VERBENA GENTRYI Moldenke

Additional bibliography: Moldenke, Phytologia 11: 58. 1965;
Moldenke, Fifth Summ, 1: 75 (1971) and 2: 91h. 1971.

Breedlove & Thorne found this plant growing on dry slopes with
Bursera, Ipomoea, lysiloma, and Quercus, at 3800 feet altitude,
flowering in September. The corollas are described as having been
white on “Breedlove & Thorne 18073.

Additional citations: MEXICO: Sinaloa: Breedlove & Thorne 18073
Cae <<

VERBENA GLABRATA H.B.K.

Emended synonymy: Verbena glabrata Humb. & Bonpl. ex Steud.,
Nom. Bot. Phan., ed. 1, 873. 1821.

Additional & emenied bibliography: Steud., Nom. Bot. Phan., ed.
1, 873. 1621; J. F. Macbr., Fielid Mus. Publ Bot. 137 (5)s 616" &
621. 1960; Hocking, Excerpt. Bot. A.7: 56. 1964; Moldenke, Biol.
Abstr. 9: 1325 (1968) and 9 (3): B.A.S.I.C. $.185. 1968; *Niolden-
ke, Phytologia 16: 53, 98, 211, & 31. 1968; Moldenke, Résumé
Suppl. 16: 5. 1968; Hocking, Excerpt. Bot. A.l): 206. 1969; Molden-

1972 Moldenke, Notes on Verbena 233

ke, Biol. Abstr. 50: 18. 1969; Wiggins & Porter, Fl. Gal4p. Isls.
26 & 997. 1971; Moldenke in Wiggins & Porter, Fl. Gald4p. Isls.
503--505 & 609. 1971; Moldenke, Fifth Summ, 1: 120, 137, & 143
(1e7l) and 2: 671 & 91). 1971.

Macbride (1960) credits this binomial to "HBK. Nov. Gen. & Sp.
2: 276. 1818", but it actually dates back to "ed. folio, 2: 223-—
22). 1817". He says of the plant "To 1 meter tall, corolla tube
red, blue-violet above, fruit red-maroon (Metcalf). Schauer says
this is more compact, more glabrate than V. litoralis HBK. var.
pycnostachys Schauer; compare V. bonariensis L., V. brasiliensis
Vell." He cites only Metcalf 3029 from Ayacucho, but records
the species also from Cuzco and Lima, Peru, and gives its extra-
limital distribution as "Venezuela; Ecuador." I have never seen
any material of it from Venezuela and cannot believe that it is
found there.

Recent collectors have encountered this plant growing among
lakeshore vegetation, on old airfields, and in secondary cloud-
forests. Holm-Nielsen & Jeppesen found it in "both humid and
more dry places near road in meadows". The corollas on Asplund
17227 are described as having been "pale-lilac", while those on
Asplund 19923 were "pale bluish-violet".

The Rose, Pachano, & Rose 22851, previously cited by me as V.
glabrata, is actually V. demissa Moldenke, while Rose, Pachano,

& Rose 22939 is V. parvula Hayek.

Additional citations: ECUADOR: Cotopaxi: Holm-Nielsen & Jeppe-
sen 117 (Ac). Imbabura: Asplund 20189 (N). Loja: Sparre 1666)
(S). Napo: Sparre 17690 (S). Pastaza: Sparre 17525 (S). Pi-
chincha: Asplund 16030 (N), 16139 (N), 17227 (N), 20151 (N);
Sparre 16899 (S). Tungurahua: Asplund 19923 (N). PERU: La Lib-
ertad: Sagdstegui A. 7192 (N).

VERBENA GLABRATA var. TENUISPICATA Moldenke

Additional bibliography: Moldenke, Phytologia 16: 53, 211, &
341. 1968; Moldenke, Résumé Suppl. 16: 5. 1968; Moldenke, Biol.
Abstr. 9: 1325 (1968), 49 (3): BA.S.I.C. S. 185 (1968), and
50: 418. 1969; Hocking, Excerpt. Bot. A.ll: 206. 1969; Wiggins &
Porter, Fl. Galdp. Isls. 26 & 997. 1971; Moldenke in Wiggins &
Porter, Fl. Galfp. Isls. 503--505 & 509. 1971; Moldenke, Fifth
Sum. 1: 138 (1971) and 2: 91). 1971.
( oe citations: GALAPAGOS ISLANDS: James: C. Darwin s.n.
Cu). 7

VERBENA GLANDULIFERA Moldenke

Additional bibliography: Moldenke, Phytologia 13: 196. 1966;
Moldenke, Fifth Summ. 1: 200 (1971) and 2: 521 & 914. 1971.

In addition to months previously reported by me, this plant
has also been found in fruit in January. The corollas on Varela
699 are described as having been "sky-blue",

Additional citations: ARGENTINA: Catamarca: Luna Risso 505 (N).

234 P HO OG ele a: Vol. 23, Ge 2

San Luis: Varela 699 (N). Santiago del Estero: Pierotti "h" (31/
III/ub)] (N), "h" (3/IV/h4] (N).

VERBENA GLUTINOSA Kuntze

Additional bibliography: Briq. in Chod. & Wilczek, Bull. Herb.
Boiss., sér. 2, 2: 5h. 1902; Troncoso in Bocher, Hjerting, &
Rahn, Dansk Bot. Arkiv 22 ite 109. 1963; Moldenke, Phytologia 1}:
283 & 287. 1967; Schnack & Rubens, Bol. Soc. Argent. Bot. 13: 207.
1970; Moldenke, Fifth Summ. 1: 18h & 200 (1971) and 2: 521, 668, &
Ue tcy ale

Briquet (1902) cites Wilczek 54. Schnack & Rubens (1970) re-
port the species from La Pampa and Neuquén, Argentina, while
Troncoso (1963) cites Bocher, Hjerting, & Rahn 676 & 768, but
notes that the material of the former number is "not typical".
These collectors found the plant growing in riverbeds and describe
the flowers as "yellow or flesh-coloured",

Additional citations: ARGENTINA: Mendoza: H. H. Bartlett 19521

(N).

xVERBENA GONZALEZI Moldenke

Additional bibliography: Moldenke, Phytologia 13: 196. 1966;
Moldenke, Fifth Summ. 1: 370 (1971) and 2: 522, 689, 69h, & 91k.
1971.

VERBENA GOODDINGII Bria. lat 4
Additional & emended synonymy: Glandularia gooddingii (Briq.)
Solbrig ex Rattenbury, Madrofio 15: 50. 1959. Verbena bracteata
gooddingii Ferris in Abrams & Ferris, Illustr. Fl. Pacific States,
: in syn. 1960.
Pr» dasth ooy’ a omonied ibliography: Rydb., Fl. Rocky Mtns., ed.
1, 739 & 740 (1917) and ed. 2, pr. 1, 739 & 70. 1922; Tidestr.,
Contrib. U. S. Nat. Herb. 25 [Fl. Utah & Nev.], pr. 1, 69. 1925;
E. C. Jaeger, Desert Wild Fls., ed. 1, 219 & 220, fig. 508 (1905
and ed.[2], pr. 1, 219 & 220, fig. 508. 1941; Abrams, Illustr.
Fl. Pacif, States, pr. 1, 3: 610, 612, & 616, fig. 43h7. 1951;
Rydb., Fl. Rocky Mtns., ed. 2, pr. 2, 739 & 70. 195k; J. A.
Clark, Card Ind. Gen. Sp. Var. issue 231. 1959; Rattenbury, Mad-
rofio 15: 50. 1959; Ferris in Abrams & Ferris, Illustr. Fl. Pacif.
States, pr. 1, h: 651 & 730. 1960; Howell & McClintock in Kearney
& Peebles, Ariz. Pl., ed. 2, 725-727. 1960; Lewis & Oliv., Am.
Journ. Bot. 8: [639]. 1961; Munz, Calif. Desert Wildfls., ed. 1,
7 & 122, pl. 62. 1962; Hocking, Excerpt. Bot. A.6: 91 {1963)
and A.7: 456. 1964; Ferris in Abrams & Ferris, Illustr. Fl. Pac-
if. States, pr. 2, h: 651 & 730. 1965; G. Taylor, Ind. Kew. Suppl.
13: 61. 1966; Abrams, Illustr. Fl. Pacif. States, pr. 2, 3: 610,
612, & 616, fig. 4347. 1967; L. C. Higgins, Fl. Beaver Dam Mtns.
223. 1967; E. C. Jaeger, Desert Wild Fls., ed. 2, pr. 9, xxvii,
219, & 220, fig. 508. 1968; Moldenke, Phytologia 16: 187, 202, &
215. 1968; Munz & Keck, Calif. Fl. 687, 688, & 1679, fig. 65.
1968; Munz, Suppl. Calif. Fl. 101. 1968; Bolkh., Grif, Matvej., &
Zakhar., Chrom. Numb. Flow, Pl. 715 & 717. 1969; Munz, Calif.

1972 Moldenke, Notes on Verbena 235

Desert Wildfls., ed. 2, 47 & 122, pl. 62. 1969; Rickett, Wild Fls.
U. S. 3 (2): 362 & [363], pl. 110. 1969; Rydb., Fl. Rocky Mtns.,

6d. 2, pr. 3, 739 &:7h0. "1969; Tidestr., Contrib. U. S. Nat. Herb.
25 (Fl. Utah & Nev.], pr. 2, he. 1969; Correll & Johnston, Man.
Vasc. Pl. Tex. (Contrib. Tex. Res. Found. Bot. 6:] 1876 & 1877.
1970; Moldenke in Correll & Johnston, Man. Vasc. Pl. Tex. [Con
trib. Tex. Res. Found. Bot. 6:] 1317 & 1325—1326. 1970; Rickett,
Wild Fls. U. S. (3): 539, (541), & 799, pl. 176. 1970; Moldenke,
Fifth Summ. 1: 32, 50, 51, 53, 58, 62, 63, 65, 75, & 370 (1971)
and 2: 521, 654, 656, 671, 706, 708, & 915. 1971; Moldenke, Phyto-
logia 22: 485 & 497 (1972) and’ 23: 188. 1972.

Additional & emended illustrations: E. C. vaneehy Desert Wild
Fls., ed. 1, 220, fig. 508 (190) and ed. [2], pr. 1, 220, fig.
508. 191; Nunz, "calif. Desert Wildfls., ed. re 7, pl. 62 (in
color). 1962; Munz & Keck, Calif. Fl. 687, fig. 65. 1968; E. C.
Jaeger, Bawert Wild Fls., " athe ce (Din Os 230, fig. 508. 1968; Munz,
Calif. Desert Wildfls., ed. 2, 47, pl. 62 (in color). 1969; Rick-
ett, Wild Fls. U. S. 3 (2): [363], pl. 110 (in color) (1969) and
i (3): [541], pl. 176 (in color). 1970.

The Madrofio (1959) reference in the bibliography above is some-
times credited to Solbrig, while Bolkhovskikh anc his associates
(1969) cite it as "Darlington, Document. Chrom, Numb. Pl....",
but it seems actually to be a continuation of a series started by
Rattenbury and not otherwise accredited.

Recent collectors describe this plant as having dark-green
foliage and have found it growing in dry mesquite deserts and
roadsides. Moran reports it as "rather scarce" at 1650 meters
altitude in Baja California, while Gentry & Fox found it "scatter-
ed along moister bottomland of open canyon bottoms" in the same
state. The Mearses describe it as "common with Astragalus, Cen-
chrus, and Gaillardia" in Santa Cruz County, Arizona, "common with
Cowania, Eriodictyon, Mirabilis, and Penstemon" in Clark County,
Nevada, and as growing with Juniperus and Yucca baccata in San
Bernardino County, California. Higgins (1967) confirms its pres-
ence in Washington County, Utah. Jaeger (1968) informs us that it
is known in California from dry limestone canyons and slopes of
the Clark and Providence Mountains and from there eastward to Tex-
as. Howell & McClintock (1960) aver that it is found "throughout
the state [of Arizona] mostly below 5,000 feet, dry slopes &
mesas, flowering throughout the year; commonest and most wide-
spread species in Arizona". They cite Jones 3901, from Yucca, Mo-
have County, as the type collection of V. ar arizonica Briq., which
is correct, but it represents var. nepetifolia Tidestr. rather
than the typical form of the species. They cite also N. ©. Wilson
95, from Diamond Creek, Mohave County, as the type collection of
Ve verna A, Nels.; this is also now regarded as var. nepetifolia.

The f following vernacular names are recorded for V. gooddingii
in addition to those previously reported by me: "desert vervain"
and "Goodding verbena",

Lewis & Oliver (1961) give 30 as the diploid chromosome number

236 1st AG ye (0) at, (0) (eh ac VN Vol. 23, no. 2

for this species, while Solbrig is said by Rattenbury (1959) to
have found it to be 20. It should be noted, however, that the
Lewis & Oliver record was for var. nepetifolia rather than the
typical form of the species.

The corollas on Gentry & Fox 11789 are described as having
been "pink", those on Beatley 9969 were "deep blue-lavender", and
those on R. V. Moran 10993 were e@ "pale-lavender". Rickett (1969)
describes the | species as "densely hairy. The leaves are cleft
into three lobes which are coarsely toothed or jaggedly cleft. The
corolla may be pink, lavender, or blue, the lobes spreading 1/3 —
1/2 inch across", says that it blossoms from "February to October"
and is found Nin’ sandy soil and mountains from southwestern Texas
to California and Nevada; and in Baja California".

Material of this species has been misidentified and distribu-
ted in some herbaria as V. neomexicana (A. Gray) Small. On the
other hand, the 0. M. Clark 1128, distributed as V. gooddingii,
is aah var. - nepetifolia Tidestr., Wiggins & Thomas 187 is
V. lilacina Greene, and H. V. Moran 11476 is V. “plicata Greene.

Additional citations: NEVADA: Clark Co.: Mears & Mears 1895
(Au--258]63). Nye Co.: Beatley 9969 (N). TEXAS: Brewster Co.:
Nelson & Nelson 5025 (Se—11876). ARIZONA: Coconino Co.: Coffey
25 (Ac); DeLuna 19 19 (Rf). Mohave Co.: N. D. Atwood 1730 (N, W--
259916) ; X. Me. Gaines 1361 (Se--228)22); C C. L. Hitchcock 2561)
(N, Se--235128, W--2580550) ; Munz 11691 (Se--126728) « Pima Coe:
C. L. Hitchcock 2550 (N, Se--236),80, W--2580529). Pinal Co.: A.
Davis 32 (Fg). Santa Cruz Co.: Keil, Pinkava, & Lehto 933) (N);_
Mears & Mears 1691 (Au--257979) . Tavaneseos oe Blalock 21 21 (Rf);
Demaree 13883 (Rf); Greenwell 63 (Ac); C. F. Harbison 11178 (Sd—
28992); D - F. Howe s.n. [2h April 1966] (Sd--64277). CALIFORNIA:
San ene ere Co.: Me “M. Beal 51h (Se—161257); Letcher s.n. [San
Bernardino, March 31, 1929] (Sd—-l1006); Moldenke & Moldenke 3206
(Ac); Munz "12860 (Se--16277) ; Munz, Johnson, & Harwood h25h
(Se--162773); C. B. Wolf 7021 (Se—-137h67), 9680 (Sd—2]96, Se—
122997), 10718 (Au--122112). MEXICO: Baja California: Gentry &
Fox 11789 (i); j R. V. Moran 10993 (Sd--53867), 13532 (Sd—-636)6) ;
ee Wiggins 1507) (Se--208881); Wiggins & Wiggins eins 16073 (Au=--
238501). CULTIVATED: California: Balls 18900 (Se—157360) .

VERBENA GOODDINGII var. NEPETIFOLIA Tidestr,

Additional synonymy: Verbena gooddingii nepetifoloia Tidestr.,
in herb,

Additional & emended bibliography: Tidestr., Contrib. U.S.
Nat. Herb. 25 [Fl. Utah & Nev.], pr. 1, 69. 1925; Lewis &
Oliv., Am. Journ, Bot. 8: [639]. 1961; Hocking, Excerpt. Bot.
A.6: 91. 1963; Moldenke, Phytologia 16: 187. 1968; Munz, Suppl.
Calif. Fl. 101. 1968; Tidestr., Contrib. U. S. Nat. Herb. 25
[Fl. Utah & Nev.], pr. 2, 69. 1969; Moldenke, Fifth Sum, 1:
BOSSI, 163506558 715 (1971) and 2: 652, 65h, 663, 671: 106,.ies

1972 Moldenke, Notes on Verbena 237

& 915. 1971; Moldenke, Phytologia 22: 85 & 497 (1972) and 23:
188. 1972.

Recent collectors have found this variety growing in arroyos,
among dry rocks, and among eroded granite boulders. Moran found
it to be "occasional along arroyos" in Baja California, while Mo-
ran, Witham, & Hommersand report it "locally common in disturbed
areas" and Henrickson says it was found "in edge of deeply cut
erosion areas in clay soil" in the same state. Mears calls it
"common in canyons with Agrostis, Erythrina, and Rhus" in Arizo—
na. The corollas are said to have been "rose" on R. V. Moran
7893 and "rose-puple" on Henrickson 2311. men

~~ Material has been misidentified and distributed in some herbar-
ia under the names V. bracteata Lag. & Rodr. and "V. cilliata
Benth." On the other hand, the Diaz L. s.n. [16 .1X.19 59], dis-
tributed as this variety, is actually ve ~delticola Small, while
Jarabek sen. [June 195] is not even verbenaceous.

Additional citations: ARIZONA: Mohave Co.: Pinkava, Lewis,
Noble, & Lehto 1129 (N). Pima Co.: L. M. Andrews 259a (N)5 (N)5 J.
A. Mears 1744 (Au--258005). Yavapai Co.: Toumey 305 (Si (Sd--52238).
County undetermined: O. M. Clark 1128 [S. Bateswell, Grawley
Mtns.] (Au--122113). "MEXICO: Baja a California: Bell be Newcomb
1356 (Se--1)8592); Bratstrom s.n. [March 17, 1951] 51) (Sd); C.F.
Harbison 27283 (Sd—-27283), 4isho (Sd—-15)9), sen. [April 2,
1950] (Sd--l,3850) ; Hastings & & Turner 63-192 (Sd——56502) ; Hen-
rickson 231] (Mi); Huey s.n. “(June 16, 1947] (Sd—-hl9hh); R. Ra Ve
Moran 7893 ( (Sd--60711) , 10226 (Sd—SL55h) ; Moran, Witham, & Hom-
mersand 1651 (Mi). Sonora: I. L. Wiggins 7225 (Se—168338),

11775 (Sd=-}}7325) .

XVERBENA GOODMANI Moldenke

Synonymy: Verbena halei Small x V. stricta Vent. ex Moldenke,
Fifth Summ. 2: 672, in syn. 1971. Verbena stricta Vent. x Vv.
halei Small ex Moldenke, Fifth Sum, 2: . 2: 698, in syn. 1971.
Verbena goodmanii Woldexks, in herb.

Additional bibliography: Moldenke, Phytologia 1): 283. 1967;
Moldenke, Fifth Summ. 1: 53 (1971) and 2: 672, 698, & 915. 1971.
Additional citations: OKLAHOMA: Marshall Coe: Goodman tes3

(Se--21251h).

VERBENA GOYAZENSIS Moldenke
Additional bibliography: Moldenke, Phytologia 9: 1)7--1)8.
1963; Moldenke, Fifth Summ. 1: 177 (1971) and 2: 915. 1971.

VERBENA GRACILESCENS (Cham.) Herter

Additional & emended bibliography: Cabrera, Man. Fl. Alred.
Buenos Aires 39) & 395. 1953; Troncoso in Cabrera, Fl. Prov.
Buenos Aires 5: 128 & 131. 1965; Meyer & Weyrauch, Inst. Mig.
Lill. Misc. 23: [Guia Excurs. Biol. Tucum.] 6) & 123. 1966; Mar-

238 Pi Ry P 0) LOG A Vol. 23, no. 2

tinez—Crovetto, Bonplandia 2: 131. 1967; Moldenke, Phytologia 16:
55 & 102. 1968; Bolkh., Grif, Matvej., & Zakhar., Chrom. Numb,
Flow. Pl. 717. 1969; Moldenke, Fifth Summ. 1: 177, 18h, 187, 189,
©: 200) (1972) candi.2s 651, 1671, 672,687, 9155 197s

The corollas of this species are described as having been
"rose" on A. T. Hunziker 6318, "lilac" on Burkart 138)1 and Luna
370 & 451, and "blue" on Ibarrola 2169. The plant has been col-
lected in fruit from July to December. Troncoso (1965) gives its
distribution as "Sudamérica templada. Comin en campos bajos y
himedos" and cites Burkart 3656, Cabrera 5701 & 7223, and Venturi
59 from Buenos Aires, deposited in the San Isidro and La Plata
herbaria.

Additional citations: ARGENTINA: Cérdoba: Balegno 256 (N); Bur-
kart 1381 (N); A. T. Hunziker 6318 (N). Corrientes: Ibarrola 2169
(N). Formosa: I. Morel 16 (N), 3459 (N), 3648 (N), 3667 (N), 3723
(N), 3778 (N). Jujuy: A. Reales 633 (N), 969 (N). Salta: Luna
451 (N); T. Meyer 8403 (N). Santa Fé: Rufz Huidobro 3080 (Se--
129885). Santiago del Estero: Luna 370 (N), 415 (N). Tucum4n:
O'Donell 5417 (N). PARAGUAY: T. Rojas 102b (Ws). URUGUAY: Ro-
sengurtt Gurvich & Gallinal 5803 (Se-—-122506).

ae

VERBENA GRACILIS Desf.

Additional & emended bibliography: Rydb., Fl. Rocky Mtns., ed.
2, Pr. 1, 739 & THO. 1922; Tidestr., Contrib. U.S. Nat. Herbesce
[Fl. Utah & Nev.], pr. 1, 469. 1925; Rydb., Fl. Rocky Mtns., ed.
2, pre 2, 739 & 7hO. 195); Howell & McClintock in Kearney & Peeb-
les, Ariz. Fl., ed. 2, 726 & 728. 1960; Moldenke, Phytologia 16:
55. 1968; Moldenke, Résumé Suppl. 16: 2. 1968; Rydb., Fl. Rocky
Mtns., ed. 2, pr. 3, 739 & 70. 1969; Tidestr., Contrib. U. S.
Nat. Herb. 25 [Fl. Utah & Nev.], pr. 2, 169. 1969; Rickett, Wild
Fls. U. S. 4 (3): S40 & 799. 19703; Moldenke, Fifth Summ. 1: 50,
63, 75, & 370 (1971) and 2: 652, 688, 693, & 915. 1971; Moldenke,
Phytologia 22: 85 & 99. 1972.

The type of V. arizonica A. Gray was collected by John Gill
Lemmon in Tanner's Cafion, near Fort Huachuca, Cochise County, Ar-
izona, in 1882, and is deposited in the Gray Herbarium at Harvard
University. Howell & McClintock (1960) state that the species
grows at altitudes of ],500--6500 feet in Arizona and flowers
there from Juhe to October. Recent collectors have found it in
flower and fruit also in December, growing in abandoned fields
and on inundated ground. The Matthews describe it as "very abun-
dant throughout open fields along railroad tracks" in Sonora,
Urueta found it in "pastizales" in México, Stuessy calls it
"scarce" in Chihuahua, Holgufn found it on "ladera basdltica con
vegetacién de matorral de Opuntia, Zaluzania y Mimosa" in México,
and Rzedowski in "charcos cercanos a la carretera" in México and
"Jadera riolftica con vegetacién de zacatal" in San Luis Potosi.

Material has been misidentified and distributed in some her—
baria as V. halei Small. The Gonzd4lez Quintero 1908 and J. Rze-

1972 Moldenke, Notes on Verbena 239

dowski 15671 represent a small-bracted densely fruited form of

the species. The corollas on Stuessy 1013 are described as having
been "blue" and on J. Rzedowski 1352, 1473, 11225, & 11265 as
"purple". The P . 0, Schallert Sone [6/16/43], di distributed as Vv.
gracilis, is aetualiy Vv. bracteata | Lag. & Rodr.

Additional citations: ARIZONA: Cochise Co.: W. W. Jones s.n.
[Aug. 10, 1926] (Sd--7326). MEXICO: Chihuahua: Stuessy 1013
(Ip). Federal District: J. Rzedowski 1352 (Au--2)1293, Ip—1615),
173 (Ip--161)). Hidalgo: Gonz4lez Quintero 1908 (Ip). México:
Holguin s.n. [)/VII/1965] (Ac); J. Rzedowski 15671 (Ip), 20288
(Rf); U Urueta 32 (Rf). San Luis Potosi: J. Rzedowski 31:29 (Ip),
1995. (Tp), 11225 (Ip), 11265 (Ip). Sonora: Matthews & Matthews
487 (Au--25993]) «

VERBENA GRISEA Robinson & Greenm,

Additional bibliography: Moldenke, Phytologia 16: 55--56 & 32.
1968; Moldenke in Wiggins & Porter, Fl. Galfp. Isls. 503 & 505.
1971; Wiggins & Porter, Fl. Galdp. Isls. 997. 1971; Moldenke,
Fifth Summ, 1: 138 (1971) and 2: 915. 1971.

VERBENA GUARANITICA (Troncoso) Moldenke, Phytologia 23: 211. 1972.

Synonymy: Glandularia guaranitica Troncoso, Darwiniana 16:
618--621, fig. 3. 1971.

Bibliography: Troncoso, Darwiniana 16: [613], 614, & 618--621,
fig. 3. 1971; Moldenke, Phytologia 23: 211. 1972.

Illustrations: Troncoso, Darwiniana 16: [619], fig. 3. 1971.

This species is based on A. Burkart 8028, collected at Tres
Cerros, dept. San Martin, Corrientes, eat ey on November 8,
1936, and is deposited in the herbarium at San Isidro. Mrs. Bur-
kart records for it the vernacular name "margarita" and gives
its geographic distribution as "Misiones y Corrientes en la Ar-
gentina; Paraguay. Habita en rozados y terrenos altos a orillas
de bosques." She makes the following comments: "Agradezco al
Ing. Benno Schnack de la Universidad Nacional de La Plata, los
ejemplares de Misiones y de Corrientes de esta especie y sus
afines, que me proporcioné para su estudio, as{ como varias ob-
servaciones sobre las mismas, las que tiene en cultivo en el
jardin experimental de la Facultad de Agronomfa de La Plata....
G. guaranitica adem4s de su afinidad ya seflalada con G. megapo-
tamica y G. phlogiflora, podrfa confundirse por su hAbito y as-
pecto con G. incisa (Hook.) Tronc., especie que difiere funda-
mentalmente por sus inflorescencias que se alargan en forma
notable después de la antesis, por la ausencia de glAndulas con-
ectivales y por sus flores de color rojo-solferino....Hassler,
en Chod. et Hassl., Pl. Hassler: 77. 190, reconoce Verbena
megapotamica var. tweediana 0. Ktze. Su interpretacién de esta
variedad corresponderfa a Glandularia guaranitica. Me baso para
afirmarle en uno de los ejemplares citados por este autor (Hass-

2),0 P. BiMrt0, Ge OaGe ark Vol. 23, no. 2

ler 3293 de Paraguay, Caraguatay), que he visto en el herbario de

Kew."
She cites the following specimens: PARAGUAY: Fiebrig 652,

Hassler 1466 & 3293, Osten 13571, T. Rojas 1882. ARGENTINA: Cor-
rientes: " Burkart + 8028 (type Selection | Misiones: Grondona &
Spegazzini ipioon,. Issouribehere s.n. [Posadas], A. de Llamas s.n.
[Santa Ana, Candelaria], F. M. Rodriguez 30.

VERBENA GYNOBASIS Wedd.

Additional & emended bibliography: Reiche & Phil., Fl. Chil.
5: 285 & 286. 1910; J. F. Macbr., Field Mus. Publ. Bot. 13 (5):
617 & 618. 1960; Moldenke, Phytologia 1): 28). 1967; Moldenke,
Fifth Summ, 1: v3, 18h, & 193 (1971) and 2: 653 & 915. 1971;
Moldenke, Phytologia 23: 191. 1972.

Recent collectors have found this plant growing at altitudes
of 00 to 300 meters, flowering and fruiting in October and No-
vember, in addition to months previously reported. Macbride
(1960) ° reduces the species to synonymy under V. calcicola Walp.,
but I regard V. calcicola as a synonym of V. clavata Rutz & Pav.
and quite distinct from the present taxon. _

Additional citations: PERU: Arequipa: Vargas Calderén 18227
(Ac). Tacna: Vargas Calderén 13068 (Rf). “CHILE: Tarapacd: Zoll-
ner 1088 (Go).

VERBENA GYNOBASIS var. STRIGOSA Wedd,
Additional bibliography: Moldenke, Phytologia 11: 59. 1965;
Moldenke, Fifth Sum, 1: 1):3 (1971) and 2: 672 & 915. 1971.

VERBENA HALEI Small, Bull. Torrey Bot. Club 25: 617-618. 1898.

Additional synonymy: Verbenna halei Small ex Moldenke, Fifth
Summ, 2: 708, in syn. 1971.

Additional & emended bibliography: J. K. Small, Bull. Torrey
Bot. Club 25: 617--618. 18985 Hocking, Excerpt. Bot. A.1: 30.
1959; Howell & McClintock in Kearney & Peebles, Ariz. Fl., ed. 2,
726 & 728. 1960; Lewis & Oliv., Am. Journ. Bot. 8: [639]--6)1,
figs Tos 196i; Hocking, Excerpt. Bot. A.6: 91. 1963; Radford,
Ahles, & Bell, Guide Vasc. Fl. Carol. 281 & 282. 196; Shinners,
Sida 3: 182. 1967; Moldenke, Phytologia 16: 187 & 190. 1968;
Moldenke, Résumé Suppl. 16: 2. 1968; Pullen, Jones, & Wats.,
Castanea’ 5320332. 1960,bolkh Gris. Matvej., & Zakhar., Chrom.
Numb. Flow. Pl. 717. 1969; Dens Costello, Prairie World 2h.
1969; Muehlenbach, Ann, Mo. Bot. Gard. 56: 169--170. 1969;
Rickett, Wild Fls. U. S. 3 (2): 36h (1969) and (2): 5hO &
(Eee, 1970; Moldenke in Correll & Johnston, Man, Vasc. Pl. Tex.
[Contrib. Tex. Res. Found. Bot. 6:] 131) & 1318--1319. 1970;
Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex. Res.
Found. Bot. 6:] 1876 & 1877. 1970; Shinners, Sida 3: 554. 1970;
Moldenke, Fifth Summ. 1: 23, 25, 27, 30, Bos 33, U5, iveeloe
53, 58, 62, 63, 75, & 370 (1971) and 2: 663, 672, 67h, 679, 680,
685, 686, 698, 708, & 915. 1971; Moldenke, Phytologia 233 193.

1972 Moldenke, Notes on Verbena 21

1972.

Emended illustrations: Lewis & Oliv., Am. Journ. Bot, 8: 6)0,
Pie, 16. 1961.

Recent collectors describe this plant as an annual herb with
fibrous roots or an erect perennial, slender and wiry, )5—60 cm.
tall, branched from the base, the branches erect, to fs cm. long,
the leaves shiny-green, the inflorescence very slender, and the
flowers small. The corollas are described as having been "pale-
bine" on R. Runyon 2358, "Light—blue" on Pilcher 196 & 236,

der-blue" on M. D. McCracken 90, "pluish-lavender™ on n Ellison &
Ellison 1010, “Wlavender" on Be }. Hutchins 453 & 1101 and Jeter | Jeter ll,
"Li ght-lavender on Correll & & Mitchell 3LL52, "violet" on isan
81 and McDaniel 3501, and " and "purple" on Gutiérrez R. 218 and R.

Runyon 2181.

Demaree 58109 is anomalous, tall, the lower leaves numerous on
the stem and only sparsely dentate, hardly lobed, and quite pil-
ose beneath -——- it may represent a hybrid, possibly with V. sin-
plex Lehm. Jones & Reynolds 1170 has some of its spikes _ fasci-
ated. Wood 721 b bears a notation that "plants similar to nos. 692
and 693 but but less hairy". B. Hutchins 53 is accompanied by a
photograph; Nixon G.60 60 is an "ecological ~ voucher", according to
its collector, while Cumbie 173 is a "voucher for. anatomical
study".

Rickett (1969) states that "There are several spikes, with
rose, blue, lavender, or purple flowers scarcely more than 1/3
inch "long": and gives its distribution as "in meadows and wood-
land and on prairies and hills and roadsides practically through-
out the state [of Texas], eastward to Alabama, northward to Ok-
lahoma, and in Mexico", flowering in Texas from "February to No-
vember". The species has been found growing at altitudes of 9--
5800 feet. Additional common names recorded for it are "slender
verbena" and "vervain".

Recent collectors have found the plant growing in loose red or
red sandy loam, loose sand in woods around bogs, calcareous clay-
loam, sandy-clay of river-bottoms, sandy loam or calcareous clay
loam of barditch highways, limestone soil, or in soil derived
from granite, in sandy loam, open pasturelands, open oak-mequite
woodlands, or open fields and disturbed areas, in fine sandy
soil, deciduous tropical woods, low moist areas of rolling plains,
wet spots along highways, and in irrigation ditches, along rail-
road tracks, in Prosopis-Opuntia communities, on road shoulders
and sandy weedy beaches, along the edges of upland mixed hardwoods,
and in hilly areas with serpentine mounds vegetated mainly by
grasses. Pilcher reports it "occasional among weeds and grasses",
while the Ellisons describe it as a "perennial in full sun, sandy
soil, abundant local population with V. brasiliensis but no hy-
bridization evident".

In Arkansas it is called "common in waste areas" by Demaree.

2h2 PRY TO OCG a Vol. 23, no. 2

Stuessy found it "scarce" in Chihuahua, while in Mississippi
Rogers found it "in full sun along roadside and in openings" and
McDaniel says that it is "common along roadsides in longleaf pine
and oak associations". Mears found it growing with Ambrosia,
Cardiospermum, Solanum, and Spartina, while Cory describes it as
"frequent" or "infrequent" on roadsides, "frequent in sandy soil
of grasslands and onion fields", and "infrequent on highway
shoulders". In the state of Texas Strother describes it as a
"weed in overgrown lots" (Brown Co.), Waller found it to be "oc-
casional in sandy clay loam of oak woodlands" (Mason Co.), Jeter
calls it "common on blackland prairies" (Ellis Co.), and Runyon
found it to be "frequent to abundant in open ground" or "occasi-
onal in sandy loam" (Kenedy Co.) and comments that it "flowers
throughout the summer months, frequent throughout the region"
(Cameron Co.).

Stanford reports V. halei from "mostly grassland of sandy to
sandy-loam soil in the Campestrian Biotic Province, frequent in
all areas" of Wichita Co., Texas. Shinners (1967) records it
from Garza County, citing his nos. ))53, 109, & 1101. Kibler en-
countered it at 2000 feet altitude in Nuevo Le6én, Mexico.

Howell & McClintock (1960) cites Peebles & Harrison h22h from
Pinal Co. and Goodding & Lusher 186-5 from Pima Co., Arizona.
Pullen, Jones, & Watson (1968) record it from Amite, Clarke, Cov-
ington, Harrison, Jackson, Jasper, Jefferson Davis, *Lauderdale,
Marion, Scott, Stone, Wayne, and Wilkinson Counties, My ssissippi .
Radford, Ahles, & Bell (196) report it as infrequent along road-
sides and in pastures in Warren County, North Carolina, amin
Aiken and Sumter Counties, South Carolina, flowering there from
April to June.

Muehlenbach (1969) has found this species as an adventive in
two freight yards in St. Louis, Missouri, in 1957, 1958, and 1960,
but in a letter to me dated March 20, 1970, he says "I tried in
the last years desperately to rediscover in St. Louis Verbena
halei, but in vain. I left even once two specimens uncollected,
but nevertheless this plant disappeared in the following summer.
But that is nothing astonishing — so frequently adventive
plants vanish on the railroads without obvious reasons."

The H. M. Parker hah, cited below, was collected in a thorn
forest and d@ dry stream m bed on Tipeetene and shale substratum and
was misidentified as Phryma leptostachya L. It is an anomalous
specimen which does not have the appearance of typical V. halei.

Material of this species has also been misidentified and dis-—-
tributed in some herbaria under the names V, canadensis L., V.

runyoni Moldenke, and Phryma sp. On the other hand, the Sosa
332, Gianna, as V. halei, is actually We ieee Moldenke;
Stuessy 1013 is V. gracilis Desf.; E. G. Marsh 68h, is V. neomexi-
cana var. hirtella Perry; Matthews & & Matthews 210 is Vix perennis
Wooton; Cuesta 61, N. C. Henderson 61-9, and J. J. 0. Perez pan are
V. plicata Greene; and 7 Thomas, Thomas, Th Thomas, & & Thomas as 337 ~ is

1972 Moldenke, Notes on Verbena 23

V. xutha Lehn,

Additional citations: SOUTH CAROLINA: Aiken Co.: Ellison & El-
lison 1010 (N). MISSISSIPPI: Forrest Co.: S. McDaniel 3501 (N).
Hancock Co.: S. B. Jones 11867 (N). Jackson Co.: K. be Rogers
1093 (N). Lawrence Co.: Jones & Reynolds 1170 (Au--260968) «
Pearl River Co.: F. He Sargent 7745 (Go). “ARKANSAS: Clark Co.:
Demaree 58570 (Ac). Little River Co.: Demaree 58109 (Rf), 58166
(Ac). Miller Co.: Demaree 58076 (Ac), 58081 58081 (Ac), 58083 (RE).
Nevada Co.: Demaree 60103 (RE). LOUISIANA: Caddo Par.: Demaree
55579 (Ac). Ouachita Par. R. D. Thomas 2825 (N). OKLAHOMA: A-
toka Co.: Hopkins, Nelson, & @ Nelson 1092 (Se—136591). Johnston
Co.: G. T. Robbins 2h10 ( (Ny Love Co.: 0.: Nelson, Nelson, Goodman,
& Waterfall 5706 (Se——136609). Pontotoc Co.: G. Te. Papkind ins 21,06
(N). TEXAS: Anderson Co.: K. E. Smith s.n. (Palestine, ,/23/35]
(Lk). Bastrop Co.: Westlund, Rowell, | & . Barkley 17f216 (Ip).

Bell Co.: M. Latham 7 (Au—2h7766, & Au-—2),80L) . Bexar Co.: J. O.
Perez 21 (Au—2))5152, Ip). Bosque Co.: Pilcher 236 (Lk). Bowie |
Co.: Correll & Mitchell 3452 (Ld). Brazoria Co.: Fleetwood

9010 (Au—260)20), 9110 (Au--260)28). Brazos Co.: Cory 50627
(Mi); Fryxell 976 (N).- Brown Co.: Calhoon 6 (Lk); M. A. Cole 120
(Lk); R. Norton n 8 (Au--2)8)0)); Strother 10 ) (Lk); J. Wheeler uh
(Au—-2l,7691). Burnet Co.: Cumbie 173 (Lk). Caldwell Co.: Mears
670 (Au—-255128). Cameron Coe: R Re Runyon 2181 (Au--268811) , » 2368
(Au--268612). Culberson Co.: Cory 53037 (Mi, N). Dimmit Co.: _
Atwood 2030 (N, N); Ramirez & Cardenas 3 (Au—2l5212, Ip). Duval
Co.: Co.: Alvarez, Guajardo, Sal Salazar, & McCart 7671 (Lk). Eastland
Co.: Wheeless ay, 17 (Au--2)7705). | Ellis Co.: Jeter 11 (Lk). Frio
Co.: Alvarez, Guajardo, Salazar, & McCart 8020 (Se—220),75) ;

Drews 6 (Au—21,7590); Ramirez & & Cardenas denas 25 (Ai (Au-—-2)5196, Ip). Gal-
veston Co.: Cory 51017 £1017 (Mi) 5 Mears 761 (Au--255271), 793 (Au—
255508). Garza Co.: B. Hutchins 453 (Lk), 109 (Ik), “101 (Lk).
Gillespie Co.: Nixon G.60 G .60 (Au—253187), S37 7 (Au--28323h) » Harris
Co.: E. Hall 432 (N) (N). Hidalgo Co.: Holloway 21 (Au—-257886). Jim
Hogg Co.: Botello & Ayala 12 (Au--2),5005). Kenedy Co.: R. Runyon
2358 (Au--266158, Au——268613). Kleberg Co.: Cory 51322 (Mi).
Lavaca Co.: Strother 185 (Au--23811). Mason Co.: Gipson 81 (Lk);
Waller 1701 (Lk). Matagorda Co.: Cory 51091 (Mi); Mears 19
(Au--25),959). McLennan Co.: L. D. Smith h 388 (Au—-122207). Medi-
na Co.: J. Cope 26 (Au--2):7823) « "Nueces Co.: Bolen 80 (Lk); Pil-
cher 30 con's Se Palo Pinto Co.: Mahler 1660 (Au). R Robertson Co.:
Rowell 1 8077 (Au--122260). Rusk Co.: M. De McCracken 90 (Lk). San
Patricio Co.: Cory 51259 (Mi, Se——123672); Gould & Hycka 7990 (Lk);
S. Sanderson 9) (N). San Saba Co.: L. J. Jones 3s 18 (Au—2:7795,
Au—2)812h) . “Starr Co.: Pilcher 196 . (Lk); A.D. “Wood 721 (Au—
247064, Au--262610). Taylor Co.: "Mahler 3393 (Au--28882). Tra-

2h P Hey 2 WsOrG srk Vol. 23, no. 2

vis Co.: Tharp son. [Austin, 5/2/35] (Lk), sen. [Austin, 5/9/35]
(Lk). Webb Co.: Novoa & Cantu 10 (Au--2))))815, Ip); E. Rodriguez
5 (Au—-2)5037, Ip). Wichita Co.: J. W. Stanford 291 (Lk). Will-
acy Co.: Cory 5189 (Mi). Wilson Coe: | Edw. Palmer r 1043 (Pa).
Young Co.: Vollum s.n. [Fort Belknap, 1855] (Pa). El Toro Island:
Tharp 9098 ; (Au—-122208, N), 49236 (Au--122206). MEXICO: Chihua-
hua: Stuessy 1013 (Au--257717, N). Coahuila: E. G. Marsh 1625
(Au--213)16); E E. M. Marsh 2128 (Au—21386)) . Guanajuato: Re
Pearce 2276 (Lk). Nuevo Leén: Kibler 26 (Au--29727)5; He Me Par-
ker 381 (Au--302762), h2h (Au--302763) ._ Veracruz: Garcia Saucedo
4S (Ip); Gutiérrez R. 218 (Ac).

VERBENA HALEI f. ALBIFLORA L. I. Davis

Synonymy: Verbena halei albiflora L. I. Davis ex Correll &
Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:]
1876. 1970.

Additional bibliography: Moldenke, Phytologia 9: 175. 1963;
Moldenke in Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib.
Tex. Res. Found. Bot. 6:] 131) & 1319. 1970; Correll & Johnston,
Man. Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:] 1876.
1970; Moldenke, Fifth Summ, 1: 59 (1971) and 2: 672 & 915. 1971.

VERBENA HALEI f. ROSEIFLORA (Benke) Moldenke

Additional synonymy: Verbena halei roseiflora (Benke) Moldenke
ex Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex. Res.
Found. Bot. 6:] 1876. 1970.

Additional bibliography: Moldenke, Phytologia 10: 210. 196);
Moldenke in Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex.
Res. Found. Bot. 6:] 131) & 1319. 1970; Correll & Johnston, Man.
Vasc. Pl. Tex. [Contrib. Tes. Res. penne Botec Ot 1876s 1970;
Moldenke, Fifth Summ. 1: 59 (1971) and 2: 672, 687, & 915. 1971.

VERBENA HASSLERANA Brig.

Additional synonymy: Glandularia hassleriana Schnack & Rubens,
Bol. Soc. Argent. Bot. 1339205211970.

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 571.
1965; Moldenke, Phytologia 1h: 28). 1967; Schnack & Rubens, Bol.
Soc. Argent. Bot. 13: 205. 1970; Moldenke, Fifth Sumn. 1: rg
187, & 201 (1971) and 2: 672, 688, 689, Hee & 915. 1971.

The species has been collected. in anthesis in January, in ad-
dition to the months previcusly reported, and Hatschbach de-
scribes it as an "erva do brejo". The corollas are said to have
been "violet" on Hatschbach 18321 and "red" on G. J. Schwarz 615.

Additional citations: BRAZIL: Parand: Hatschbach 18321 (Ft, N,
W--2536532). ARGENTINA: Misiones: Ruiz Huidobro 753 (Se—-
12988); G. J. Schwarz 615 (N), 5162 (N).

VERBENA HASSLERANA var. OVATIFOLIA Moldenke
Additional bibliography: Moldenke, Phytologia 13: 199. 1966;

1972 Moldenke, Notes on Verbena 25
Moldenke, Fifth Sum. 1: 177 (1971) and 2: 915. 1971.

VERBENA HASTATA L.

Additional & emended synonymy: Verbena americana, spici
multiplici, foliis urticae angustissimis, floribus purpureis
Tourn., Compl. Herb. 358. 1719. Verbena hastata ® pinnatifida
A. Wood, Class-book, ed. 2, pr. 1, 412. 1847. Verbena hastata f.
hastata Seymour, Fl. New Engl. 56. 1969. Verbena urticifolio-
hastata Bebb, in herb.

Additional & emended bibliography: Tourn., Compl. Herb. 358.
1719; Crantz, Inst. Rei Herb. 1: 573. 1766; [Retz.], Nom. Bot. ll:
Diieeeet «ciel. 1n,L., Syste Nat. ed. 13, pr. 1, .2 (1): 2
(1789) and ed. 13, pr. 2, 2 (1): 42. 1796; Balbis, Cat. Pl. Hort.
Bot. Taur. 8. 180; Balbis, Cat. Stirp. Hort. Acad, Taur. 80.
1813; Pers., Sp. Pl. 3: 346 & 347. 1819; Steud., Nom. Bot. Phan.,
ed. 1, 873. 1821; Jan, Elench. Pl. 1. 182); Beck, Bot., ed. 1,
283--28. 1833; A. Wood, Class-book, ed. 1, 268 (1845) and ed. 2,
pr. 1, 12. 1817; Beck, Bot., ed. 2, pr. 1, 285. 1848; A. Gray,
Man. Bot., ed. 1, 311. 1848; A. Wood, Class-book, ed. 2, pr. 2,
412 (18483, ed. 10, pr. 1, £22 (1885, ed. 10, pr. 2, li2 (18h9),
ed. 10, pr. 3, 412 (1850), ed. 17, 12 (1851), ed. 23, 112
(1851), ed. 29, 12 (1853), ed. 35, l12 (1854), ed. Ai, pr. 1,
hie (1855), and ed. Wl, pr. 2, 412. 1856; Beck, Bot., ed. 2, pr.
2, 285. 1856; A. Gray, Man. Bot., ed. 2, pr. 1, 298 (1856), ed.
2, pr. 2, 298 (1858), and ed. 2, pr. 3, 298. 1859; A. Wood,
Class-book, [ed. 2], pr. 1, 537. 1861; A. Gray, Man. Bot., ed.
3, 298 (1862) and ed. h, pr. 1, 298. 1863; A. iiood, Class—book,
feds ho] prs 2, 537- 18633 A. Gray, Man. Bot., ed. h, pr. 2,
298. 1864; A. Wood, Class-book, [ed. 2], pr. 3, 537. 1865; Dar-
by, Bot. South. States 7), 1866; A. Gray, Man. Bot., ed. 5, pr.
1, 340. 1867; A. Wood, Class—book, [ed. 42], pr. h, 537. 1967;
Beck, Bot., ed. 2, pr. 3, 285. 1868; A. Gray, Man. Bot., ed. 5,
pr. 2, 340. 1868; A. Wood, Class-book, [ed. h21, Disb, 5376
1868; A. Gray, Field For. & Gard. Bot., ed. 1, pr. 1, 22 (1868)
and ed. 1, pr. 2, 22. 1869; A. Wood, Class-book, [ed. 2], pr.
6, 537 (1869) and [ed. 2], pr. 7, 537. 1870; A. Gray, Man. Bot.,
ed. h, pr. 3, 298. 1870; A. Wood, Am, Bot. & Flor., ed. 1, pr. 1,
236 (1870) and ed. 1, pr. 2, 236. 1871; A. Wood, Class-book, [ed.
42], pr. 8, 537. 1872; A. Wood, Am. Bot. & Flor., ed. 1, pr. 3,
236 (1872), ed. 1, pr. lh, 236 (1873), ed. 1, pr. 5, 236 (187h),
and ed. 1, pr. 6, 236. 1875; A. Wood, Class—book, [ed. 2], pr.
9, 537. 1876; A. Gray, Man. Bot., ed. 5, pr. 8, 340 (1878) and
ed. 5, pr. 8 [9], 340. 1880; A. Gray, Field For. & Gard. Bot.,
ed. 1, pr. 3, 242. 1880; A. Wood, Class—book, [ed. 2], pr. 10,
537. 1881; Meyncke, Bull. Brooksville Soc. Nat. Hist. 1: [Fl.
Franklin Co.] 31, 1885; S. Wats. & Covlt. in A. Gray, Man. Bot.,
ed. 6, pr. 1, 402. 1889; 0. R. Willis in A. Wood, Am. Bot. &
Flor., ed. 2, 236. 1889; S. Wats. & Coult. in A. Gray, Man. Bot.,
ed. 6, pr. 2, 02. 1890; Gattinger, Med. Pl. Tenn. 64. 189); L.
H. Bailey in A. Gray, Field For. & Gard. Bot., ed. 2, 31. 1895;

2h6 PHT ON Oe A Vol. 23, no. 2

W. A. Wheeler, Minn, Bot. Stud. 2: 403. 1900; H. Kraemer, Text-
book Bot. & Pharmacog., ed. 1, 368 (1902), ed. 2, 368 (1907), ed.
3, 368 (1908), and ed. , 368. 1910; Graves, Eames, Bissell, An-
Grews, Harger, & Weatherby, Bull. Conn. Geol. & Nat. Hist. Surv.
1: [Cat. Flow. Pl.] 331. 1910; Beal, Bull. Mich. Agric. Exp. Sta.
267 [Mich. Weeds], ed. 1, 396. 1911; Britton & Br., Illustr. Fl.,
edi. 2.pr. 2a 39. 95.6 599, cid. 635, figs 355k. 19133 Ha Die
Keeler, Wayside Fls. Sum. 173--175 & 28). 1917; Rydb., Fl. Rocky
Mtns., ed. 1, 739--70. 1917; Lowe, Miss. State Geol. Surv. Bull.
17: 237. 1921; Rydb., Fl. Rocky Mtns., ed. 2, pr. 1, 739--7h0.
1922; C. A. Reed, Flow. Guide, pr. 2, 152. 1923; Tidestr., Con-
trib. U. S. Nat. Herb. 25 [Fl. Utah & Nev.], pr. 1, 469. 1925;
Pammel & King, Iowa Geol. Surv. Bull. ) (rev.): 267--269, fig.
152, 152A, & 152B. 1926; Tischler, Tabul. Biol. : 2). 1927; Har-
ger, Bull. Conn. Geol. & Nat. Hist. Surv. 48: 7h. 1930; Pellett,
Nat. Mag. 18: 185. 1931; Sefferien, Torreya 32: 125. 1932; Brit-
ton & Bre, Tllusir. Fl.., ed. 2, pr. 2, 3: 9h, 95, 59956 aes
fig. 3554. 1936; Cheymol, Bull. Soc. Chim. Biol. 19: 167—1653.
1937; St. John, Fl. SE. Wash. & Adj. Ida., ed. 1, 351, 352, & 530.
1937; Noack, Biol. Zentralbl. 57: 383--388. 1937; Anon., Chen.
Abstr. 32: 2977. 1938; F. H. & H. He Hillman, Seed Trade Buyers
Guide 1938: 137, pl. 12, fig. 6. 1938; Fischer & Harshberger,
Flower Fam. Alb. 86 & 87. 191; Gathercoal, Checklist Nat. & In-
trod. Drug Pl. [22]. 192; Britton & Br., Illustr. Fl., ed. 2;
pr. 3, 3: 9h, 95, 599, 618, & 635, fig. 355k. 193; Harvey, Er-
ickson, & Larson, Seed Trade Buyers Guide 195: 86. 195; Savage,
Cat. Linn. Herb. Lond. }. 1915; Britton & Br., Illustr. Fl., ed.
2 pre, 320 9, 95, 599; 618, & 635, far. 5554. TON Ts eee
Palmer, Fieldbook Nat. Hist., ed. 1, pr. 3, 297 & 663. 199;
Beston, Herbs & Earth 61. 1950; Abrams, Illustr. Fl. Pacific
States, pr. 1, 3: 608 & 610--611, fig. 431. 1951; R. J. Davis,
Fl. Ida. 596. 1952; H. N. & A. L. Moldenke, Pl. Bible, pr. 1
[Chron. Bot.], 5 & 326 (1952) and pr. 2 [Ronald], 5 & 326. 1952;
R. W. Br., Comp. Scient. Words 833. 195h; Rydb., Fl. Rocky lMtns.,
ed. 2, pr. 2, 739--70. 1954; St. John, Fl. SE. Wash. & Adj.
Ida., ed. 2, 351, 352, & 530. 1956; Fogg, Weeds Law & Gard. 11.
1956; S. W. Bailey, Barth, Cobble Fl. n.p. 1957; Scoggan, Nat.
Mus, Canada Bull. 1)0: [Fl. Manit.] 463. 1957; Chansler, Wild
Flow. 3: 14. 1958; Jacobs & Burlage, Ind. Pl. N. C. 221 & 251.
1958; R. McVaugh, Bull. N. Y. State Mus. 360: 196. 1958; F.
Bartley in J. C. Bartley, Bull. Ohio Biol. Surv., new ser., 1:
181. 1959; Hitchc., Cronq., & Ownbey, Vasc. Pl. Pacif. Northwest
he BON. 1959; Winter, Winter, & Van Bruggen, Check List
Vasc. Pl. S. D. 12. 1959; Ferris in Abrams & Ferris, Illustr.
Fl. Pacific States, pr. 1, : 651 & 730. 1960; Howell & McClin-
tock in Kearney & Peebles, Ariz. Fl., ed. 2, 726 & 727. 1960;
Jantzen, Emporia State Res. Stud. 9 (2): 19. 1960; Rattenbury,
Madrofio 15: 220. 1960; Martin & Barkley, Seed Ident. Man. 37 &
19h, fig. 260, pl. 235. 1961; R. M. Carleton, Ind. Common Names
Herb. Pl. 17. 1962; Hylander, Fls. Field & Forest 187, 190, &
229, fig. 6. 1962; Poindexter, Trans. Kans. Acad. Sci. 65: 09,
W10, & W12--19, fig. 2, l, & 6. 1962; St. John, Fl. SE. Wash. &

1972 Moldenke, Notes on Verbena 27

Adj. Ida., ed. 3, 380-381. 1962; Dobbs, Fl. Henry Co. 230. 1963;
E. L. D. Seymour, Wise Gard. Encycl., ed. 6, 1279. 1963; Radford,
Ahles, & Bell, Guide Vasc. Fl. Carol. 281 & 282. 1964; Rouleau in
Marie-Victorin, Fl. Laurent., ed. 2, 489 & 490, fig. 170. 196);
Ferris in Abrams & Ferris, Illustr. Fl. Pacific States, pr. 2, :
651 & 730. 1965; H. S. Fitch, Univ. Kans. Nat. Hist. Reserv. 9.
1965; Heimans, Heinsius, & Thijsee, Geillustr. Fl. Nederl. 908.
1965; Hocking, Excerpt. Bot. A.B: 226 & 537 (1965) and A.9: 290 &
367. 1965; Lakela, Fl. Northeast. Minn. 110. 1965; Sexsmith,
Canad. Field Nat. 79: 110. 1965; Gaiser & Moore, Surv. Vasc. Pl.
Lambton Co. 100. 1966; Hirata, Host Range & Geogr. Distrib. Powd.
Mild. 276. 1966; Mohlenbrock, Castanea 31: 22). 1966; F. H. Mont-
gomery, Plants from Sea to Sea 261 & 262, fig. 528. 1966; Thorn-
berry, U. S. Dept. Agr. Agric. Handb. 165: 79. 1966; Wunderlin,
Trans, Ill. Acad. Sci. 59: 143. 1966; Yotaro, Gard. Pl. World 3:
128, pl. 64, fig. 3. 1966; Abrams, Illustr. Fl. Pacific States,
pr. 2, 3: 608 & 610--611, fig. 43h. 1967; Gilkey & Dennis,

Handb, NW. Pl. 352. 1967; L. & M. Milne, Living Pl. World 212.
1967; Shinn, Univ. Kans. Sci. Bull. 46: 790, 791, 886, 887, & 928.
1967; H. Ward, Herb. Man., ed. 2, 107. 1967; W. A. Weber, Rocky
Mtn. Fl. 306. 1967; Wherry, Bartonia 37: 13. 1967; Boivin, Pro-
vanch, 2: 19) & 195. 1968; Boivin, Phytologia 16: 39 & 0. 1968;
Bulkeley, Berkshire Week Aug. 10-18 p. [17A]. 1968; Burlage, Ind.
Pl. Tex. 18h, 206, 209, 212, 218, 222, 230, & 237. 1968; W. C.
Grimm, Recog. Flow, Wild Pl. 228 & 229. 1968; Hinds & Hathaway,
Wildfls. Cape Cod 152 & 172, fig. 138. 1968; Hocking, Excerpt.
Bot. A.13: 571. 1968; Hutton, liiller, & Conrad, Castanea 22: 2h).
1968; MacKeever, Native & Naturl. Pl. Nantucket 101. 1968; Mohlen-
brock, Trans. Ill. Acad. Sci. 61: 71. 1968; Moldenke, Phytologia
16: 187. 1968; Moldenke, Résumé Suppl. 16: 1. 1968; lMunz & Keck,
Calif. Fl. 686, 687, & 1679. 1968; Munz, Suppl. Calif. Fl. 101.
1968; Peterson & McKenny, Field Guide Wildfls. 286, 316, [317], &
418. 1968; Streams, Shahjahan, & LeMasurier, Journ. Econ. Ent. 61:
997. 1968; Uphof, Dict. Econ. Pl., ed. 2, 31, Sl, 52, & 552.
1968; Anon., Checklist Vasc. Pl. West-cent. Wash. 33. 1969; Bolkh.,
Grif, Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 717. 1969; Cody,
Ind, Sem. Bot. Gard. Ottawa 1969: 22. 1969; Farnsworth, Blomster,
Quimby, & Schermerhorn, Lynn Index 6: 262 & 267. 1969; W. E. Hop-
kins, Castanea 34: 6. 1969; Jervis, Castanea 3: 115. 1969;
Krochmal, Walters, & Doughty, U. S. Dept. Agr. Forest Serv. Res.
Pap. NE-138: 266-267, 282, 285, 288, & 290. 1969; Krochmal, Wal-
ters, & Doughty, U. S. Dept. Agr. Forest Serv. Agric. Handb. 00,
pr. 1, 9, 266, & 267. 1969; R. M. Lowden, Ohio Journ, Sci. 69: 262,
263, & 280. 1969; H. L. Mason, Fl. Marshes Calif., pr. 2, 677&
877. 1969; Miller & Ehrle, Proc. Rochester Acad. Sci. 12: 62. 1969;
Rickett, Wild Fls. U. S. 3 (2): 365 & [367], pl. 111. 1969; Rydb.,
Fl. Rocky Mtns., ed. 2, pr. 3, 739--7)0. 1969; F. C. Seymour, Fl.
New Engl. 56. 1969; Swink, Pl. Chicago Reg. 427. 1969; Tidestr.,
Contrib. U. S. Nat. Herb. 25 [Fl. Utah & Nev.], pr. 2, 69. 1969;
pravton & Br., Illustr. Fl., ed. 2, pr.'5, 3: 94, 95, 599, 618, &
635, fig. 355k. 1970. [to be continued]

THE TAXONOMY OF ZEA MAYS (GRAMINEAE)

Hoch He aebtis
Department of Botany, University of Wisconsin
Madison, Wisconsin, 53706

The origin of maize has long been disputed. Of
the various theories, the oldest postulates its direct
origin by ancient human selection from a wild grass of
the genus Euchlaena, the "Teosinte" of Mexico and
Guatemala; i.e. maize is Simply regarded as a highly
domesticated and variable cultivar of Euchlaena. The
morphological steps, first clearly outlined by G.N.
Collins (J. Agr. Res. 17: 127-135. 1919), were dis-
cussed but not accepted by P.C. Mangelsdorf (Bot. Mus.
Leafl. Harvard Univ. 12: 33-75. 1945) and amplified
by W.C. Galinat (An. Rev. Gen. 5: 447-478. 1971) and
myself (H.H, Iltis The Maize Mystique. 5 pp. mimeo MS.
1976: ef: Galmnat, loc.(je2t. pp. 400), 462). athe
genetic reasons were e first outlined by G. Beadle (J.
Hered. 30: 245-247. 1939).

The close relationship of Euchlaena to Zea was
taxonomically formalized by Reeves and Mangelsdorf
(Am. J. Bot. 29: 815-817. 1942), who transferred its
two accepted species to Zea. Of these, the perennial
tetraploid Z. perennis iS the most primitive and dis-
tinctive in the genus. The diploid annual Z. mexicana
sensu lato, however, contains two major elements: the
Tripsacum-like, more primitive "Florida" Teosinte
(E. L[uxurians) from Guatemala, which does not cross as
readily with maize and other Teosinte forms, and which
deserves separate subspecific status (see below); and
the remaining races (cf. H.G. Wilkes, Teosinte: the
closest relative of Maize. Bussey Inst. Harvard. 158
pp. 1967) which cross with close to 100% fertility
with maize, and are best considered, for the time
being, under but one subspecies, conspecific with
maize as suggested by C.D. Darlington (Chromosome
Botany. Allen and Unwin, London. pp. 130-131. 1956):

"The breeding and chromosome evidence ...
agree in requiring that Euchlaena mexicana and

Zea mays should be described aS one Species,

the wild grass being today an authentic and

scarcely changed representative of what the

ancestor of Zea mays must have been."

The new combination, made informally by Il1tis
(loc. cit.), was validly published in Galinat (loc.
cit. p. 450), who, however, neglected to cite the
basSonym date and place of publication, an oversight
rectified below. Further study may in addition demand
separate subspecific recognition of other Zea mays

2),8

1972 Tltis, Taxonomy of Zea mays 29

races, especially the "Rio Balsas" and "'Chalco" popu-
iutions. (cf. Wilkes, loc. cit.).

Synopsis of Zea

1. Zea mays L. ssp. mays

2. Zea mays L. ssp. mexicana (Schrad.) Iltis, in
Galinat, An. Rev. Gen. 5: 450. 1971.
(based on Euchlaena mexicana Schrader, Index

Sem. Hort. Goett. 1832; Linnaea 8: 25. 1833).

3. Zea mays L. ssp. luxurians (Durieu) Iltis, stat.
PAA

et comb. nov.

(based on Euchlaena luxurians Durieu, Bull.
Soc. d'Acclimat. 19: 581. 1872).

4. Zea perennis (Hitchcock) Reeves et Mangelsdorf,
(based on Euchlaena perennis Hitchcock).

BOOK REVIEWS

Alma L. Moldenke

"ORGANISM AND ENVIRONMENT —- A Manual of Quantitative Ecology" by
Rezneat M. Darnell, x & 290 pp., illus., F. H. Freeman &
Co., San Francisco, California 9410). 1971. $5.75 paper-
back with looseleaf adaptation.

This intelligently planned mammal adaptable to advanced ecology
courses plans 25 laboratory and/or field experiments covering a
wide range of organisms and the following branches of ecology --
morphological, physiological, behavioral, population and commun-
ity. Illustrations, careful directional questions and biblio-
graphies enrich this work which has been previously tried out on
the author's students.

There is also the author's )7-page "Instructor's Guide"
available from the same publisher. It could also serve as an
additional guide to advanced students doing some or one of the
long term experiments.

"HOW TO KNOW THE TREES", 2nd edition, revised, by Howard A. Mil-
ler & H. E. Jaques, viii & 302 pp., illus., William C. Brown
Company, Publishers, Dubuque, Iowa 52001. 1972. $h.25
paper covered & spirally bound.

This is a reborn member of that most helpful "Picture Key
Nature Series" for amateurs, students and scientists wanting help
in easy but accurate identification work. To the first edition
of 191 and its reprinting in 196 have been added new species,
updated taxonomy, and a whole new orientation of the tree not
only as an individual specimen but also as a member of the forest
ecosystem. There is also carefully garnered material on forest
insect and fungal pests, results of air pollution, dendrochro-
nology and record specimens.

There are pictured keys for the identification of the more
common trees and for winter time identification, the typical pic-
ture glossary of this series, indexes of common and scientific
names covering 312 species, varieties and hybrids of native and
exotic trees. On p. 7 medullary is misspelled and color is
given in its British form. The line drawings of key character=
istics and the geographic distribution maps are well done.

"ENVIRONMENT AND EVOLUTION -- A Symposium on the Occasion of the
One-Hundredth Anniversary of the Foundation of the Peabody
Museum of Natural History at Yale University", edited by
Ellen T. Drake, xvi & 70 pp., illus., Yale University Press,
London & New Haven, Connecticut 06520. 1968. 135 sh or
$15 .00

250

1972 Moldenke, Book reviews 251

These papers are 16 of the invitation papers presented by out-
standing scientists at this symposium in October 1966. They cover
basic problems in many different fields of natural history such
as origins of the Metazoa, early history of land plants, paleo-
ecology of fossil spores, origins of thermoregulation, environmen-
tal photoperiod and the programming of insect development, and
outcrossing in natural populations. They all make thoughtful
reading. They all are meant to be "examples of the type of ex-
citing and stimulating research work that we wish to see under-
taken within or associated with museums of earth and life
sciences in future years."

"FUNDAMENTALS OF MYCOLOGY" by J. H. Burnett, xiii & 516 pp.,
illus. St. Martin's Press, New York, N. Y. 10010. 1968.
$13.95.

Since the classic De Bary work, now dated in part, there has
been a long wait for this excellent, well organized, definitive
text which enumerates the general features common to all (or
most) fungi and to expound the broad trends in structure, func-
tion and behavior which can be discerned in the group "of living
organisms constructed wholly of fine tubes".

The book is organized into four sections: (1) on structure
and growth of cells, hyphae, colonies and reproductive structures;
(2) on function in mutrition, transportation, translocation,
transpiration, carbohydrate catabolism, other metabolites and
ecological factors; (3) on recombination in mitosis, meiosis,
parasexual and sexual cycles; and (4) on speciation and evolution.

There are an appendix with classification of the fungi men-
tioned and indexes of references, authors and subjects. The il-
lustrations are plentiful and of fine quality.

"MORPHOLOGY OF VASCULAR PLANTS" by David W. Bierhorst, xii & 560
pp., illus., Macmillan Company, London & New York, N. Y.
10022. 1971. $14.95.

With over 2000 illustrations, including 70 percent original
publications of the author, and with a critically fine text, Dr.
Bierhorst has presented one of the two most outstanding texts
in this field. The other is by Dr. Katherine Esau, is not even
mentioned in this work, and has a different emphasis. Therefore
they do not duplicate but rather supplement each other very
well for advanced study in this field,

On the market now there are a few new books in this field
with excellent illustrations. They are mainly pictorial albums.
This one gives the readers a careful survey of the author's
studies on Equisetum, Psilotum and Schizaea. It shows the in-
tegrative relationship between ontogeny, phylogeny, physiology
and systematics, each with morphology. The book stresses the
pre-angiosperm vascular plants starting with the Lycopodiaceae.

252 PoE PS "O°LO"G. Lk Vol. 23, no. 2

"ANALYSIS OF TEMPERATE FOREST ECOSYSTEMS" edited by David Reichle,
xii & 30) pp., illus., Springer Verlag, Heidelberg, Germany
& New York, N. Y. 10010. 1970. $1.50.

This first volume of "Ecological Studies" is composed of
eighteen thoughtful papers presented at the workshop-seminar at
Gatlinburg, Tennessee, in the summer of 1968 co-hosted by the
United States International Biological Program's analysis of eco—
systems project and the Oak Ridge National Laboratory. The out-
standing contributors are about two-thirds from the United States
with the rest from eight different countries.

The editor in his preface claims "What the authors have striv-
en for has been the conceptual development and practical applica-
tion of an 'attitude' — the interrelation of ecosystem parts
which form a whole greater than the summation of its structures
and processes, and that this whole, the ecosystem, is the meaning-
ful organizational unit for ecological understanding of the en-
vironment." The papers are grouped as follows: (1) analysis of
ecosystems — quantitatively, qualitatively, phenologically; (2)
primary producers — biomass, harvesting methods, metabolic pro-
ductivity models; (3) consumer organisms in their roles as sec—
ondary production — canopy insects, birds, mammals; () decom-
poser populations — floral and faunal; (53 nutrient cycling —
micronutrients, minerals, carbon; (6) hydrologic cycles — pre-
cipitation, evapotranspiration, models.

Each paper has its own carefully prepared bibliography. There
is a subject index of world ecosystems and back cover map based
On one of Bazilevich's works of 1969.

"PLANT SPECIATION" by Verne Grant, x & 35 pp., illus., Columbia
University Press, London & New York, N. Y. 10025. 1971.
$15.00.

This book will surely become a biological classic as has its
predecessor, "The Origin of Adaptations", of 1963 which dealt
mostly with diploid sexual organisms. "Plant Speciation" de-
scribes the "evolutionary processes and patterns across the
board in higher plants (naturally neither diploid nor sexual) as
they occur in the various diverse types of genetic systems,"
dealing "at some length with the divergence and refusion of
plant species" and "the bearing of these processes on plant
macroevolution."

The book is divided into the following parts: (1)Nature of
Species, (2) Divergence of Species, (3) Refusion and its Con=
sequences, (l) Derived Genetic Systems, and (5) Evolution of
Hybrid Complexes. The bibliography is of real value. The
separate indexes are for organisms, authors, and subjects.

Extrapolating from plant microevolution to angiosperm phylo—
geny, the pattern of reticulate evolution revealed by genetic
and taxonomic studies of species groups can shed some light
[convergence] on the problems surrounding the phylogenetic

1972 Moldenke, Book reviews 253

classification of the higher categories....Evolutionary conver-
gence can be brought about by selection without hybridization, or
by the joint action of hybridization and selection...The facility
with which different plant groups can make a transition from one
ecological niche to another, so as to yield convergences in whole
character combinations, is demonstrated by hundreds of examples..
Change-overs or quantum shifts which are major in ecological
terms but relatively minor in genetical terms, can occur repeated-
ly in different phyletic lines under the guiding influence of
parallel selection alone. Within the limits of crossability the
production of convergences is stimulated still further by hybrid-
ization,"

"THE PESTS OF PROTECTED CULTIVATION — The Biology and Control
of Glasshouse and Mushroom Pests" by N. W. Hussey, W. H.
Read & J. J. Hesling, viii & 40) pp., illus., American
Elsevier Publishing Company, New York, N. Y. 10017. 1970.
$26.00.

The authors! stated purpose is the updating and enlarging of
the now 20 years old "Insect Pests of Glasshouse Crops" by Miles.
This they have achieved very effectively by adding assistance for
the expanding mushroom industry, ample newer illustrations, the
phytotoxicity factors of pesticides, immunizations to certain
chemical controls, and the advantages of biological controls.
Glasshouse pests, their effects and control, are arranged sys-
tematically, as are those of mushroom cultures.

There are appendices for chemicals used in pest control,
tolerances of plants to pesticides, abbreviations, equivalents
and dilution tables. There is also a good index.

While planned for the United Kingdom, the information is ap-
plicable to many other areas.

"BRITISH STEM- AND LEAF-FUNCI (Coelomycetes)" by W. B. Grove,
Volume I, xx & 88 pp., illus., Vol. II, ix & 07 pp., il-
lus., Verlag von J. Cramer, Lehre 3301, Germany. Reprint
1967. $16.50 paperbound,

The first volume of this fine work initially appeared from
the Cambridge University Press in 1935 and the second volume in
1937. Therefore they have not been easily available for quite
some time. Volume I covers the Sphaeropsidales to the end of
the Sphaerioideae with colorless spores, Volume II completes the
Sphaeropsidales and treats the Melanconiales -- ail in the Fungi
Imperfecti. The very clear text helps make up for the paucity
of illustrations.

There are helpful indexes of the Ascomycetes considered by
some other authors to be Coelomycetes, of hosts, and of the bi-
nomial names of the Coelomycetes treated.

It is indeed fortunate to have this valuable mycological
study readily available again. Now more folks will be able to

254 PHY? 0 LOG EX Vol. 23, no. 2

enjoy reading the peppery and valid epilogue. It is available
not only from the publisher but also from Wheldon & Wesley Ltd.
in Codicote and from Stechert-Hafner in New York City.

"GLOBAL ECOLOGY — Reading toward a Rational Strategy for Man"
by John P. Holdren & Paul R. Ehrlich, viii & 295 pp., illus.,
Harcourt Brace Jovanovich, Inc., Chicago, San Francisco, At-
lanta & New York, N.Y. 1971. $h.50 paperback.

"The book should be suitable for supplementary reading in
courses in both the biological and sociological sciences, and in
the many multidisciplinary programs in human ecology now spring—
ing up in colleges across the country. The concerned layman
with some previous exposure to discussions of populations,
resources and environment will also find much of interest here."
True indeed for this proverbial 'needle in the haystack'} It
points to and pricks the important, not over-simplified issues
poignantly and intelligently in sharp contrast to so many of the
ecologically oriented paperbacks with only straw-like contents.
The editors and many of the authors have long been leaders of
world renown for their efforts to alert scientists, political
leaders and the general public to resource realities, environ-
mental roulette, threatened species, technological circuses,
psycho-social complications, prospects for a sane economics, a
population policy, what we must do, and the cost of failure —
all the topics under which this important collection of 39 es—
says is organized.

"ELECTRON MICROSCOPY AND PLANT ULTRASTRUCTURE", by A. W. Robards,
iv & 298 pp., illus., McGraw-Hill Book Company, London, To-
ronto, Sydney, Mexico, Panama, Johannesburg, Singapore and
New York, N. Y. 10036. 1970. $10.95.

’

This book should supplement effectively many different botany
and microbiology courses and the reading of many scientists and
educated persons.

"Biological electron microscopy is an exciting and expanding
field of research, but it cannot be studied most successfully in
isolation. It is but a single technique in the armoury of the
modern biologist." The beginning three chapters with their clear-
cut exposition can make "the reader thoroughly familiar with
current methods of electron microscopy so that he may appreciate
with greater perception and enjoyment the micrographs presented
for study". Later chapters describe the following individual
components of plant cells: membranes, nucleus, mitochondria,
plastids, golgi apparatus, endoplasmic reticulum and their rela-
ted ribosomes, lysosomes, spherosomes, flagella and their rela-
ted microtubules, cell walls, and variations in ultrastructure
of different types of higher plant cells. The final chapters by
other authors deal with algae, fungi, bacteria, bluegreen algae
and viruses.

1972 Yoldenke, Book reviews 255

Many fine illustrations, selective bibliography, glossary and
index are provided.

"HAYFEVER PLANTS — their Appearance, Distribution, Time of Flow-
ering and their Role in Hayfever", 2nd revised edition, by
Roger P. Wodehouse, ix & 280 pp., illus., Hafner Publishing
Company, New York, N. Y. 10022. 1971. $11.95.

This valuable book has been brought up-to-date mainly by ad-
ditional information about causal species and by a greater, near-—
ly worldwide geographic range. The United States and especially
its southern California still register the greatest number of
cases. The causal plants have anemophilous or amphiphilous
flowers with buoyant, abundant and allergenic toxic pollen which
tends to be genus- rather than species-specific. The most in-
portant are grasses and weeds in the Gramineae, Compositae, Cheno-
podiaceae, Amaranthaceae, Plantaginaceae, Polygonaceae and trees
and shrubs in the Cupressineae, Betulaceae, Platanaceae, Fagaceae,

ceae,

The chapter on botanical literature, while sufficient for its
purpose, does not list any of the outstanding works which appeared
after 1950.

The plants are arranged systematically by families, described,
located, often illustrated for inflorescences and pollen grains
with the time of their shedding also recorded.

Regional surveys, compiled largely from reports throughout
large areas of our earth with ten sections for the United States,
give seasonal reports of affecting pollen, fungal spores, "dust"
and climate asthma,

There is a particularly good bibliography (with the word dif-
fusion misspelled on p. 268). There are a helpful glossary and a
far from complete index.

"OUR NORTHERN SHRUBS and How to Identify Them" by Harriet L.
Keeler, xxx & 539 pp., illus., Replication Edition by Dover
Publications, Inc., New York, N. Y. 1001). 1969. $3.75.

This is a welcome unabridged republication of Charles Scribner's
Sons edition of 1903 to which has been added fortunately a new
appendix of nomenclatural changes by Dr. Voss of the University
of Michigan Herbarium. It includes about 230 native, naturalized
and garden shrubs in almost 50 different plant families. All are
simply, accurately and attractively described and located. There
are over 200 full-page black/white photographs that have lost very
little definition in this printing process; diagnostic features
show up clearly.

This book has had much appeal to armchair and field botanists
and naturalists throughout this century. With this new and de-
lightfully inexpensive printing, many more can be pleased.

256 PY Tr’ Oh OGL Vol. 23, no. 2

"PLANT LIPID BIOCHEMISTRY — The Biochemistry of Fatty Acids and
Acyl Lipids with Particular Reference to Higher Plants and
Algae" by C. Hitchcock and B. W. Nichols, xiii & 387 pp.,
illus., Academic Press, London NW1L7DU & New York, N. Y. 10003.
1971. £6.50 or $19.00.

This is the fourth volume of Experimental Botany: An Internati-
onal Series of Monographs under the consulting editorship of J. F.
Sutcliffe and P. Mahlberg. It encompasses the research work es—
pecially of the past fifteen years into a coordinated, well-
explained picture of the structure, distribution and metabolism
of acyl lipids and fatty acids in higher plants and algae. It al-
so makes comparisons with related results obtained in mammalian
and microbial metabolism.

The chapters cover (1) major, minor and unusual fatty acids,
(2) acyl lipids such as glycerides, diols, acyl esters, sphingo-
lipids and cutin, (3) their presence in specific tissues of
higher plants, mosses, ferns and algae, () lipids in seeds, un-
derground structures and chlorophyllous tissue, (5) biosynthesis
of carboxylic acids, saturated and unsaturated fatty acids, (6)
biosynthesis of glyceride and other acyl lipids, (7) lipolytic
enzymes, (8) biological degradation of fatty acids by Alpha-,
Beta- and Omega-oxidation, (9) metabolic changes during senes-
cence and organogenesis, (10) lipids in membrane structure and
function, energy storage, ethylene production, and (11) fracti-
onation and analysis of the lipids and their components.

In addition to a very detailed bibliography, author index and
subject index there are "Supplementary Readings" of a more gener-
al scope and "More Recent Publications Not Considered in the
Text" arranged according to the chapters. The work is a veri-
table storehouse of information on plant lipid metabolism.

"ECOLOGY OF LEAF SURFACE MICRO-ORGANISMS" edited by T. F. Preece
& C. H. Dickinson, xvii & 640 pp., illus., Academic Press,
London NW17DU & New York 10003. 1971. $26.00.

The )\7 papers printed here were presented as the proceedings
of an international symposium at the University of Newcastle up-
on Tyne, September 1970 by active workers in the field of leaf
surface studies. They are grouped into the following five sec-
tions, each followed by the recorded interesting discussions:

(1) characteristics of leaf surfaces - anatomically, chemically,
physically, climatically - with several excellent electron micro=-
graphs, (25 saprophytes on leaf surfaces, (3) pathogens on leaf
surfaces, (1) microbiology of senescing leaves, and (5) inter-
actions on the leaf surface - host exudates, phytoalexins, antag-
onisms, inhibitions, nitrogen fixation.

Each paper carries its own bibliography. Author, systematic
and subject indexes are carefully prepared. Under the cover title
"ecology" much valuable isolated material is herein well integra-
ted in terms of basic science and is well organized for those study-
ing the role of leaf surface in crop protection & plant pathology.

B23
a <
.>
Designed to expedite botanical publication
Vol. 23 May, 1972 No. 3
;
: CONTENTS
MOLDENKE, H. N., Additional notes on the genus Verbena. XII...... Sdn
SIEREN, D. J., A new combination in the genus Euthamia .......... 304
KING, R. M., & ROBINSON, H., Studies in the Eupatorieae
4 (Asteraceae). LXVII. Grazielia nom. nov. for Dimorpholepis . 305
KING, R. M., & ROBINSON, H., Studies in the Eupatorieae
7 (Asteraceae). LX VL. A new genus, Conocliniopsis ....... 307
KING, R. M., & ROBINSON, H., Studies in the Eupatorieae
a (Asteraceae). LXIX. A new genus, Gyptidium ........... 310
“ING, R. M., & ROBINSON, H., Studies in the Eupatorieae
(Asteraceae } LXX. A new genus, PPT, i a a P 312
[OLDENKE, H. N., Five more verbenaceous novelties ............ 314
oA
MOLDENKE, PPI SERIOUS ct it RC yw ths mss yie oe d a hemes 316

f

a

a

‘ il Giger

Published by Harold N. Moldenke and Alma L. Moldenke

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

‘

Price of this number, $1; per volume, $7.50, in advance,

MAY 34 1972 or $8, at close of volume §

we -erepaARY

(EIR no ascapiarenidter ee Peet

ADDITIONAL NOTES ON THE GENUS VERBENA. XII

Harold N. Moldenke

VERBENA [Dorst.] L.

Additional synonymy: Vervena Kundu & De, Bull. Bot. Surv. In-
dia 10: 399, sphalm. 1968.

Additional & emended bibliography: Virgil., Eccl. 8: v. 64--65.
B.C.37; Propertius, Eleg. (3): v. 57--58. B.C.26; Virgil.,
Aeneid 12: v. 119--120. B.C.19; Plin. Secund., Hist. Mund. Nat.,
lib. 22, cap. 2. A.D.77; Camer. in Matth., Pl. Epit. Util. 717.
1586; L., Mat. Med., ed. 1, 6, 208, & 218. 179; Manetti, Virid.
Florent. 98. 1751; Schreb. in L., Mat. Med., ed. 2, pr. 1, 38 &
[272]. 1772; W. Curtis, Fl. Lond., ed. 1, 1 (1): pl. 41 ("2").
1775; O. F. Mill. in Oeder, Icon. Pl. Fl. Dan. h: 5, pl. 628.
1775; Bulliard, Herb. France pl. 215 & table. 1785; Schreb. in L.,
Mat. Med., ed. 5, 1 & [324]. 1787; Mavor, Gen. View Agr. Berks.
1809; Sibth. & Sm., Fl. Graec. Prodr. 1 (2): 402. 1809; Targ.-
Tozz., Diz. Bot. Ital. 2: 233. 1809; W. Curtis, Fl. Lond., ed. 2,
1 (1): pl. 41. 1817; Zerapha, Fl. Melit. Thes. 2: 77. 1831; W.
Curtis, Fl. Lond., ed. 2, 3: pl. 90. 1835; Bertol., Fl. Ital. 6:
261. 184; Schau. in A. DC., Prodr. 11: 535--557 & 736. 187;
Grech Delicata, Fl. Melit. 27. 1853; Gulia Gavino, Repert. Bot.
Malt. 7. 1855; Reichenb., Icon. Fl. Germ. 18: 52--53, pl. 91
(1292) & 103. 1857; Boswell Syme in Sowerby, Engl. Bot., ed. 3,

6: 202--203, pl. 1018. 1866; Nyman, Consp. Fl. Eur. 563. 1881;
Arcang., Compend. Fl. Ital., ed. 1, 561 & 885. 1882; Baker & New-
bould in H. C. Wats., Topogr. Bot., ed. 2, 302. 1883; Druce, Fl.
Oxf. 22). 1886; Arcang., Compend. Fl. Ital., ed. 2, i y——Lhs &
31. 1894; T. Cooke, Fl. Presid. Bomb. 2: 37. 1908; Hatton,
Craftman's Plant-book , 368, 369, & 538, fig. I & 729. 1909;
Dur. & Barr., Fl. Lib. Prodr. 193. 1910; Béguinot & Vacc., Monog.
Rapp. Colon, Minist. Aff. Est. Roma 16: [Contrib. Fl. Libia] 62.
1912; liubert, Verbénac. Util. Mat. Méd. 8--15. 1921; Holste,
Arch. Exp. Path. Pharmakol. 101: 6. 1924; Pamp., Libya 3: 145.
1927; Miller, Journ. Am, Pharm. Assoc. 17: 7h. 1928; Grove,
Brit. Stem- & Leaf-fungi, pr. 1, 1: 1) & 470. 1935; Noack, Biol.
Zentralbl. 57: [383]--3388, fig. 1—17. 1937; Cheymol, Bull. Soc.
Chem. 5: 633 & 642. 1938; Furusato, Bot. & Zool. Theoret. & Appl.
{Syokobutu Oyobi Dobutu] 8: [1303]--1311 (39--7). 190; W. Hoffm,,
Arch, Pharm. 231: 269. 1943; E. L. D. Seymour, New Gard. Encycl.,
ed. 3, 1279 & 1281 (1944) and ed. h, 1279 & 1281. 1946; E. L. Jor-
dan, Hamond Nat. Atl., pr. 1, 2, & 3, 219 (1952) and pr. h, 5, & 6,
219. 1953; Schnack & Solbrig, Revist. Fac. Agr. La Plata 29:
[255]—-266, fig. 1--h. 1953; E. L. Jordan, Hamond Nat. Atl., pr.
% 219 (195k), pr. 8, 219 (1955), pr. 9 & 10, 219 (1956), pr. 11,
219 (1957), pr. 12, 219 (1958), and pr. 13, 219. 1959; Hatton,
Handb. Pl. & Flor. Ornament , 368--369, & 538, fig. I & 729.
1960; Buchi & Manning, Tetrahedron Lett. 1960: 5. 1960; E. L. Jor-
dan, Hamond Nat. Atl., pr. 14, 219 (1961) and pr. 15, 219. 1962;
257

258 PHEYTOLOGIA Vol. 23, no. 3

Biichi & Manning, Tetrahedron 18: 109. 1962; Newcomb, Pocket Key
Comm. Wild Fls. 2h, 56, 6h, & 65. 1963; E. L. D. Seymour, New
Gard. Encycl., ed. 6, 1279 & 1281, pl. 59 (1963) and ed. 7, 1279
& 1281, pl. 59. 1964; E. L. Jordan, Hamond Nat. Atl., pr. 17, 219
(1964) and pr. 18, 219. 1965; Huynh, Denkschr. Schweiz. Natur-
forsch. Gesel. [Mém. Soc. Helv. Sci. Nat.] 85: 100. 1965; E. L.
Jordan, Hamond Nat. Atl., pr. 19, 219 (1966) and pr. 20, 219.
1967; Grove, Biblioth. Mycol. 2: 1) & 470. 1967; Kundu & De,
Bull. Bot. Surv. India 10: 399. 1968; R. I. Patel, For. Fl. Mel-
ghat 271. 1968; E. L. Jordan, Hamond Nat. Atl., pr. 21, 219.
1969; E. L. D. Seymour, New Gard. Encycl., ed. 8, 1279 & 1281.
1970; Angely, Fl. Anal. Fitogeogr. Est. S. Paulo, ed. 1, h: 825,
826, 836--8)0, & xix, maps 1391--1395. 1970; Reed & Hughes, Com—
mon Weeds U. S., pr. 1, 306--309, fig. 151 & 152 (1970) and pr. 2,
306--309, fig. 151 & 152. 1971; Hocking, Excerpt. Bot. A.18: hhh.
1971; E. L. Jordan, Hamond Nat. Atl., pr. 22, 219. 1971; Vyas &
Garg, Zeit. Pflanzenphysiol. 65: 189--19). 1971; Priszter, Delect.
Sem. Spor. Pl. Hort. Bot. Univ. Hung. 59 & 81. 1971; C. D. Adams,
Flow. Pl. Jam. 626--628, 630--632, & 846. 1972; Anon., Biol. Ab-
str. 53 (3): BA SLC. S.cfl. 1972; Vyas & Garg; Biol. Abstr.
53: 1650. 1972; Moldenke, Phytologia 23: 211--2)7. 1972.

The Sibthorp & Smith reference (1809) is often cited as "1806",
but pages 219--\)2 of this work were issued in 1809. The Boswell
Syme reference (1366) is sometimes cited as "1863", but volume 6
of this edition was not issued until 1866.

VERBENA ALATA Sweet

Additional bibliography: Angely, Fl. Anal. Fitogeogr. Est. S.
Paulo, ed. 1, 4: 838 & xix. 1970; Moldenke, Phytologia 23: 213.
Rey ges

VERBENA AMBROSIFOLIA Rydb.

Additional & emended bibliography: Darlington & Wylie, Chrom.
Atl., pr. 1, 323 (1956) and pr. 2, 323. 1961; Moldenke, Phytolo-
giae3: 213) & 228. 1972.

VERBENA BERTERII (Meisn.) Schau.

Additional bibliography: Moldenke, Phytologia 23: 182 & 230.
alts 1 (Ar

Additional citations: CHILE: Colchagua: Zollner 5351 (Ac).

VERBENA BIPINNATIFIDA Nutt.

Additional bibliography: Vyas & Garg, Zeit. Pflanzenphysiol.
65: 189--19). 1971; Vyas & Garg, Biol. Abstr. 53: 1650. 1972; Mol-
denke, Phytologia 23: 215--216 & 226. 1972; Anon., Biol. Abstr.

53 (3): BeAsS.1.C. 5.271. 1972.

Vyas & Garg (1971) report that seeds of this species stored dry
for 7 weeks developed light sensitivity. "GA at a 100-ppm concen-
tration and 30 C incubation temperature provided the optimum con-
ditions for germination. GA was capable not only of bypassing the
light requirements but also of overcoming some of the temperature
blocks of germination. Coumarin and far-red reversed the Stimula-

1972 Moldenke, Notes on Verbena 259

tion caused by GA. Preheating of seeds at 60 C abolished subse-
quent GA sensitivity of germination of those seeds in the dark.
Nitrogenous substances accelerated the dark germination of the

seeds induced by GA." "GA", of cogrse, is gibberelic acid.

VERBENA BONARIENSIS L.

Additional & emended bibliography: Noack, Biol. Zentralbl. 57:
38 & 386, fig. 11. 1937; Darlington & Wylie, Chrom. Atl., pr. 1,
323 (19565 and pr. 2, 323. 1961; Huynh, Denkschr. Schweiz. Natur—
forsch. Gesel. [Mém. Soc. Helv. Sci. Nat.] 85: 100. 1965; Angely,
Fl. Anal. Fitogeogr. Est. S. Paulo, ed. 1, : 838 & xix, map 1391.
1970; Moldenke, Phytologia 23: 216—-217 & 233. 1972; C. D. Adams,
Flow, Pl. Jam. 627, 628, & 846. 1972.

Additional illustrations: Noack, Biol. Zentralbl. 57: 386, fig.
ah 1937.

Adams (1972) states that in Jamaica this species is "Locally
common.....at roadsides and in old cultivations or gravelly waste
open places", growing ut altitudes of 1700 to 5200 feet, flowering
and fruiting for most of the year. He gives its overall distribu-
tion as "Native of subtropical S. Amer., introduced into Bermuda,
United States, Puerto Rico and elsewhere and escaping from culti-
vation". He cites from Jamaica Adams 631, Harris 9132, Jamaican
Plants 1169, and Proctor 23518.

VERBENA BONARIENSIS var. CONGLOMERATA Briq.

Additional bibliography: Angely, Fl. Anal. Fitogeogr. Est. S.
Paulo, ed. 1, 4: 838 & xix. 1970; Moldenke, Phytolo-ia 23: 217.
1972.

VERBENA BRACTEATA Lag. & Rodr.

Additional & emended bibliography: Noack, Biol. Zentralbl. 57:
[383], 384, & 386, fig. 6. 1937; Reed & Hughes, Common Weeds U.
S., pr. 1, 306--307, fig. 151 (1970) and pr. 2, 306--307, fig.
151. 1971; Moldenke, Phytologia 23: 217, 234, 237, & 239. 1972.

Additional illustrations: Noack, Biol. Zentralbl. 57: 386, fig.
6. 1937; Reed & Hughes, Common Weeds U. S., pr. 1, 307, fig. 151
(1970) and pr. 2, 307, fig. 151. 1971.

VERBENA BRASILIENSIS Vell.

Additional bibliography: Angely, Fl. Anal. Fitogeogr. Est. S.
Paulo, ed. 1, : 838 & xix, map 1391. 1970; Moldenke, Phytologia
23: 217--218, 233, & 21. 1972.

Bayliss describes this plant as 15 inches tall, with rough
stems and leaves, growing in sandy grasslands in South Africa.

Additional citations: SOUTH AFRICA: Cape Province: Bayliss
BS .3045 (W--2616807) .

VERBENA CANADENSIS (L.) Britton

Additional & emended bibliography: Noack, Biol. Zentralbl. 57:
38) & 387, fig. 15. 1937; E. L. D. Seymour, New Gard. Encycl.,
ed. 3, 1279 (1944), ed. h, 1279 (1946), and ed. 5, 1279. 1951;
Darlington & Wylie, Chrom, Atl., pr. 1, 323. 1956; E. L. D. Sey-

260 PR YT, 0 i, 0 1G 4A Vol. 23, now

mour, New Gard. Encycl., ed. 6, 1279 (1963), ed. 7, 1279 (196k),
and ed. 8, 1279. 1970; Moldenke, Phytologia 23: 218--219 & 2h2.
1972.

VERBENA CORYMBOSA Rufz & Pav.

Additional & emended bibliography: Noack, Biol. Zentralbl. 57:
38h & 386, fig. 12. 1937; Darlington & Wylie, Chrom. Atl., pr. 1,
323 (19565 and pr. 2, 323. 1961; Moldenke, Phytologia 23: 220.
1972 e

Additional illustrations: Noack, Biol. Zentralbl. 57: 386, fig.
dee ASB

Although the Darlington & Wylie work (1956) cited above and in
the bibliographies of various other species is often cited as
"1955" and appears to be so dated by the Library of Congress, the
title-page date is "1956" and Mr. Reed, Librarian of the New York
Botanical Garden, states that he sees no reason for adopting the
earlier date.

VERBENA CRITHNIFOLIA Gill. & Hook.

Additional & emended bibliography: Darlington & Wylie, Chrom.
Atl., pr. 1, 323 (1956) and pr. 2, 323. 1961; Bolkh., Grif, Mat-
vej., & Zakhar., Chrom. Numb. Flow. Pl. 715 & 716. 1969; Moldenke,
Phytologia 23: 195. 1972.

VERBENA DISSECTA Willd.

Additional & emended bibliography: Darlington & Wylie, Chrom.
Atl., pr. 1, 322 (1956) and pr. 2, 322. 1961; Angely, Fl. Anal.
Fitogeogr. Est. S. Paulo, ed. 1, h: 838 & xix. 1970; Moldenke,
Phytologia 23: 222—-22). 1972.

VERBENA DOMINGENSIS Urb.
Additional bibliography: Moldenke, Phytologia 23: 22). 1972.
Additional citations: HISPANIOLA: Dominican Republic: Gastony,
Jones, & Norris 363 (W—2657619) .

VERBENA EPHEDROIDES Cham.
Additional synonymy: Verbena sphedroides Angely, Fl. Anal.
Fitogeogr. Est. S, Paulo, ed. 1, 4: xix, sphalm. 1970.
Additional bibliography: Angely, Fl. Anal. Fitogeogr. Est. S.
Paulo, ed. 1, ): 839 & xix, map 1392. 1970; Moldenke, Phytologia
232230089722

VERBENA FILICAULIS Schau.

Additional bibliography: Angely, Fl. Anal. Fitogeogr. Est. S.
Paulo, ed. 1, : 839 & xix, map 1392. 1970; Moldenke, Phytologia
233 2316 1o0i2<

VERBENA FLAVA Gill. & Hook.

Additional & emended bibliography: Darlington & Wylie, Chrom,
Atl., pr. 1, 322 (1956) and pr. 2, 322. 1961; Moldenke, Phytolo-
gia 23: 231—-232. 1972.

|
|

}
|

1972 Moldenke, Notes on Verbena 261

VERBENA GRACILESCENS (Cham.) Herter

Additional synonymy: Verbena offinalis var. gracilescens Cham.
ex Angely, Fl. Anal. Fitogeogr. Est. S. Paulo, ed. 1, k: 839,
sphalm. 1970.

Additional & emended bibliography: Darlington & Wylie, Chrom,
Atl., pr. 1, 323 (1956) and pr. 2, 323. 1961; Angely, Fl. Anal.
Fitogeogr. Est. S. Paulo, ed. l, i: 839. 1970; Moldenke, Phyto-
logia 23: 237--238. 1972.

VERBENA HASTATA L.

Additional & emended bibliography: Noack, Biol. Zentralbl. 57:
384--386, fig. 1. 1937; E. L. D. Seymour, New Gard. Encycl., ed.
3, 1279 (1944), ed. h, 1279 (1946), ed. 5, 1279 (1951), and ed. 6,
1279. 1963; Newcomb, Pocket Key Comm, Wild Fls. 56, 64, & 65.
1963; E. L. D. Seymour, New Gard. Encycl., ed. 7, 1279 (196) and
ed. 8, 1279. 1970; Domville & Dunbar, John Burroughs Nat. Hist.
Soc. Bull. 8: 94. 1970; Duncan & Stuckey, Mich. Bot. 9: 190. 1970;
El-Gazzar & Wats., New Phytol. 69: 483 & 485. 1970; Harrington in
Frankel & Bennett, Genetic Resources 516. 1970; Joyal, Natur.
Canad. 97: 577. 1970; Moldenke in Correll & Johnston, Man. Vasc.
Pl. Tex. (Contrib. Tex. Res. Found. Bot. 6:] 1314 & 1319. 1970;
Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex. Res. Found.
Bot. 6:] 1876 & 1877. 1970; Rickett, Wild Fls. U. S. (3): 5h0,
[543], & 799, pl. 177. 1970; Wise, Rhodora 72: 52). 1970; Young-
ken & Tashiro, Rep. Contr. Research Prop. U. S. Army Med. Res.
Develop. Command Washington Contract DA9193MD3037. 1970; Brown &
Wherry, Bartonia 0: 13. 1971; Cody, Ind. Sem. Bot. Gard. Ottawa
1971: 20. 1971; Cochrane, W. E. & M. M. Rice, Mich. Bot. 10: 183.
1971; Farnsworth, Pharmacog. Titles 5: Cumul. Gen. Ind. 1971;
Farnsworth, Pharmacog. Titles 7 (7): xiv & title 12547 (1971) and
7 (8): xix & title 15169. 1971; Krochmal, Walters, & Doughty, U.
S. Dept. Agr. Forest Serv. Agric. Handb. 00, pr. 2, 9, 266, &
267. 1971; Ownbey & Monserud, Common Wild Fls. Minn. 312. 1971;
Rickett, Wild Fls. U.S. 5 (2): [h55), 456, & 665, pl. 152. 1971;
Thilenius, U. S. Dept. Afr. Forest Serv. Res. Paper RM.71: 2.
1971; Moldenke, Fifth Sum, 1: 1-23, 25, 27, 30, 3h--43, 45
49--53, 59, 62, 63, 101, 203-205, & 370 (1971) and 2: 548, 6h9—
652, 656, 660, 661, 668, 669, 672—67h, 679, 686, 688, 690, 695,
696, 698, 702, 704, 705, 708, 732, 793, 915, & 967. 1971; Molden-
ke, Phytologia 22: 97 (1972) and 23: 195. 1972

Additional & emended illustrations: Britton & Br., Illustr.
wi., 6d. 2, pr. 1, 3: 95, fig. 355). 1913; C. A. Reed, Flow.
Guide, pr. 1, 152 [in color]. 1916; H. L. Keeler, Wayside Fls.
Sum. 174. 1917; C. A. Reed, Flow. Guide, pr. 2, 152 [in color].
1923; Pammel & King, Iowa Geol. Surv. Bull. ); (rev.): 267 & 268,
peg. 152 & 152A. 1926; Britton & Br., Illustr. Fl., ed. 2, pr. 2,
3: 95, fig. 3554. 1936; Noack, Biol. Zentralbl. 57: 386, fig. 1.
1937; F. H. & H. H. Hillman, Seed Trade Buyers Guide 1938: 137,
pl. 12, fig. 6. 1938; Fischer & Harshberger, Flower Fam. Alb. 87.
191; Britton & Br., Illustr. Fl., ed. 2, pr. 3, 3: 95, fig.
3554. 1943; Harvey, Erickson, & Larson, Sced Trade Buyers Guide
1945: 86. 1945; Britton & Br., Illustr. Fl., ed. 2, pr. 4, 3: 95,

262 P HOY tO! Le rk Vol. 23, no. 3

fig. 3554. 1917; Abrams, Illustr. Fl. Pacific States, pr. 1, 3:
608, fig. 431. 1951; Hitchc., Cronq., & Ownbey, Vasc. Pl. Pacif.
Northwest : 2)6. 1959; Martin & Barkley, Seed Ident. Man. 37,
pl. 235, & 194, fig. 260. 1961; Hylander, Fls. Field & Forest 187,
fig. 6. 1962; Rouleau in Marie-Victorin, Fl. Laurent., ed. 2,
489, fig. 170. 1964; F. H. Montgomery, Plants from Sea to Sea
261, fig. 528. 1966; Yotaro, Gard. Pl. World 3: pl. 6h, fig. 3
[in color]. 1966; Abrams, Illustr. Fl. Pacific States, pr. 2, 3:
608, fig. 431. 1967; Gilkey & Dennis, Handb. NW. Pl. 352. 1967;
Bulkeley, Berkshire Week Aug. 10-18, p. [17A]. 1968; W. C. Grim,
Recog. Flow. Wild Pl. 229. 1968; Hinds & Hathaway, Wildfls. Cape
Cod 152, fig. 138. 1968; Peterson & McKenny, Field Guide Wildfls.
[317] (in color). 1968; Krochmal, Walters, & Doughty, U. S. Dept.
Agr. Forest Serv. Agric. Handb. 400, pr. 1, 267. 1969; Krochmal,
Walters, & Doughty, U. S. Dept. Agr. Forest Serv. Res. Paper NE.
138: 267. 1969; Rickett, Wild Fls. U. S. 3 (2): [367], pl.
{in color] (1969) and h (3): [543], pl. 177 [in color]. 1970;
Britton & Br., Illustr. Fl., ed. 2, pr. 5, 3: 95, fig. 355k.
1970; Ownbey & Monserud, Common Wild Fls. Minn. 312. 1971; Rick-
ett, Wild) Fls.U.'S. 5 (2): [455], pl. 152) [in color]. 197ae

The Madrofio (1960) reference in the bibliography of this spe-
cies is sometimes credited to Solbrig, while Bolkhovskikh (1969)
cites it as "Darlington, Document. Chrom. Numb. Pl......", but it
appears to be a continuation of a series begun by Rattenbury and
not otherwise accredited since then.

Weber (1967) characterizes V. hastata as follows: "Spikes
slender, acute at the apex; leaves deep green; flowers deep blue
or purple" and describes its habitat as "Swamps and irrigation
ditches, piedmont valleys and plains. Common, flowering in late
summer" in the Rocky Mountains region. Harrington (1970) has
found that its seeds have a life span of 39 years. Poindexter
195-1, was used to determine a chromosome count of n = 7 and pol-
len fertility of 99 percent. Ownbey & Monserud (1971) report
that the corollas are "sometimes violet or pink", implying that
the normal form is blue. Gilkey & Dennis (1967) describe the
flowers as "purplish-blue, opening upward on the spike, only a few
flowers of a spike open at one time". Cumbie 89 was used for an-
atomical studies. aS

Gattinger (189) tells us that in Tennessee this species is
"Very common in sandy soils", flowering in July, "The herb and
root used medicinally".

Grimm (1968) describes the inflorescence as follows: "The
small violet-blue flowers, hardly more than 1/8 inch across, are
arranged in several long and very slender clusters at the tip of
the stem and its branches" and says of the plant that "It is usu-
ally common in moist fields, meadows, and along streams; bloom-
ing between June and September", giving its range as "N. B. to B.
C. south to Fla., Tex., and Calif." Hitchcock, Cronquist, & Own-
bey (1959) describe the corolla [of what is probably var. scabra]
as "blue or violet" and give its habitat in the Pacific Northwest
as "Ditch banks and moist low ground" and its range as "B. C. to

1972 Moldenke. Notes on Verbena 263

N. S., s. to Calif., Ariz., and Fla.", blooming from July to Sep-
tember. Hausman (198) describes the corollas as "violet-blue,
rarely white or pink" and describes the habitat as "Moist fields,
meadows, roadsides, waste places. Nova Scotia south to Florida:
west to British Columbia, Nebraska, Arizona. June—Sept.".
Violet-purple actually seems to be the corolla color in the typi-
cal form of this species, but this color usually appears as blue
to observers who are red-colorblind. There are, however, distinct
blue-flowered, pink-flowered, and white-flowered forms.

Recent collectors have found this plant growing on brushy
hillsides, in damp open weedy fields, in cattail thickets along
rivulets (in Missouri), in sandy muck (in Oklahoma), on lake mar-
gins and in deep sand of dune areas (in Texas), and in damp soils
generally. Lowe (1921) found it in "low damp open pastures" in
Benton and Pontotoc Counties, Mississippi; Wheeler (1900) records
it from Houston County, Minnesota; Dobbs (1963) describes it as
"Frequent to common on moist shores and in low moist to wet de-
pressions" in Henry County, Illinois. Rickett (1969) describes
its habitat and distribution as "in meadows, fields, prairies,
swamps from Nova Scotia across Canada and southward to Florida,
Tennessee, Missouri, northern Texas, and California" and comments
that "This has been called simpler's-joy, from some medicinal
use" [actually this name belongs to V. officinalis L. and has
been misapplied to the American V. hastata only in books].

Poindexter encountered V. hastata in "areas characterized by
sandy wastes and stabilized sand dune pastures". Radford, Ahles,
& Bell (1964) record it from Ashe, Orange, Warren, Washington,
and Watauga Counties, North Carolina, and report that the plant
grows there in low fields, bogs, and marshes, and is infrequent
in North Carolina, blooming there from June to October. Lakela
(1965) cites Lakela 11172 & 1562) from Minnesota, while Hutton,
Miller, & Conrad (1968) record it from Jefferson County, West
Virginia, and Wherry (1967) from Delaware County, Pennsylvania.
Hopkins (1969) cites his no. 1022 from Pope County, Illinois, and
Mohlenbrock (1968) also reports it from that county; Domville &
Dunbar (1970) describe it as "common in damp places" in Ulster
County, New York, flowering there "in summer", while Ownbey & Mon-
serud (1971) aver that it inhabits "Pastures, wet meadows and
swamps.....throughout the state" of Minnesota. Cochrane (1971)
refers to it as "common" in "low open ground" in Rock County, Wis-
consin, while Jantzen (1960) reports it from Stafford County,
Kansas.

Lowden (1969) found it growing on Squaw Island, Ohio; Joyal
(1970) cites Joyal 1217 and Rouleau 1125 from Soeur Island, Que-
bec; Wise (1970) found it on Isle au Haut in the outermost part
of Penobscot Bay, Knox County, Maine, while Duncan & Stuckey
(1970) tell us that it was once on Big Chicken and Green Islands,
Michigan, since they saw it there in 1939, but that it had dis-
appeared from both islands by 1970. Howell & McClintock (1960)
cite Macdougal 566 from Coconino County and Covers & Palmer 279
and Toumey s.n. from Yavapai County, Arizona. St. John (1962)

264 P Hieyor.0 "0 "G*r iA Vol. 23, no. 3

says that the species is "occasional in meadows and streambeds,
Upper Sonoran life zone" in southeastern Washington and adjacent
Idaho and in his 1937 work that it is "occasional, meadows and
stream banks" in southwestern Washington — in both cases it is
undoubtedly var. scabra Moldenke to which reference is being made.
Winter and his associates (1959) state that V. hastata grows in
"moist valleys and draws over the state" of South Dakota, Bartley
(1959) records it from Jackson County, Ohio, and Tatnall (196)
avers that it is "common in meadows" in the Piedmont and on the
Coastal Plain of Delaware and Maryland "as far south as Wicomico
Co.", blooming there from "late June through August". Swink (1969)
tells us that it is found in "All 22 counties" of the Chicago
region and is there "Frequent in marshy ground, associating with
Asclepias incarnata, Boehmeria cylindrica, Calamagrostis canaden-
sis, Eupatorium maculatum, Eupatorium perfoliatum, Impatiens cap-
ensis, Leersia oryzoides, Lycopus americanus, Lythrum alatun,
Penthorum secoides, Scirpus americanus, Scirpus atrovirens, and
Typha latifolia. In moist meadows its associates include Agrostis

ior, Pycnanthemum virginianum, Spartina pectinata, and Veronicas—
trum virginicum." Montgomery (1966) gives its Canadian distribu-
tion as "N. S. to B. C.", which asrees with what most authors
state. Wunderlin (1966) affirms that it is "local in open ground”
in Carroll County, Illinois, and cites his nos. 148 and 208,
while Rouleau (196);) tells us that it occurs only in western and
central Quebec and Sexsmith (1965) records it from Grosse Ile in
that province.

Fischer & Harshberger (191) describe the flowers of this
plant very interestingly: "The individual flowers attempt to be
2-sided like the mints but have never quite succeeded in getting
their mouths open". Ward (1967) almost unbelievably reduces V.
hastata to synonymy under V. officinalis L.!

Dobbs (1963) reports that V. hastata is "One of the many old
time medicinal plants and still said to be of some commercial im-
portance, the herb being used"; Krochmal, Walters, & Doughty
(1969) say that in Appalachia the "herbaceous parts [of this
plant are] reportedly used as an astringent, antipyretic, vulner-
ary, antirheumatic, tonic, and expectorant", while Uphof (1968)
says "Dried herb, collected at flowering time, used medicinally
as expectorant, diaphoretic. Contains verbenalin, a glucoside".
Others have reported that the root is used as an emetic, tonic,
expectorant, sudorific, and vulnerary, and that it contains irri-
tant properties. Chemists report that the plant contains gluco-
sides, verbenalin, a bitter principle, and tannin. Verbenaloside
is reported from it by Cheymol (1937).

Keeler (1917) has found that V. hastata is pollinated by bees
and flies and is nectar-bearing. Shinn (1967) found it to be
visited regularly by the bee, Calliopsis nebraskensis. Streams
and his associates (1968) report that nymphs of the tarnished

1972 Moldenke, Notes on Verbena 265

plant-bug infest it in Connecticut, as they also do V. urticifol-
ia, Erigeron annuus, E, strigosus, Oenothera biennis, , Daucus car—
ota, and species ies of Solidago, and are there east tiaed by the
wasp, Leiophron pallipes, forming a biologic control of the bug
that would otherwise spread from weeds to cultivated crops.

Hirata (1966) records the powdery mildew, Erysiphe cichoracea-
rum P, DC., infesting V. hastata in Canada and the United States,
while Thornberry (1966) records the same fungus and also the fol—
lowing: Phyllosticta verbenicola G. Martin (in New Jersey), Puc-
cinia vilfae Arth. & Holw,. (a rust) in South Dakota and from In-
diana to to Oklahoma, and Septoria verbenae Rob. (a leaf-spot) from
Vermont to Wississipp!, Texas, and South Dakota,

Savage (195) tells us that in the Linnean Herbarium, London,
sheet 12 under genus 35 is labeled "hastata" in Linnaeus' own
handwriting. The "blue vervain" illustrated by Pellett (1931) is
surely V. stricta Vent., rather than V. hastatal

Additional common names reported for Verbena hastata, besides

those previously recorded by me, are "American blue vervain",
"simplers joy", "tall vervain", *ntall wild verbena", and tyerbain's

Material of V. hastata has been misidentified and distributed
in some herbaria as Ve s V. engelmannii Moldenke, Veronica longifolia
L., and Teucrium canadense var. occidentale Te Gray) McClintock
& Epl. On the other hand, the Prater 60 and A. E. Radford 51,
distributed as V. hastata, are actually - Vv. bonariensis L.; L.; ‘eu A.
Duke 0196 is V. hau iioasis Vell.; Lindzey 293 TS Ve canadensis
(L.) Brit Britton; "Cory 50298, Daubenmire s.n. [August i Patol 328
lon 842, Elmer 532, ME F. Henderson s.n. wn. [June 19, 192], Kreager
469, E. E. P. She: Sheldon S.n. pS eT I, Stephens & Brooks 25064, and
D. R. Swingle s.n. [July ra 1933] are V. hastata var. . scabra Mol-
denke; “Mahler 861; is V. macdougal ii Heller; Drumm Drummond s.n. ne (St.
Louis, N. Am. N. An.] “is xV. “Moechina Moldenke; Hapeman s.n S$.n. ~[Minden,
Aug. 23, 1939] is xV. perriana Moldenke; Bolen 59 is. is V. pumila
Rydb.; B. Martin 86 4. is V. stricta Vent.; Barkley & Ha'if 003 is
V. supina L.; L.; and H H. B. “Parks s.n. [June 23, 1946] is V. xutha
Lehm. The G. N. Jones 17655, | cited below, has its leaves slightly
roughened and mi may possibly | be var. scabras

Additional citations: QUEBEC: Huntingdon Co.: Ernest & LeBlanc
61-200 (Go). ONTARIO: Carleton Co.: Senn 1958 (Ld d). MAINE: Cum-
berland Co.: H. N. Moldenke 18893 (Se——18),850) . NEW HAMPSHIRE:
Cheshire Co.: “Ottley 5750 (Lk). Merrimack Co.: Barkley & Court-
ney 0USO39 (Ac, Go). VERMONT: Chittenden Co.: S. F. Blake 1957
[Herb. Blake 3058] (Ld). MASSACHUSETTS: Berkshire Co.: Jones &
Jones 17129 (N). NEW YORK: Jefferson Co.: J. I. Northrop s.n.
[Thousand Islands, July 16, 1889] (N). Otsego Co.: Collector un-
determined s.n. [E. Springfield, 7.76] (N). Seneca Co.: Chicker-
ing s.n. (Ovid, July 1857] (W--2605971). NEW JERSEY: Morris Co.:

266 P EY T0 Eb 0"G t's Vol. 23, no. 3

Moldenke & Moldenke 25633 (Ac, Ft, Rf, Z)- Somerset Co.: A. L.
Moldenke s.n. [August 2, 1969] (Ps—1019) ; H. N. Moldenke 8090
(Se--113522). Union Co.: A. L. Moldenke s.n. [Mountainside, A Aug.
20, 1968] (Ps--39). PENNSYLVANIA: Huntingdon Co.: H. A. Wahl
1388 (Se). MARYLAND: Montgomery Co.: Killip 3671 (Au--122289).
DISTRICT OF COLUMBIA: Sudworth 605 (Mi), son. [22 June 1890] (Mi).
WEST VIRGINIA: Preston Co.: Davis } & Davis 897k (Se—127868) .

OHIO: Butler Co.: J. Ferguson s.n s.n. ~ [July | 1S. 9, 1932] (Go). ILLINOIS:
Champaign Co.: R. A. Conover s.n. wn. [August 7, 1946] (N). Cook Co.:
H. R. Bennett s.n. ~fAugust 2h, 1957] (Se--1789)3). McHenry Co.:
He Re Bennett s.n. [August 31, 1957] (Se--180360). Tazewell Co.:
Vette Chase Chase 3229 29 (Se--2011)9). INDIANA: Greene Co.: Friesner
2229) (Au--12228)). Pulaski Co.: Friesner 9768 (Sd=--317))). IOWA:
Allamakee Co.: Snead s.n, [August 12, 19],6] (Au--122285). KEN-
TUCKY: Bath Co.: M. E. '. Wharton 3226 (Mi). Estill Co.: M. E. Whar-
ton 3015 (Mi). Jefferson Co.: Gunn 2020 [South. App. Bot. Club _
19: 1831] (W--2)33751). Lewis Cow: Me. M. E. Wharton 5086 (Mi). Mad-
ison Co.: M. E. Wharton 5633b (Mi). “MICHIGAN: Branch | Cow: Co Ae
Brown 2605 CM est. Joseph Co.: H. R. Bennett 2707 (W--2))5852) «
WISCONSIN: Dane Co.: G. N. Jones 17655 (N). Dodge | Co.: Preston
19 (Ws). Grant Co.: Beetham s.n SNe ~[Sept. 19, 1966] (Ws). Manito-
woc Co.: Demler 30 (Ws). Ro Rock Co: P. J. Scott san. [9-17-66]
(Ws). MINNESOTA: Martin Co.: Bartlett & | Grayson | 1382 (N). Mur-
ray Co.: Jensen-Haarup s.n. (Slayton, summer 1913] (Ac). Ramsey
Co.: S, F. Blake 161 [Herb. Blake 1200] (Ld). KANSAS: Reno Co.:
Poindexter 195-14 (N). MISSOURI: Clark Co.: J. A. Steyermark
68918 (N). St. Louis: Muehlenbach 3695 (Ac). NEBRASKA : Dundy
Co.: Richardson & Robertson 1256 (N). OKLAHOMA: Harper Co.:
Stratton 3368 (Lk). TEXAS: Hemphill Co.: Cumbie 89 (Lk); E. L.
Reed 1,03 (Lk); C. M. Rowell 10596 (Lk). Oldham Co.: York & &
Rodgers 363 (Lk). LOCALITY OF COL COLLECTION UNDETERMINED: : Herb. A.
Brown s.n. (N) -

VERBENA HASTATA f. ALBIFLORA Moldenke

Additional bibliography: Hausman, Begin. Guide Wild Fls. 303.
1948; Moldenke, Phytologia 16: 91. 1968; Moldenke, Fifth Summ, 1:
1618, Ei 0; & 45 (1971) and 2: 650, 673, & 915, 1971.

VERBENA HASTATA f. CAERULEA Moldenke

Synonymy: Verbena americana, spici multiplici, foliis urticae
angustissimis, floribus caeruleis Tourn., Compl. Herb. 358. 1719.

Bibliography: Tourn., Compl. Herb. 358. 1719; Hocking, Excerpt.
Bot. A.9: 290. 1965; Moldenke, Phytologia 13: 200. 1966; Moldenke,
Fifth Summ. 1: 15 (1971) and 2: 649—651 & 915. 1971

Many authors have spoken of the corollas of Verbena hastata L.

as being "blue", and the standard common name for the species is

1972 Moldenke, Notes on Verbena 267

"Blue Vervain". In reality, the corollas of the typical and wide-
spread form of the species are violet-purple, but usually appear
as blue to persons who are more or less colorblind to red. It is
not certain how widely the truly blue color-form of the species is
distributed since only the reports of definitely non-colorblind
observers can be relied on in this matter.

VERBENA HASTATA f. ROSEA Cheney

Additional bibliography: Graves, Eames, Bissell, Andrews, Har-
ger, & Weatherby, Bull. Conn. Geol. & Nat. Hist. Surv. 1: {cat.
Flow. Pl. Conn.] 331. 1910; Harger, Bull. Conn. Geol. & Nat.
Hist. Surv. 8: 7). 1930; Hausman, Begin. Guide Wild Fls. 303.
1948; Dobbs, Fl. Henry Co. 230. 1963; Moldenke, Phytologia 1):
285. "1967; F - C. Seymour, Fl. New Engl. 56. 1969; Ownbey & Monse-
rud, Common Wild Fls. Minn. 312. 1971; Moldenke, Fifth Sum. 1:
15—19, Big 23, Ol, HO, Ud, eS (1971) and 2: 673 & 915. 1971.

Dobbs (19635 records this form from "low moist south border of
Shadow Lake", Henry County, Illinois. Ownbey & Monserud (1971)
record it from Minnesota, but, unfortunately, do not give a pre-
cise locality.

VERBENA HASTATA var. SCABRA Moldenke

Synonymy: Verbena hastata scabra Moldenke ex Correll & John-
ston, Man. Vasc. Pl. Tex. (Contrib. Tex. Res. Found. Bot. 6:]
1876. 1970.

Additional bibliography: Moldenke, Phytologia 16: 91. 1968;
Munz, Suppl. Calif. Fl. 101. 1968; Moldenke in Correll & Johnston,
Man. "Vasc. Pl. Tex. (Contrib. Tex. Res. Found. Bot. 6:] 131, &
1319. 1970; Correll & Johnston, Man. Vasc. Pl. Tex. (Contrib.

Tex. Res. Found, Bot. 6:] 1876. 1970; Moldenke, Fifth Sum. 1:
15, hO—3, 50, 52, 59, 62, 6h, & 65 (1971) and 2: 67h, 793, &
915. 1971.

Recent collectors have found this variety growing in wet
meadows, on sandhill prairies, and in roadside ditches, flowering
and fruiting in September and October. Crampton describes it as
a perennial; Cory calls it "abundant in moist floodplain of riv-
er", growing to 18 dm, tall, while Andrew & Alison Moldenke refer
to it as a "weed in horse pasture, sympatric, no hybrids". The
only recorded common name for it is "blue vervain" and it has
hitherto been misidentified as typical V. hastata L.; in fact,
the Dillon 842, L. F. Henderson Son. {June 19, '92], Kraeger 169,

E. P. Shel Sheldon S$ -11167, and D. R. ” Swingle s.n. [July 21, 1, 1933),
cited below, were previously reported by me as typical V. hastata
before I recognized the validity of the variety. It is very
probable that most, if not all, of the far western collections
previously cited by me under V. hastata will prove to represent
this variety. Munz (1968) describes the variety as "Lf.-—bases
more rigid; lvs. conspicuously scabrous on upper surface, often +t
conspicuously pubescent beneath. - Cited from Modoc, San Joaquin
and Shasta cos.; to B. C., Mont."

Additional & emended citations: MANITOBA: Fisher Distr.: J. M.

268 PUBS LeOKGer A Vol. 23, no. 3

Macoun s.n. [July 30th, 188] (Pa). MINNESOTA: Hennepin Co.:
Daubenmire s.n. [August 1, 1933] (Se--177625). MONTANA: Gallatin
Co.: D. R. Swingle s.n. [July 21, 1933] (Se--26665). Ravalli Co.:
Hitchcock & Muhlick 21937 (Se--19602). NEBRASKA: Logan Co.:
Stephens & Brooks 2506) (N). TEXAS: Hemphill Co.: Cory 50298
(Mi). WASHINGTON: Grant Co.: T. H. Scheffer s.n. [July 31, 195]
(Se--102238). Okanogan Co.: Dillon 8h2 (Se--9),50) ; Elmer 537
(Vt). Skamania Co.: Suksdorf 10572 (Se--118381). Stevens Co.:
Kreager 69 (Se--68697). Yakima Co.: L. F. Henderson s.n. [June
19, 192] (Se—119),3) ; Kruckeberg 255 (Se--181162); Moldenke &
Moldenke 3123 (Rf, Z). OREGON: Multnomah Co.: Gilkey s.n. [Port-
land, July 31, 1935] (Au--122290, Se--131497); E. P. Sheldon S.
11167 (Se--10920). CALIFORNIA: San Joaquin Co.: Crampton 319
(Ca--1278077) .

VERBENA HATSCHBACHI Moldenke

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 571.
1965; Moldenke, Phytologia 13: 249. 1966; Moldenke, Fifth Sum.
1: 177 (1971) and 2: 67) & 915. 1971.

VERBENA HAYEKII Moldenke

Additional bibliography: J. F. Macbr., Field Mus. Publ. Bot.
13 (5): 614. 1960; Moldenke, Phytologia 13: 29. 1966; Moldenke,
Résumé Suppl. 17: 3. 1968; Moldenke, Fifth Sum. 1: 1h3 (1971)
and 2: 691 & 915. 1971; Moldenke, Phytologia 23: 185. 1972.

Vargas Calderén describes this plant as "prostrate", while
Hodge refers to it as a "creeping roadside plant, flowers blue-
purple", It has been collected in fruit in December (in addition
to the months previously reported by me) and material has been
misidentified and distributed in herbaria under the name V. poly-
stachya H.B.K. Te

Additional citations: PERU: Apurimac: Vargas Calderén 8712
(Ac). Cuzco: Vargas Calderén 20066 (Ac). Junin: Hodge 6237 (W—

eee

VERBENA HERTERI Moldenke

Additional synonymy: Verbena herteri "Moldenke ex Herter" a-
pud Hocking, Excerpt. Bot. A.8: 226. 1965.

Additional bibliography: Moldenke, Phytologia 16: 91. 1968;
Moldenke, Fifth Summ. 1: 177 & 189 (1971) and 2: 67) & 915. 1971.

VERBENA HIRTA Spreng.

Additional bibliography: Barroso, Rodriguésia 32: 70. 1957;
Angely, Fl. Anal. Paran., ed. 1, 571. 1965; Dombrowski & Kuniyo-
shi, Araucariana 1: 14. 1967; Moldenke, Phytologia 16: 91, 199,
& 205. 1968; Angely, Fl. Anal. Fitogeogr. Est. S. Paulo, ed. 1,
h: 839, map 1392. 1970; Moldenke, Fifth Summ. 1: 177, 201, & 370
(1971) and 2: 690 & 915. 1971.

1972 Moldenke, Notes on Verbena 269

According to Hatschbach this species grows from a xylopodiun.
Its corollas are described as "lilac" on Hatschbach 14781 & 22)87
and Hatschbach & Guimardes 18439. Barroso (1957) cites a
15664, Burret & - Brade 16042, Occhioni s.n. [Abril de 1921], a
"RB" "RB" 161,58, 22564, 32 32898, & & 35233, giving the geographic reer
bution of the spe species as Sdo Paulo, Minas Gerais, and Rio de Jan-
eiro. Actually it is also found in Bahia, Parand, Rio Grande do
Sul, and Santa Catarina, Brazil, as well as in Misiones, Argenti-
na, "and cultivated in Germany.

* Additional citations: BRAZIL: Parand: Angely 3579 (Ca—1276799) ;
Hatschbach 1781 (W--256728), 22487 (Mi); Hatschbach & Guimar&es
18439 (Ac); Hatschbach & Haas 15669 (Rf); Reitz & Klein 1748) (N,
W—258331). Rio de Janeiro: Pabst 9317 (Herb. Brade 4202] (Ac).
Santa Catarina: Smith & Klein 13885 @).

VERBENA HIRTA var. GRACILIS Dusén

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 571.
1965; Moldenke, Phytologia 16: 91 & 205. 1968; Moldenke, Fifth
Summ. 1: 177 (i971) and 2: 915. 1971.

Additional citations: BRAZIL: Paran4: Reitz & Kiein 17616 (N,
W--25),8333), 17883 (W--25),8332) . 7 tT

VERBENA HISPIDA Ruiz & Pav.

Additional & emended bibliograp as Persie, Ope PL. 32. 3hfs
1819; Steud., Nom. Bot. Phan., ed. 1, 873. 1821; Reiche & Phil.,
Fl. Chil. 5: 283 & 28h. 1910; es "Biol. Zentralbl. 57: 38h &
386, fig. 9. 1937; Cheymol, Bull. Soc. Chim. Biol. 19: 1647—-
1653. 1937; Anon., Chem. Abstr. 32: 2977. 1938; J. F. Macbr.,
Field Mus. Publ. Bot. 13 (5): 613, 615, & 622. "1960; Padmanabhan,
Phytomorph. 14: 49. 1964; Meyer & Weyrauch, Inst. Mig. Lill.
Misc. 23: 64 & 123. 1966; Alzate, Pl. Medic., ed. 7, 264--265.
1968; Kunkel, TEP hy: 351. 1968; Moldenke, Phytologia 16:
91--92. 1968; Moldenke, Résumé Suppl. 16: ie 1968; Farnsworth,
Blomster, Quimby, & Schermerh., Lynn Index 6: 267. 1969; Bolkh.,
Grif, Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 717. 1969; An-
gely, Fl. Anal. Fitogeogr. Est. S. Paulo, ed. 1, ): 839. 1970;
Moldenke, Fifth Summ. 1: 137, 143, 177, 184, 187, 193, 201, 203,
205, 206, & 370 (1971) and 2: 655, 665, 671, 673, 67h, 679, 680,
686, 695, 702, 705, & 915. 1971.

Additional’ illustrations: Noack, Biol. Zentralbl. 57: 386, fig.
9. 1937.

Recent collectors have found this plant growing in the beds of
dry brooks, fruiting (in addition to the months previously re-
ported by me) in Jamuary and February. The corollas are descri-
bed as "lilac" on Luna 382 & 453 and Terribile 238. The Cabrera
(1957) reference in the - bibliography of this species is sometimes
cited as "1958".

Alzate (1968) affirms that this species can be used as a sub-
stitute for V. officinalis L. (which see) in New World medicine.
Cheymol (1937) reports the isolation of stachyose from this plant.

270 P BY TO LoOnG igh Vol. 23, no. 3

Kunkel (1968) informs us that the so-called V. hispida Rufz &
Pav. of Reiche in Anal. Mus. Nac. Chile (1903) is actually Vv.
litoralis H.B.K. Macbride (1960) cites the following Peruvian
collections: Arequipa: Isern 165. Cajamarca: Raimondi s.n,. [Cas-
cas]. Cuzco: Herrera s.n. wn. [Prov. de Cercado], Hicken sen. [Sicu-
ani], Pennell 1 1 118), Soukup 259. Huancavelica: Stork & & Horton
10820. Hudnuco: Macbride ride 170k, Rufz & Pavon s.n. [near Hudmuco].
Junin: Killip & Smith 21855 & | & 21925, Macbride 1006. Lima: Rai-
mondi s.n. [near Lima], Pennell T 1582. Moquegua: Isern 166. Pr Puno:
Pennell 1309. He gives "Ecuador, Ch Chile and Argentina" as the
extra-Peruvian distribution, but, of course, it occurs also in
Brazil, Paraguay, and Bolivia, is naturalized in Germany and
Switzerland, and is widely distributed elsewhere in cultivation,

Material of V. hispida has been misidentified and distributed
in some herbaria as Junellia sp. On the other hand, the Hodge
6237, distributed as V. hispida, is actually V. ayo Moldenke.

~~ Additional citations: ECUADOR: Azuay: Asplund 17805 (N). PERU:
Apurimac: Vargas Calderén 8881 (Ac). Arequipa: Vargas Calderén
19426 (Ac), 19530 (Ac). Cuzco: Vargas Calderén 12722 (W--
2520259) , 13902 (Ac). Moquegua: Vargas Calderén 17985 (Ac). Puno:
Ugent & Ugent L543 (W--2558165). BOLIVIA: Cochabamba: R. F.
Steinbach mB (W—2533070) . CHILE: Tarapac4: Zollner you (Ac).
ARGENTINA: Catamarca: Luna Risso 875 (N), s.n. [1-1-1917] (N);
A. Reales 1039 (N). Chaco: Buratovich 577 (N), 752 (N). Formosa:
ie Morel 2908 (N), 3476 (N), 3893 (N), )102 (1), “139k (N), 6331
(N), > 6392 (N) (N). Jujuy: A. Reales les 615 (njceal Mendoza: S Semper s.n.
(7/v/ul (N). Salta: Luna Ra02 (N), | 453 (N). Santiago del Este-
ro: Balegno 96 (N); Luna na 257 ( (N). Tucum4n: A. Reales 891 (N,

Rf) ; Terribile 238 Cs, a we

VERBENA HOOKERIANA (Covas & Schnack) Moldenke

Additional & emended bibliography: J. A. Clark, Card Ind. Gen.
Sp. Var. issue 191. 1945; Darlington & Wylie, Chrom. Atl. Pravde
323 (1956) and pr. 2, 323. 1961; Rufz Leal, Revist. Facult. Cienc.
Agrar. Mendoza 11: 173 & 17h, fig. Bs 1963 Troncoso in Cabrera,
Fl. Prov. Buenos Aires 5: 133 & 137. 1965; Moldenke Phytologia.
16: 92. 1968; Rufz Leal, Biol. Abstr. 9: 3256. 1968; Bolkh.,
Grif, vatvej., & Zakhar ., Chrom. Numb. Flow. Pl. 71S" & 717. 1969;
Moldenke, Fifth Summ. 1:°201 (1971) and 2: 521, 66h, & 915. 1971;
Voldenke, Phytologia 23: 195. 1972.

Additional illustrations: Ruiz Leal, Revist. Facult. Cienc.
Agrar. Mendoza 11: 17), fig. 8. 196).

Bartlett found this plant growing "in flat Larrea cuneifolia
association, loess soil" in Catamarca, Argentina. na. It has been
collected in fruit in December (in addition to the months previ-
ously reported by me). The corollas are described as having been
"yellow" on P. Garcia 927, but this was probably a mistake in ob-

1972 Moldenke, Notes on Verbena 271

servation or transcription. Troncoso (1965) states that they are
normally violet in color. She gives the distribution of the spe-
cies as "Regiones secas y 4ridas del centro y oeste del pafs [Ar-
gentina], extendiéndose hasta el extremo mAs austral de las pro-
vincias fitogeogréficas del Monte y Espinal", citing Bartlett
19918 (at San Isidro) and J. F. Molino s.n. (at Buenos Aires), and
commenting that "Todas las citas de V. Vv. crithmifolia Gill. et Hook.
dadas para el Provincia [Buenos Aires] deben referirse a esta es-
pecie". Material has been widely misidentified and distributed in
herbaria, as she says, as V. crithmifolia.

Additional citations: ARGENTINA: Buenos Aires: Eyerdam, Beetle,
& Grondona 2343 (Se—119015). Catamarca: H. H. Bartlett 20231
(N); Brizuela la lOh (Se—130303), 614 (XN), 101) ("), 106), (N); 0" or
Donell & Meye & Meyer 5155 5155 (N), 5220 (Ny. Cérdoba: | Balegno 0 35 (N); He
He H. Bartlett 201h9 (. (N); Sota ta 317 (N), 407 (N), L43 (N). La Rioja:
M. P. Gomez s.n. (Herb. Inst. Miguel Lillo 1068Lh) (N). Mendoza:
Krapovickas & Cristébal 1462) (Z). Rio Negro: O'Donell 1693 (N),
1749 (N). San Luis: Varela 547 (N), 62) (N), 653 (N), 732 (N).
Santiago del Estero: Balegno 210 (N, S); P. Garcia 927 (N).

VERBENA HUMIFUSA Cham.

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 571.
1965; Moldenke, Phytologia 16: 92 & 100. 1968; Moldenke, Fifth
Summ. 1: 177, 187, & 190 (1971) and 2: 662, 675, 692, 693, & 915.
1971.

Additional citations: PARAGUAY: Hassler 979 (Ca—95056l) .

XVERBENA HYBRIDA Voss

Additional synonymy: xVerbena hybrida "Voss ex Riimpler" apud
Hocking, Excerpt. Bot. A.8: 226. 1965.

Additional & emended bibliography: E. Twining, Ill. Nat. Ord.
Pl. 2: 104, fig. 2. 1855; Paque, Fl. Anal. & Descr. Prov. Namr
& Luxemb. 47. 1902; Blakeslee, Science, new ser., 8: 298--299.
1918; Tischler, Tabul. Biol. h: 2h. 1927; Gough, Gard. Book Mala-
ya ahs. 1928; Navarro de Haydon, Flor. Comun. Puerto Rico [16].
1936; Noack, Biol. Zentralbl. o7: (383]—387, fig. 1h. 1937;
Scott-Moncrieff & Sturgess, Chem. Abstr. 3): 119. 190; Furusa-
to, Bot. & Zool. Theoret. & Appl. [Syokobutu Oyobi Dobutu] ai
{(2303]—13n2 (39:7). 1940; Fischer & Harshberger, Flower Fam.

. 87. 191; E. L. D. Seymour, New Gard. Encycl., ed. 3, 1279
(isis) and ed. 4, 1279. 1946; Blank, Bot. Rev. 13: 260, 287, &
288. 1947; E. L. Palmer, Fieldbook Hat. Hist., ed. l, pr. es 297
& 663. 1919; Bu le, De Seymour, New Gard. Encycl., ed. ee eae
1951; Bodger, Libr. Hort. Tllustr. Electros of Fls. ay a
& 105, pl. 5Shg, hile, 573c, 956a, 1137f, & 3018f. 1952; Geissman
& Hinreiner, Bot. Rev. 18: 133. 1952; Mauser, Samen [zurich]
54>, 1, fig. 8053. 1953; Kuck & Tongg, Mod. "Trop. Gard. 213.
1955; Thomas & Hendricks in Magill, Holden, & Ackley, Air Pollu-
tion Handb. 9: 1—l. 1956; Darlington & Wylie, Chrom. Atl., pr.

272 P Bey. 5. 0) By 0) Geek Vol. 23, noses

1, 323. 1956; Ledbetter, Zimmerm., & Hitchc., Contrib. Boyce
Thomp. Inst. 20: 275 & 277. 1959; Withamfogg in Kiaer & Hancke,
Gard. Fls. in Colour, pr. 1, 127 & 190, fig. 487--l89 & 91. 1959;
Howell & McClintock in Kearney & Peebles, Ariz. Fl., ed. 2, 72h.
1960; Buia, Fl. Mic. Illustr. Rep. Pop. Rom. [50] & Ol, pl. 16.
1961; Darlington & Wylie, Chrom. Atl., pr. 2, 323. 1961; Haramaki,
Proc. 17th Northeast. Weed Control Conf, 213—217. 1963; S. R. &
R. S. Fujimoto, U. S. Pl. Patent 2271. 1963; E. L. D. Seymour,
New Gard, Encycl., ed. 6, 1279, pl. 59 (1963) and ed. 7, 1279,
pl. 59. 1964; Anon., Hortic. Abstr. 3h: 732. 1964; Bardi, Trop.
Gard. Burle Marx 123, pl. 177. 1964; Buckley & Cavaye, Greenhouse
Gard. Grass : 20. 1964; Radford, Ahles, & Bell, Guide Vasc. Fl.
Carol. 281 & 282. 196; Fugii, U.S. Pl. Patent 2)0 (196) and
2537. 1965; Anon., Hortic. Abstr. 35: 518. 1965; J. & L. Bush-
Brown, Am. Gard. Book, ed. 4, 315, 327, & 671. 1965; Haramaki,
Weed Abstr. 1): 256. 1965; Heimans, Heinsius, & Thijsee, Geillustr.
Fl. Nederl. 908. 1965; Mitchell & Knight, Journ. Exper. Biol. 16:
11. 1965; Ohwi, Fl. Jap. 763. 1965; Stefan, Babes, & Juga, Lucr.
Sti. Stat. Exp. Dobregea, Vol. Omag. 19-196), 265-—-275. 1965;
Yotaro, Gard. Pl. World 1: 131, pl. 66, fig. 1. 1965; Altman &
Dittmer, Environ. Biol. 316, 62), & 641. 1966; Anon., Hortic. Ab-
str. 36: 582. 1966; J. G. Barton in Novdc, Pict. Encycl. Pl. &
Fls. 403 & OS, fig. 801. 1966; Burkill, Dict. Econ. Prod. Malay
Penins. 2: 2266. 1966; T. H. Everett, Reader's Digest Compl. Book
Gard. 113, 153, 553, & 569. 1966; Greensill, Trop. Gardening 79,
98, & 22. 1966; Hirata, Host Range & Geogr. Distrib. Powd. Mild.
276-277. 1966; Mishra, Sci. & Cult. 32: 199--201. 1966; Thorn-
berry, U. S. Dept. Agr. Agric. Handb. 165: 79. 1966; G. Abraham,
Green Thumb Book 195 & 220. 1967; Anon., Hortic. Abstr. 37: 17h.
1967; Arora & Khoshoo, Indian Journ, Genet. & Pl. Breed. 27:
275--277. 1967; Coon, Fragrance & Frag. Pl. 142. 1967; H.C. D.
de Wit, Pl. World High. Pl. 2: 183. 1967; G. W. Kelly, Rocky Mtn.
Hort., ed. 2, 125--127, 148, & 223. 1967; L. & M. Milne, Living
Pl. World 212. 1967; Tingle, Check List Hong Kong Pl. 38. 1967;
Zukowski in Pawlowskiego, Fl. Polsk. 11: 65. 1967; Anon., Hortic.
Abstr. 38: 779 & 1037. 1968; Anon., Deutsch. Gartnerborse 68:
72--78. 1968; Faroy Inc. Houston Tex. Card 25-89. 1968; Hartmann
& Kester, Pl. Prop., ed. 2, 683. 1968; Horodysky, Diss. Abstr.
28: 4937. 1968; Horodysky, Bioresearch Ind. : 5682. 1968; Lau-
rie, Kiplinger, & Nelson, Commercial Flow. Forcing 83. 1968;
Moldenke, Phytologia 16: 187--188, 193, & 195. 1968; Moldenke,
Résumé Suppl. 16: 6, 7, 13, & 28 (1968) and 17: 3 & 7. 1968; A. L.
Moldenke, Phytologia 16: 55. 1968; A. & I. Nehrling, Easy Gard.
Drought-resist. Pl. 254. 1968; Roos, Natl. Geogr. 133: 726 & 732.
1968; Stucchi, Fiori 11: 131. 1968; Verbena Snow White, Northrop
King Seeds 526. 1968; R. Webber, Early Horticult. 133 & 208.
1968; Withamfogg in Kiaer & Hancke, Gard. Fls. in Colour, pr. 5,
127 & 190, fig. 487—l89 & 491. 1968; Bolkh., Grif, Matvej., &
Zakhar., Chrom. Numb. Flow. Pl. 717. 1969; Farnsworth, Blomster,
Quimby, & Schermerh., Lynn Index 6: 267. 1969; Fior, Nostr. Fl.,
ed. 3, 570. 1969; Hay & Synge, Dict. Gard. Pl. 49 & 369, pl.

392. 1969; Liogier, Fl. Cub. Supl. 123. 1969; Shumway, Ann. Cat.

1972 Moldenke, Notes on Verbena 273

99: . 1969; Rickett, Wild Fls. U. S. 3 (2): 362 (1969) and (3):
539 & 799. 1970; Angely, Fl. Anal. Fitogeogr. Est. S. Paulo, ed.
1, 839. 1970; E. L. D. Seymour, New Gard. Encycl., ed. 8, 1279.
1970; Graf, Exot. Pl. Man., ed. 1, 411. 1970; Moldenke in Correll
& Johnston, Man. Vasc. Pl. Tex. (Contrib. Tex. Res. Found. Bot.
6:] 1316 & "1323—~132h. 1970; Moldenke in Menninger, Flow. Vines
338. 1970; Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex.
Res. Found. Bot. 6:] 1876. 1970; Bright of America, Summersville,
W. Va., 25.GC.70 Petunias & Verbenas. 1970; Anon., Biol. Abstr.
52 (16): BeA.S.I.C. S.269. 1971; Furia, Biol. Abstr. 52: 8978.
1971; G. W. Park Seed Co., Park's Flow. Book 1971: 75. 1971; Mol-
denke, Pitth*Somm. Lt 19, 25, 30) 353750535 59, (6552755 9d 98,
120, 137, 143, 177, 20, 231, 238, 262, & 370 (1971) and 2: 651,
657, 659, 662, 663, 665, 666, 670, 672, 674-677, 680, 682, 683,
689, 690, 693, 697, 700, 703, 706, 733, & 915. 1971; Priszter,
Delect. Sem. Spor. Pl. Hort. Bot. Univ. Hung. 59. 1971; Westcott,
Pl. Disease Handb., ed. 3, 745. 1971; Moldenke, Phytologia 22:
31), & 497 (1972) and 23: 221. 1972; D. Burpee, Burpee Seeds 1972:
1, 56, & 57, pl. 3177 & 4345. 1972; W. J. Park, Park's Flower
Book 1972: 87. 1972.

Additional & emended illustrations: E. Twining, Ill. Nat. Ord.
Pl. 2: 104, fig. 2 (as V. chamaedrifolia) [in color]. 1855; Noack,
Biol. Zentralbl. 57: 386, fig. 1). 1937; Fischer & Harshberger,
Flower Fam. Alb. 87. loki; Bodger, Libr. Hort. Illustr. Electros
of Fls. 61 & 95, pl. She, “Wits. 573¢ {in color], aa 1137f, &
3018f [in color]. 1952; Viauser, Samen [Zurich] 1953: ue fig. 8053
[in color]. 1953; Withanfogg in Kiaer & Hancke, Gard. Fls. in
Colour, pr. 1, 127, fig. 487--489 & 91 [in color] . 1959; Buia,
Fl. Mic. Illustr. Rep. Pop. Roms [50], pl. 16.1961; S. R. & R.
S. Fujimoto, U. S. Pl. Patent 2271. 1963; Be Lb. D. Seymour, Wise
Gard. Encycl., ed. 6, pl. 59 [in color] (1963) and ed. 7, pl. 59
[in color]. 1964; Bardi, Trop. Gard. Burle Marx 123, pl. 177.
1964; Fugii, U. S. Pl. Patent 240 [in color] (1964) and 2537
{in color]. 1965; Heimans, Heinsius, & Thijsee, Geillustr. Fl.
Nederl. 908 (as V. chamaedrifolia) . "1965; Yotaro, Gard. Pl. World
Bren 66.) fig. 1 [in color]. 1965; J. G. Barton in Novak, Pict.
Encycl. Pl. & Fls. 03, fig. 801. 1966; Greensill, Trop. Garden-
ing 98. 1966; Roos, Natl. Geogr. 133: 726 & 732 [in color]. 1968;
Faroy Inc. Houston Tex. Card 25-89 [in color]. 1968; Verbena Snow
White, Northrop King Seeds 526 [in color]. 1968; Withamfogg in
Kiaer & Hancke, Gard. Fls. in Colour, pr. 5, 127, ite 487—h89 &
491 [in color]. 1968; Hay & Synge, Dict. Gard. Pl. 49, pl. 392
[in color]. 1969; Shumway , Ann. Cat. 99: h. 1969; Bright of Amer-
ica, Summersville, W. Va., 25.GC.70 Petunias & Verbenas [in color].
1970; Graf, Exot. Pl. Man., ed. 1, 11. 1970; G. W. Park Seed Co.,
Park's Flower Book 1971: 75 [in color]. 1971; D. Burpee, Burpee
Seeds 1972: 1 [in color], 5, 6,:& 57, pl. 3177 [in color] & 434s
fan color]. 1972; W. J. Park, Park's Flower Book 1972: 87 [in
color]. 1972.

The Tischler (1927) reference in the bibliography above is
cited by Bolkhovskikh and his associates (1969) as "Tischler,

27h PHY DO LeOrGriva Vol. 23, no. 3

Pflanzl. Chrom. eee on

Pennington tells us that in Chihuahua, Mexico, this entire
plant is decocted into a potion taken for "dolor del estémago"
(stomache-ache). Blank (197) reports that investigations have
shown that pelargonidin derivatives are sometimes dominant and
sometimes recessive to delphinidin derivatives in this plant; al-
so, monosides are sometimes dominant and sometimes recessive to
dimonosides. Mixtures of anthocyanins occur due to incomplete
dominance or to modifying factors. Geissman & Hinreiner (1952)
report the presence in this plant of a naturally-occurring antho-
cyanin called delphin, which is a 3,5-diglucoside of delphinidin.
Farnsworth and his associates (19695 record the isolation by
Scott-Moncrieff & Sturgess (190) of delphinidin-3,5-diglycoside,
delphinidin-3-monoglycoside, and glucose in the flowers of xV.
hybrida. Ledbetter and his associates (1959) report that "Ver-
bena sp. var. L. A." is one of 32 species and varieties of plants
susceptible on exposure to ozone at concentrations of 0.1 to 1.0
ppm. in a circulating stream of washed air. Very young leaves
with no functional stomata were resistant to ozone, but others
spotted necrotically.

Thornberry (1966) and Westcott (1971) record the following
disease-causing organisms as attacking this plant: Botrytis cin-
erea Pers. (a flower blight, in Massachusetts), Erysiphe cichor-
acearum P. DC. (a powdery mildew, general), Macrophomina phaseoli
(a charcoal rot, Oklahoma), Phymatotrichum omnivorum (Shear) Dug.
(a root rot, Texas), Rhizoctonia solani Kuehn (a root rot, New
York), Thielaviopsis basicola (a root rot, Pennsylvania), Meloi-
dogyne sp. (a root rot nematode, Maryland), Sclerotium bataticola
Taub. (a charcoal stem rot, Oklahoma), Sphaerotheca humuli (P.DC.)
Burr. (a powdery mildew, Puerto Rico), and Heterodera marioni
(Cornu) Goodey (a root knot nematode, Maryland) .

According to Hirata (1966) the following fungi attack xV. hy-
brida: Erysiphe cichoracearum (in Australia, Bermuda, Sweden, and
United States), E. polyphaga (in France, comma Italy, and Swit-
zerland) , Sphaerotheca fuliginea and 5S. verbenae (in Armenia,
Australia, Italy, Japan, Lithuania, Puerto Rico, Romania, Se a
ted States), Microsphaera ferruginea (in Italy and Sweden) ,

Oidium verbenae (in Argentina, Australia, Egypt, New reerenane
Portugal, Rhodesia, South Africa, Switzerland, Tasmania, and Uni-
ted States), all powdery mildews.

The Bush=-Browns (1965) add the oblique-banded leaf-roller to
the known pests of this plant. This is a small caterpillar,
banded with yellow and light-green, feeding on the foliage and
flowers, rolling and webbing the leaves with silken threads. The
female moth deposits her eggs in tiny masses on the foliage, where
they hatch in a few weeks. The larvae are full-grown in one
month, pupate within the rolled leaves, and emerge as adults sev-
eral weeks later. There are normally two broods per season. Kelly
(1967) reports that a blister beetle constitutes a pest of this

1972 Moldenke, Notes on Verbena 275

plant in the Rocky Mountain region.

The Nehrlings (1968) list this among their annual drought-
resistant plants. Laurie, Kiplinger, & Belson (1968) aver that
verbenas of this type can be sown in early March for spring sale
by dealers, but better plants with more flowers can be produced
by sowing the seed in early February and carrying the plants at
53° F. Picking will make compact bushy plants of the taller
grandiflora types, although natural dwarf varieties are avail-
able. Tingle (1967) reports it as being cultivated in Hongkong.
Thornberry (1966) notes that it is "widely grown for ornament as
a summer annual in the North [of the United States] and as a win-
ter annual in the South."

According to Mishra (1967) terminal shoot—tips of this plant
were treated experimentally with colchicine at concentrations of
up to 0.1 percent for h, 8, and 16 hours. All the treated seed-
lings survived, although growth was temporarily retarded. They
later became taller than control seedlings, had 12--23 branches
as compared to only 5—8 on the controls, and produced larger and
more serrated leaves. Stomatal size increased considerably in
the treated plants with a corresponding decrease in mumber per
unit of leaf-surface and the pollen-grains were significantly
larger with a high sterility percentage. The treated plants re-
mained vegetative longer, but the flowering period was also more
prolonged than in the controls.

Coon (1967) notes that "The flowers are not noted for frag-
rance, yet there's a special kind of quiet, haunting odor to
verbenas which you'll not get to know unless you grow them."
Burkill (1966) says that "V. hybrida does not fruit in Malaysia,
as the climate is too moist. The presumed parents are perenni-
al, but V. hybrida must be grown as an annual from imported seed,
and......is impatient of damp and not always a success in wet
weather." Ohwi (1965) records it as cultivated in Japan. Rad-
ford, Ahles, & Bell (196) affirm that the plant is commonly
cultivated in the Carolinas and rarely escaped along roadsides
and in sandy alluvium in Darlington and Kershaw Counties, South
Carolina, flowering there from late March to May.

Webber (1968) informs us that verbenas were introduced into
England from South America in the mid-sighteenth century and
then used mainly as bedding plants. Stucchi (1968) adds that the
hybrid was first introduced into Italy between 1829 and 1839.
Kuck & Tongg (1955) reiterate that it is very difficult to grow
in tropical gardens. It has apparently escaped from cultivation
and become naturalized in Paran4, Brazil, where it is now found
in "Locaes umidos de varzea", according to Hatschbach. The cor-
ollas on these naturalized specimens are "violet" (Hatschbach
14769] or "rosa avermelhada" [Manfrin 4]. In Honduras the natur-
alized plants have "rose-red" corollas [Molina R. 1708].

Culture directions given by Northrup King (1968) are "use:
cut flower, low bedding and border. Where to plant: any garden
soil under full sunlight. When & how to plant: indoors: For
earliest flowering, start seeds in flats, peat pots or boxes in

276 P Wr o, LOG rs. Vol. 23, no. 3

early spring. Cover seed 1/8 inch deep, pressing firmly into

soil. Keep moist and in sunny window. When seedling [is] 2 in-

ches tall, thin or transplant. When weather [is] warm, transplant
into garden spacing about 1 foot apart. Water well and protect
from sun a few days. Outdoors: Seed can be sown as soon as weath—
er [is] warm outdoors covering about 1/) inch deep. Firm soil
well over seeds. Later thin plants to 1 foot apart....germina-
tion [is] slow on Verbena, usually 3--l weeks so allow ample time
for sprouts to appear." Park (1971) calls the plants "Delicious-
ly fragrant - Always in bloom. Brilliantly colored blooms, fine
plants for beds, borders, edgings and cuttings. Seed started
outside in May will bloom from midsummer until frost; started
early inside, June until frost. Fine for covering the bare spot
left by early spring bulbs or ground cover among summer or fall
bulbs. Easily grown." Priszter (1971) offers seed as his no,

1669.

The following horticultural strains of xV. hybrida are listed
in recent horticultural literature and are recorded here because
they are either new or their descriptions differ somewhat from
the descriptions previously recorded by me:

Amethyst (Park) -- "that lovely sought-after mid-blue, very dwarf
8--12-inch compact plants", Sparkle type; (Burpee) -- "a
dwarf Sparkle of low cushionlike plants covered with laven-
der-blue fls. ideal for low-growing edges, borders, and
beds ."

Blaze (Hay & Synge) -- dwarf, 9-inch plants, with brilliant scar-
let flowers; (P) All-American Selection winner, with large,
bold, bright, solid, scarlet flower-heads 3 inches across;
(Burpee) -- "a Dwarf Sparkle of dazzling bright scarlet
color",

Bush Type Mixed (Park) -- broad flat-tonped bushy plants 8--10
inches tall, 12--15 inches across, flower-heads almost
touching each other entirely covering the plant.

Candidissima (Withamfogg) -- a dwarf hybrid, 10 inches tall, with
white flowers.

Chiquita —- this is V. tenera var. maonetti Regel, which see.

Compliment (Park) -- Bush type, with salmon-orange flowers, each
with a yellow eye.

Crystal (Park) -—- Sparkle type, with white flowers.

Dannebrog (Withamfogg) -- plants 10--16 inches tall, blooming
early, the flowers light-red with white centers.

Dazzle (Park) -- Sparkle type, with red flowers.

Defiance -- see under xV. corrupta Moldenke.

Delight (Park) -- Sparkle type, with coral flowers.

Dwarf Sparkle -- free-blooming compact spreading strain forming a
floral carpet all season, 6 inches tall (Burpee).

Firefly (Greensill) — bright scarlet flowers, meritorious, seed
not easy to harvest, shake seed—heads into a bag daily when
the sun is hottest.

Firelight (Mauser, Park) -- Bush type, with red flowers.

Giant Salmon Queen (Mauser) -- with salmon-red flowers.

1972 Moldenke, Notes on Verbena 277

Ideal Florist Mixture (Park) —— early flowering plants weaving a
carpet of color in summer gardens, only 8 inches tall but
spreading to 18 inches across the ground, "the plants clothe
themselves in a dazzling array of colors ranging from white
to scarlet and pink to purple, each with a dash of white in
the center".

Madame Du Barry (Park) -- "an entirely new color in verbenas,
brilliant purplish-red", vigorous plants 10 inches tall,
“large umbels on strong stems, free-branching but distinctly
compact in habit, good for bedding and cutting".

Mammoth Royal Bouquet (Hay & Synge) -- colorful mixed strain with
white Auricula eye, plants 12 inches tall.

Multiflora Gigantea (Burpee) —— free-blooming bush type bearing
large trusses of big flowers 10 inches tall. "Finest Mixed"
colors include rose-pink, lavender, blue, white, violet,
salmon-pink, scarlet, and red shades, many "eyed". Burpee
No. 435.

Rainbow Mixed (Hay & Synge) -- compact, 9 inches tall, very
early flowering.

Roselight (Mauser, Park) -- Bush tyne, rose-colored flowers.

Royal Blue (Withamfogg) -- 18 inches tall, with deep purple-blue
flowers.

Royal Carpet (Fugii) -- see Plant Patent description.

Ruffled Pink (Burpee) -- a lovely deep salmon-pink with neat
clusters of ruffled flowers and handsome deep-green foliage;
the colorful plants are ideal for edgings, beds, ground
covers, rock gardens and window-boxes, thriving even in poor
soil.

Salmon Queen (Park) -- Dush type, with salmon-pink flowers.

Snow Carpet (Fugii) -- see Plant Patent description.

Snow White (Mauser, Northrup, Park) -- Bush type, with compact
clusters of fragrant flowers on attractive compact plants,
pure glistening white flowers, plants 1 foot tall.

Sparkle (Park) —- with scarlet flowers.

Sparkle Mixed (Hay & Synge, Park) —- dwarf, compact, mound-
shaped plants, 6--10 inches tall, free-flowering and complete-
ly covered with bloom (no open spots), the full color mixture
includes warm scarlet through rose and pink to softer salmon,
lavender, and white, midc-blue, purple, and bright combina-
tions.

Splendor (Park) -- Sparkle type, with royal purple flowers with a
clear white eye.

Verbena Candycane (Fujimoto) -- see Plant Patent description.

Verbena Grandiflora Brightness (Bodger).

Verbena Grandiflora Mixed (Bodger).

Verbena Grandiflora Royale (Bodger).

Verbena Hybrida Compacta Leuchtfunk (Mauser).

To record the selling retail price of verbena seeds in 1972, it
may be mentioned here that the prices charged by Burpee (1972) for
"Ruffled Pink" may be taken as typical: "No. 3660 pkt. 75 cents;
1/16 oz. $2.25; 1/8 oz. $4.25; 1/h oz. $8.00."

An additional vernacular name for these plants is "tuinverbena"

278 P) Ret pFoO) L0G. TA Vol. 23, no. 3

in Dutch-speaking countries. The plant has been cultivated at 2800
feet altitude in Mexico. Material has been misidentified and
distributed in some herbaria under the names V. chamaedrifolia
Juss., V. delticola Small, and V. teucrioides Gill. & Hook.
Additional citations: MEXICO: : Chihuahua: Pennington 27 (Au—
264116). PERU: Lima: Camona 1590 (Rf). VENEZUELA: Mérida:
Lépez—Palacios 2212 (Z). BRAZIL: Paran&: Hatschbach 14769 (W—
256,569); Manfrin in (Herb. Fac. Farmacia 5861] (W--25277h)9) «
CULTIVATED: Honduras: Molina R. 14708 (N). New Jersey: A. L. Mol-
denke s.n. [July 9, 1968] 68] (Ps—-121)._ New Zealand: W. R. . Sykes
106/64 (N (Nz--153295). Pennsylvania: H. N. Moldenke 14979 (Se--
135328). MOUNTED ILLUSTRATIONS: fig. 2761 [in color] Jy (Z); Ry
fig. 2793 [in color] (Z); Vaughan Seed Cat. fig. 5715, 5716,
5725 [in color] (Z).

XVERBENA ILLICITA Molcenke

Additional & emended bibliography: J. D. Poindexter, Trans.
Kans. Acad. Sci. 65: O09, 10, 15, 17, & 418. 1962; Moldenke,
Phytologia 16: 94. 1968; Swink, Pl. Chicago Reg. 428. 1969; Nol-
denke, Fifth Sum. 1: B55) ha, 43, 45, 47, & 52 (1971) and 2:
673, 698, 70h, 705, & 915. 1971.

Poindexter, in a note appended to Poindexter 207-21, cited
below, states that pollen fertility in the plant of which this
is a specimen was only 17 percent and the chromosome number was
n= 7.

The color photographs taken by my son, Dr. Andrew R. Moldenke,
and cited by me in a previous installment of these notes, are all
of A. R. Moldenke 1275 from Calhoun County, Illinois.

Additional citations: ILLINOIS: Henderson Co.: H. N. Patterson
s.n. [Oquawka, July] (Pa). KANSAS: Cherokee Co.: Poindexter 207-
21 (N). MISSOURI: Saint Louis: Eggert s.n. [Prairies, 1) August
1875] (Pa).

VERBENA INAMOENA Bria.
Additional bibliography: Moldenke, Phytologia 13: 203. 1966;
Moldenke, Fifth Summ. 1: 188 (1971) and 2: 915. 1971.

VERBENA INCISA Hook.

Additional synonymy: Glandularia incisa (Hook.) Troncoso ex
Cabrera, Fl. Prov. Buenos Aires Ba: 135. 1965.

Additional & emended bibliography: A. Gray, Man. Bot., ed. 3,
levi. (1862), ed. lh, pr. 1; Ixvi.,(1863),, and: ed... pr. 2.) Java
186; A. Gray, Field For, & Gard. Bot., ede) 2 pres 2), (1868)
and ed. 1, pr. 2, 242. 1869; A. Wood, Am. Bot. & Flor., mG as
pr. 1, 235 (1870), ed. 1, pr. 2, 235 (1871), ed. 1, pr. 3, 235
(1872), ed. 1, pr. h, 235 (1873), ed. 1, pr. 5, 235 (187k), and
ed. 1, pr. 6, 235. 1875; A. Gray, Field For. & Gard. Bot., oan Le
pr. 3, 22. 1880; 0. R. Willis in A. Wood, Am. Bot. & Flor., ed.
2, 235. 1889; Burkart, Excerpt. Bot. A.5: 586. 1962; Angely, Fl.

1972 Moldenke, Notes on Verbena 279

Anal, Paran., ed. 1, 571. 1965; Troncoso in Cabrera, Fl. Prov.
Buenos Aires 52: 133 & 135. 1965; J. A. Clark, Card Ind. Gen. Sp.
Var. issue 29. 1965; Yotaro, Gard. Pl. World 1: 131. 1965; Bur-
kill, Dict. Econ. Prod. Malay Penins. 2: 2266. 1966; Moldenke,
Phytologia 16: 188, 19h, 208, & 212. 1968; Moldenke, Résumé Sup-
pl. 16:- 22 & 28. 1968; Moldenke in Menninger, Flow. Vines 338 &
339. 1970; Schnack & Rubens, Bol. Soc. Argent. Bot. 13: 205 & 206.
1970; Angely, Fl. Anal. Fitogeogr. Est. S. Paulo, ed. 1, ): 839,
map 1393. 1970; G. Taylor, Ind. Kew. Suppl. 1h: 63. 1970; Molden-
ke, Fifth Summ. 1: 30, 177, 18h, 188, 190, 201, & 371 (1971) and
ae "521, 652, 657, 662; 663, 670, 677, 683, 700, 703, 783, & 915.
1971; Moldenke, Phytologia 22: h71 &’h91 (1972) ana’ 23: 239. 1972.

Recent collectors have found this plant growing in high fertile
campos. According to Religa, it is found abundantly in sunny,
sandy, semitropical areas in Paraguay. Troncoso (1965) gives its
overall distribution as "Brasil austral, Uruguay y litoral argen-
tino. Barrancas del Parand" and cites Boelcke 1968, Burkart

1276, and Nicora 351 from Buenos Aires, Argentina, deposited in

the San Isidro herbarium. Schnack & Rubens (1970) record it from
Buenos Aires, Corrientes, Entre Rfos, and Santa Fé provinces.
Krapovickas is of the opinion that the natural hybrid between
this species and V. temuisecta Briq. is V. calliantha Briq.,
which he says that % he found growing very sparingly among large
numbers of the two supposed parental species and which he says
fails to produce fertile seed. I have been regarding xV. trini-
tensis Moldenke as the natural hybrid between these two species.
The corollas of V. incisa are described as having been "red"
on Aguilar 55, Herrera 322, T. Rojas 322, and Terribile 871 &
882. Material has been misidentified and distributed in some
herbaria under the name Glandularia peruviana (L.) Small.
Additional citations: BRAZIL: Rio Grande do Sul: H. M. Filho
353 (W--283827). PARAGUAY: Religa s.n. [June 30, 1966] (Ws, Ws, Ws);
Bas Rojas 358a (Ws); Woolston 731 (N), » 1353 (N). ARGENTINA: Bue-
nos Aires: Fabris 4958 (N). Catamarca: Rodriguez V. 907 (Se—
130299). Chaco: Aguilar 55 (S), 1056 (N), 1316 (N); Buratovich
72 (N), 167 (N); Malv4rez 1 4155 (Ny; T T. Rojas 2111 [Herb. Inst.
Miguel Lillo 107893] (N). Cérdoba: Terribile 571 (N), 882 (N).
Corrientes: Malvarez 152 (N). Formosa: Montes Ss 2816 (Ny; I. Mo-
rel 2652 (N), 269) (N), 2! 2896 (N), 3113 (N), 352 3521 (N) (N), 3566 (. (N),
360 (N), 3624 (N), 3671 (N), 3705 (N), 3825 (N), 384u (N), 3883
(ny), 412 (Ny), 4677 (N),, 1,869 (Ny), 5149 (XN), 5317 (N),, 56146 Q),
5751 @), 5853 QD, 5925 Q), 6136 (%), 6319 (i), 6433 (), s son.
T22-x-19),8) (N (N); Pierotti 4123 (Se (Se--130298), 652 (N). M;siones:
Montes 11662 (Ac, N, Rf); G. G. Je Schwarz 4717 (N). Santa Fé: Al-
varez 987 87 (N). Caen: Herrera 322 (N)j O Olea 278 (N); Rocha
3683 (S).

VERBENA INCISA f. ALBIFLORA Osten & Moldenke
Additional bibliography: Moldenke, Phytologia 10: 112. 196k;

280 P BYE? oO LOG Vol. 23, udne
Moldenke, Fifth Summ, 1: 201 (1971) and 2: 677 & 915. 1971.

xVERBENA INHONESTA Moldenke
Additional bibliography: Moldenke, Phytologia 11: 68. 1965;
Moldenke, Fifth Summ, 1: 371 (1971) and 2: 679, 705, & 915. 1971.

VERBENA INTEGRIFOLIA Sessé & Moc.

Additional & emended bibliography: Lewis & Oliv., Am. Journ.
Bot. 8: [639]. 1961; Hocking, Excerpt. Bot. A.6: 91. 1963; Mol-
denke, Biol. Abstr. ho: 4697 (1968) and 49.(10): B.A.S.I.C. S.71
& S.18). 1968; Moldenke, Phytologia 16: 188. 1968; Bolkh., Grif,
Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 717. 1969; Moldenke,
Fifth Summ. 1: 76 (1971) and 2: 645, 678, & 915. 1971.

VERBENA INTEGRIFOLIA f. ALBIFLORA Moldenke

Bibliography: Moldenke, Résumé Suppl. 15: 3. 1967; Moldenke,
Biol. Abstr. 9: 4697. 1968; Anon., Biol. Abstr. 9 (10): B.A.S.I.
C. S.18). 1968; Moldenke, Phytologia 16: 95. 1968; Moldenke,
Fifth Summ. 1: 76 (1971) and 2: 915. 1971.

XVERBENA INTERCEDENS Briq.

Additional bibliography: Moldenke, Phytologia 1: 286. 1967;
Moldenke, Fifth Summ. 1: 177, 188, 190, 201, & 371 (1971) and 2:
655, 688, & 915. 1971.

Additional citations: PARAGUAY: Woolston 785 (Go, N).

VERBENA INTERMEDIA Gill. & Hook.

Additional & emended bibliography: Reiche & Phil., Fl. Chil.
5: 295. 1910; Cabrera, Man. Fl. Alred. Buenos Aires 295 & 396,
fig. 148 a--e. 1953; Darlington & Wylie, Chrom. Atl., pr. 1, 323.
1956; Schnack, Fehleisen, & Cocucci, Revist. Fac. Agron. La Plata
35: [lu7] & ce, fig. 3. 1959; Darlington & Wylie, Chrom. Atl.,
pr. 2, 323. 1961; Martinez—Crovetto, Bonplandia 1: 19 & 203.
1963; Troncoso in Cabrera, Fl. Prov. Buenos Aires 5: 127-130,
fig. 45 E—G. 1965; Huynh, Denkschr. Schweiz. Naturforsch. Gesel.
[Mém. Soc. Helv. Sci. Nat.] 85: 100. 1965; Martinez-Crovetto,
Bonplandia 2: 39, 52, 53, 59, & 70 (1965) and 2: 131. 1967; Mol-
denke, Résumé Suppl. 16: 28. 1968; Moldenke, Phytologia 16: 95 &
18). 1968; Bolkh., Grif, Matvej., & Zakhar., Chrom. Numb. Flow.
Pl. 717. 1969; Moldenke, Fifth Sum. 1: 177, 190, 201, & 371
(1971) and 2: 654, 655, 661, 663, 664, 672, 678, 693, 699, 700,
& 915--916. 1971; Moldenke, Phytologia 22: 47h. 1972.

Additional illustrations: Cabrera, Man. Fl. Alred. Buenos
Aires 396, fig. 148 a—e. 1953; Schnack, Fehleisen, & Cocucci,
Revist. Fac. Agron. La Plata 35: [54], fig. 3. 1959; Troncoso in
Cabrera, Fl. Prov. Buenos Aires 130, fig. 45 E—G. 1965.

Recent collectors have found this plant growing on campos and
report the vernacular name "verbena". It has been collected in
fruit in April and December (in addition to the months previous-
ly reported by me). The corollas are described as "violet" on
Burkart 19557, "blue" on Montes 1860 and A. G. Schulz 582, and

"lilac" on Krapovickas, Cristébal, Marufiak, Pire, & Tressens

1972 foldenke, Notes on Verbena 281

15319. Troncoso (1965) gives its overall distribution as "Sur del
Brasil, Uruguay, NE. y centro de la poesia Abunda en la pra-
dera primitiva, al borde de arroyos, vias férreas y en los valles
de las sierras del S. de la Provincia [Buenos Aires]". She cites
Cabrera 5630 and Hicken 475 & s.n. [Herb. San Isidro 3438] from
Buenos Aires. Schnack and his associates (1959) report that this
plant is apomictic in its natural form of reproduction.
Additional citations: URUGUAY: L. He Bailey B.765 B.765 (Se—-113022) ;
Rosengurtt Gurvich B.765 (W--2562150); Rosengurtt Gurvich & Gal-
linal 6018 (Se--1293hh) « ARGENTINA: Corrientes: Burkart 19557
(W--256797h) ; Krapovickas & Cristébal 16302 (Rf); Tressens & s & Al-
bizzatri 2 (Rf). La Pampa: Fortuna 13 3 (N), 30 (N); A. Ge Schulz
5842 (N). ~ Mendoza: Lourteig 852 (N) (N). ~~ Misiones: Krapovickas,
Cristébal, Marufiak, Pire, & Tressens 15319 (Rf); J. E. Montes
1,866 11,866 (Au-270806, N, |, RE, W--2556116). San Luis: Varela 692 (N).

VERBENA INTERMEDIA f. ALBIFLORA Moldenke
Additional bibliography: Moldenke, Phytologia 9: 38h. 1963;
Moldenke, Fifth Summ. 1: 201 (1971) and 2: 916. 1971.

VERBENA INTERMEDIA f. GLABRESCENS Hauman-Merck
Additional bibliography: Moldenke, Phytologia 9: 38). 1963;
Moldenke, Fifth Sum, 1: 201 (1971) and 2: 916. 1971.

VERBENA INTERMEDIA var. LANUGINOSA Moldenke
Additional bibliography: Moldenke, Phytologia 9: 38)—385.
1963; Moldenke, Fifth Sum, 1: 201 (ism and 2: 916. 1971.

VERBENA JORDANENSIS Moldenke

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 572.
1965; Moldenke, Phytologia 16: “ee 1968; Angely, Fl. Anal. Fito-
geogr. Est. S. *Paulo, ed. 1, h: 839 & xix, map 1393. 1970; Mol-
denke, Fifth Sum. 1: 177 (1971) and 2: 692 & 916. 1971.

Hatschbach has collected this species on rocky grassy campos.
The corollas are described as "violet" on Hatschbach 22833,
"lilac" on Hatschbach 22419, and "dark lilac" on Hatschbach 11883.

Additional citations: BRAZIL: Paran4: Hatschbach 14883 (Weaaks
2564554), 22833 (N, Rf), 33419 (Ac). Santa Catarina: Smith &
Reitz 1279 (N). ’

XVERBENA KONDAI Moldenke
Additional bibliography: Moldenke, Phytologia 11: 68. 1965;
Moldenke, Fifth Summ. 1: 371 (1971) and 2: 692, 700, & 916. 1971.

VERBENA KUNTZEANA Moldenke

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 572.
1965; Moldenke, Phytologia 11: 68. 1965; Moldenke, Fifth Sum.
1: 177, 188, a 201 (1971) and 2: 683 & 916. 1971.

282 PB) HNS, Ts (0) 5 ONG, aoa Vol. 23, no. 3

VERBENA LACINIATA (L.) Briq.

Additional synonymy: Verbena erinoideas Lam. apud Baez, Anal.
Asoc. Estud. Mus. Pop. Parand 1920: 37, sphalm. 1920. Verbena
lacinata (L.) Briq. ex Moldenke, Résumé Suppl. 3: 39, in syn.
1962. Verbena laciniata (Lam.) Briq. ex Moldenke, Résumé Suppl.
3: 39, in syn. 1962. Veraena erinoides Lam. ex Tawada, Biol. Mag.
Okinawa ) (6): 36, sphalm. 1967. Verbena laciniata (H.B.K.) Brigq.
ex Moldenke, Résumé Suppl. 16: 28, in syn. 1968. Erimus lacinia-
tus L. apud Angely, Fl. Anal. Fitogeogr. Est. S. Paulo, ed. 1, h:
839 & vii, sphalm. 1970.

Additional & emended bibliography: Balbis, Cat. Stirp. Hort.
Acad. Taur. 80. 1813; Pers., Sp. Pl. 3: 346. 1819; Steud., Nom.
Bot. Phan., ed. 1, 873. 1821; Jan, Elench. Pl. 1. 1831; A. Gray,
Field For. & Gard. Bot., ed. 1, pr. 1, 22 (1868) and ed. 1, pr.
2, 242. 1869; A. Wood, Am. Bot. & Flor., ed. 1, pr. 1, 235 (1870),
ed. 1, pr. 2, 235 (1871), ed. 1, pr. 3, 235 (1872), ed. 1, pr. h,
235 (1873), ed. 1, pra\5). 235 (187), and.ed..1, pro 16) 2950
1875; A. Gray, Field For. & Gard. Bot., ed. 1, pr. 3, 242. 1880;
O. R. Willis in A. Wood, Am. Bot. & Flor., ed. 2, 235. 1889; L.

H. Bailey in A. Gray, Field For. & Gard. Bot., ed. 2, 31. 1895;
Briq. in Chod. & Wilczek, Bull. Herb. Boiss., sér. 2, 2: 5h.
1902; T. Peckolt, Bericht. Deutsch. Pharm. Gesell. 1h: 166. 190);
Reiche & Phil., Fl. Chil. 5: 287, 289, & 29--295. 1910; Baez,
Anal. Asoc. Estud. Mus. Pop. Parand 1920: 37. 1920; Noack, Biol.
Zentralbl. 57: [383], 38), & 387, fig. 17. 1937; Baez, hius. Entre
Rios Cart. Herb. Paran. 3. 1938; Fischer & Harshberger, Flower
Fam. Alb. 86. 191; J. A. Clark, Card Ind. Gen. Sp. Var. issue
183. 1944; E. L. D. Seymour, New Gard. Encycl., ed. 3, 1279
(19h), ed. h, 1279 (1946), and ed. 5, 1279. 1951; Kuck & Tongg,
Mod. Trop. Gard. 213. 1955; Darlington & Wylie, Chrom. Atl., pr.
1, 323. 1956; Schnack, Fehleisen, & Cocucci, Revist. Argent.
Agron. 2h: 132, 13h, & 135. 1957; J. F. Macbr., Field Mus. Publ.
Bot. 13 (5): 612, 61h, 619, 623-626, & 629. 1960; Darlington &
Wylie, Chrom. Atl., pr. 2, 323. 1961; Harler, Gard. Plains, ed.

h, 23, 2h, 29, 238, & 250. 1962; E. L. D. Seymour, Wise Gard. En-
cycl., ed. 6, 1279. 1963; Troncoso in Bocher, Hjerting, & Rahn,
Dansk Bot. Arkiv 22 (1): 109. 1963; E. L. D. Seymour, New Gard.
Encycl., ed. 6, 1279 (1963) and ed. 7, 1279. 1964; R. Good, Geogr.
Flow. Pl. 218. 1964; Backer & Bakh., Fl. Jav. 2: 596. 1965; J. &
L. Bush-Brown, Am. Gard. Book, ed. , 327. 1965; Solbrig, Castanea
30: 173. 1965; Burkill, Dict. Econ. Prod. Malay Penins. 2: 2266.
1966; Greensill, Trop. Gardening 79. 1966; Hirata, Host Range &
Geogr. Distrib. Powd. Mild. 277. 1966; E. Lawrence, South. Gard.,
ed. 2, 115, 135, 172, & 21). 1967; Tawada, Biol. Mag. Okinawa
[Okinawa Seibutsugakkai] (6): 36--37. 1967; Moldenke, Phytologia
16: 188 & 207. 1968; Moldenke, Résumé Suppl. 16: 28. 1968; Sol-
brig, Passani, & Glass, Am. Journ. Bot. 55: 1239. 1968; Bolkh.,
Grif, Matvej., & Zakhar., Chrom, Numb. Flow. Pl. 715--717. 1969;
R. F. Ve Cooper in Pastore, Bol. Soc. Argent. Hort. 157: 123--125.
1969; Angely, Fl. Anal. Fitogeogr. Est. S. Paulo, ed. 1, h: 839,

1972 Moldenke, Notes on Verbena 283

vii, & xix, map 1393. 1970; E. L. D. Seymour, New Gard. Encycl.,
ed. 8, 1279. 1970; Moldenke, Fifth Sum, 1: 137, 143, 178 ver
190, 193, 201, & 371 (1971) and 2: 492, 521, 522, 569, 66h, 66
668, 671, 678, 684, 686, 691, 696, & 916. 1971; Moldenke, Rea
logia eat 222, 223, & 231. 1972

Additional illustrations: Noack, Biol. Zentralbl. 57: 387, fig.
L7. 1937.

Sparre found this plant growing on dry slopes in Ecuador,
while Fosberg refers to it as "prostrate, common in grassy places
on top of low hill grazed by goats". The corollas are described
as having been "violet" on Angulo & Lépez ae, "pale-violet" on
Asplund 20147, "lavender" on mn Fosberg 2765, and "light to rather
dark reddish-violet" on Asplund nd 20463, whi while Lourteig says "flor
amarilla violacea muy aromAtica".

Macbride (1960) does not attempt to separate V. tenuisecta
Briq., V. dissecta Willd., and V. laciniata (L. ) Briq. in his
key, lumping them all together in the line reading "Leaves mostly
or all larger" [than 1 cm.]. Actually, a casual examination of
the calyx or fruiting-calyx of each of these taxa under a hand-
lens reveals quite distinct pubescence characters for separating
them with comparative ease. He comments about V. laciniata:

"Type from Peru by Feuillée. Stems wiry, prostrate, forming
loose mats to a meter or more wide, the odorless, clear, mauve to
violet flowers in flattened heads to 5 cm. across (Balls); as in-
terpreted, leaves trifid or much divided, the Linnaean plant was
said to have scarlet flowers (Hooker); red-violet (Schauer) ." He
cites the following specimens from Peru: Cuzco: Herrera 18, 32,
& 3450, Hicken s.n. [Sicuani], Pennell 13677, Vargas 2k. "uaa
co: : Macbride ride 1260. La Libertad: West 8115. Lima: Ferreyra 6502,
Killip & Smi & Smith 21754, Macbride 678, BR Raimondi 10703. Briquet (1902)
cites Wilczek ezek hh, while Troncoso © (1963) cites Hicken 65.

Hirata (1966) records the fungus, Oidium verbenae, as infes-
ting V. laciniata in Java, but it is not at all certain that the
plant ci cultivated in Java aod described by Backer & Bakhuizen van
den Brink (1965) is really this species. I suspect that it is
really V. tenuisecta Briq., as I suspect are also the plants re-
ferred to as V. erinoides by Lawrence (1967) and by Tawada (1967),
while that discussed by Harler (1962) as "white verbena" and
"moss verbena" is almost certainly V. tenuisecta var. alba Mol-
denke. It should be noted, in passing, that Tawada's paper is
sometimes cited as "1968", but the first page of the issue is
plainly dated December 25, 1967, as is also the first page of
Tawada's article. ile records the vernacular name "karakusa-
baabena", but, again, it is most probable that this applies to
V. temiisecta rather than to the true V. laciniata.

Peckolt (190) records a plant which he calls V. erinoides
Lam. as being cultivated in Rio de Janeiro under the name "cha
do Brasil", which he translates as "brasilianischer Tee". He de-
scribes the plant as a "Niederliegende Pflanze mit an der Basis

28h P Hy 0 LyOrG sna Vol. 23, no. 3

keilformigen, in den Blattstiel verlaufenden, dreiteilig-fieder-
spaltigen Blattern. Bluten lilafarben. Die Infusion gilt beim
Volke als menstuations—- und lochienbeforderndes Mittel. Schwach
gerostet, wird sie von den Familien der Arbeiter als Ersatz des
indischen Tees benutzt." It seems most probable that this, also,
as with most of the many other references in literature to "V.
erinoides Lam.", refers to V. tenuisecta Briq., rather than to
V. laciniata.

The Pfister 2177, distributed as V. laciniata, is actually V.
berterii (Meisn.) S Schau., while Soukup 4960 is v. occulta Molden-
ke, Troncoso 353 is V. santiaguensis nsis (Covas & Schnack) Moldenke,
and © - Ow Schallert 2 251 is V. tenuisecta Briq.

heaton citations: ECUADOR: Chimborazo: Asplund 20163 (N);
F. R. Fosberg 27645 (Rf). Cotopaxi: Sparre 15690 (S). Tungurag-
ua: Asplund 201),7 (N). PERU: Arequipa: Vargas Calderén 19434
(Ac). La Libertad: Angulo & Lépez 1346 (Ac). Lima: Lépez Guil-
16n 670 (Rf); Riccio & Chumpitdz 3718 (Rf). ARGENTINA: Mendoza:
Lourteig 772 (N).

VERBENA LACINIATA var. CONTRACTA (Lindl.) Moldenke

Additional bibliography: Moldenke, Phytologia 11: 69. 1965;
Moldenke, Fifth Summ. 1: 193, 201, & 371 (1971) and 2: 667, 668,
68h, & 916. 1971.

VERBENA LACINIATA var. SABINI (Sweet) Moldenke

Additional bibliography: Moldenke, Phytologia 11: 469. 1965;
Moldenke, Fifth Summ. 1: 371 (1971) and 2: 667, 668, 678, 68h,
694, & 916. 1971.

VERBENA LAEVIS Salmon ex Hirata, Host Range & Geogr. Distrib.
Powd. Mild. 277, nom. nud. 1966.
Bibliography: Hirata, Host pane & Geogr. Distrib. Powd. Mild.
277. 1966; Moldenke, Fifth Sum, 1: 375 (1971) and 2: 916. 1971.
Nothing is known to me of this ae except that Hirata (1966)
states that it is attacked by the ubiquitous fungus, Erysiphe
cichoracearum P. DC., giving "Salmon" as authority.

VERBENA LANDBECKI R. A. Phil.

Additional & emended bibliography: Reiche & Phil., Fl. Chil.
5: 285 & 287. 1910; lioldenke, Phytologia 13: 251. 1966; Moldenke,
Fifth Sum, 1: 193 (1971) and 2: 679 & 916. 1971.

VERBENA LASIOSTACHYS Link

Additional & emended bibliography: Steud., Nom. Bot. Phan., ed.
1, 873 & 874. 1821; Cheymol, Bull. Soc. Chim. Biol. 19: 167--
1653. 1937; Noack, ’Biol. Zentralbl. 57: 36h & 306, fig.-2.d9ans
Anon., Chem. Abstr. 32: 2977. 1938; Howell, Marin Fl., ed. 1,
232--233. 1949; Abrams, Illustr. Fl. Pacif. States, pr. 1, 3: 610,
611, & 616, fig. 43h. 1951; Rattenbury, Madrofio 15: 51. 1959;
Ferris in Abrams & Ferris, Illustr. Fl. Pacif. States, pr. 1, :

1972 Moldenke, Notes on Verbena 285

651 & 730. 1960; J. H. Thomas, Fl. Santa Cruz Mtns., pr. 1, 29h &
43h, fig. 192. 1961; Pusateri, Fl. Sierra Natl. Parks 104 & 105,
fig. 138. 1963; Ferris in Abrams & Ferris, Illustr. Fl. Pacif.
States, pr. 2, 4: 651 & 730. 1965; Hocking, Excerpt. Bot. A.9:
36. 1965; Raven, Kyhos, & Hill, Aliso 6: 113. 1965; Abrams, Il-
lustr. Fl. Pacif. States, pr. 2, 3: 610, 611, & 616, fig. 43h.
1967; Boivin, Naturaliste Canad. 9): 642. 1967; Ornduff, Reg. Veg.
50: 86 & 123. 1967; Boughey, Mus. Syst. Bio. Univ. Calif. Irvine
Res. Ser. 1: 82. 1968; J. T. Howell, Calif. Bot. Club Trans-
Sierran Phyt. Exped. 1968 p. 14. 1968; Moldenke, Phytologia 16:
188 & 200. 1968; Moldenke, Résumé Suppl. 16: 1 & 28. 1968; Munz,
Suppl. Calif. Fl. 101. 1968; Munz & Keck, Calif. Fl. 686—688 &
1679. 1968; J. H. Thomas, Fl. Santa Cruz Mtns., pr. 2, 29h & 43h,
fig. 192. 1968; Bolkh., Grif, Matvej., & Zakhar., Chrom. Numb.
Flow. Pl. 717. 1969; Farnsworth, Blomster, Quimby, & Schermerh.,
Lynn Index 6: 267. 1969; H. L. Mason, Fl. Marshes Calif., pr. 2,
677 & 877. 1969; J. T. Howell, Marin Fl., ed. 2, 232--233. 1970;
Moldenke, Fifth Summ. 1: 18, 64-66, 76, & 371 (1971) and 2: 672,
67h, 679, 680, 691, 693, 705, & 916. 1971; Rickett, Wild Fls. U.
S. 5 (2): [455], 456, & 665, pl. 152. 1971; Moldenke, Phytologia
22: 458 & 467 (1972) and 23: 226 & 228. 1972.

Additional illustrations: Noack, Biol. Zentralbl. 57: 386, fig.
2. 1937; Abrams, Illustr. Fl. Pacif. States, pr. 1, 3: 616, fig.
4344. 1951; J. H. Thomas, Fl. Santa Cruz Mtns., pr. 1, 29, fig.
192. 1961; Pusateri, Fl. Sierra Natl. Parks 104, fig. 138. 1963;
Abrams, Illustr. Fl. Pacif. States, pr. 2, 3: 616, fig. 43h.
1967; J. H. Thomas, Fl. Santa Cruz Mtns., pr. 2, 294, fig. 192.
1968; Rickett, Wild Fls. U. S. 5 (2): [455], pl. 152 [in color].
1971.

The Madrofio (1959) reference cited above is sometimes credit-
ed to Solbrig, but appears to be part of a series begun by Rat-
tenbury and not otherwise accredited since.

Recent collectors have found V. lasiostachys growing on open
hills and on the banks of streams. Howell (199) says that it is
rather rare in clay soil of open grassy places or the edge of
brush in Marin County, California. Boughey (1968) refers to it as
"occasional" in wet places in Orange County, while Thomas (1968)
reports it from dry ground of disturbed areas, creek-bottoms,
roadsides, and along the edge of brushy vegetation in the Santa
Cruz Kountains, blooming there from May to September. He separ-
ates it from V. robusta Greene as follows:

Vv. lasiostachys — "Adaxial leaf-surfaces not scabrous; spikes
loosely flowered in fruit."
V. robusta -~ "Adaxial leaf-surfaces scabrous; spikes densely

~ ~~¥fTowered in fruit."

A. R. Moldenke 3396 is a very abnormal collection with foliace-
ous bracts in the young flower-spikes; according to the collector
it appeared in the Stanford University greenhouse after insecti-
cide sprays had been used therein -—- the plants were normal before
the spraying and shoots which developed after the spraying had been
completed were also normal.

286 PY EO 2 OG rs Vol. 23, no. 3

Raven, Kyhos, & Hill (1965) report the chromosome number for
V. lasiostachys as "2n = 7__" based on Raven 75411 from Los An-
geles County, California, and deposited in the herbarium of the
University of California at Los Angeles. Munz (1968) quotes this
as "2n = 7 pairs". Verbenaloside is reported from this species by
Cheymol (1937). Boughey (1968) cites Booth 1171 from Orange Coun-
ty, California.

Boivin (1967) is of the opinion that Née 111, preserved in the
Madrid herbarium and labeled as having been collected on Nootka
Island, is actually mis-labeled and did not originate on that is-
land, where, he maintains, this species of vervain is unknown.
This assumption is very possible, since numerous others of Née's
collections have been shown to be accompanied by erroneous lo-
cality data.

My son, Dr. Andrew R. Moldenke, reports that during the past
several years! field work in California he has observed the fol-
lowing insects visiting the flowers of Verbena lasiostachys:
Ashmeadiella cactorum basalis, A. californica, Bombus vosnesen-
skii, Ceratina acantha, C. a a Cc. aaa Cc. pacifica, C.
sequoiae, Chelostoma Socks rerae Gheladtonopets rubifloris,
Epeolus americanus, Eulonchus =the Geron sp., feviades
occidentalis, Hesperapis regularis, Hippodamia convergens, Hop-
litis producta gracilis, Hylaeus cressoni, Lepidanthrax lauta,
Megachile brevis onobrychia, M. gentilis, Melissodes lupina,
Phycioides campestris, and Polites sabuleti. Of these, Ashmead-
iella, Chelostoma, Chelostomopsis, Heriades, Hoplitis, and Mega-
chile are ecserinia bees, Bombus is a aie eesee Ceratina is a
carpenter bee, Epeolus ae Melissodes are dushasiosta bees,
Eulonchus is a cyrtid fly, Geron and Lepidanthrax are ponbyliid
flies, Hesperapis is a melittid bee, Hippodamia is a lady beetle,
Hylaeus is a colletid bee, and Phycioides and Polites are butter-
flies.

The McCulloch 2063c and Purer 538, distributed as V. lasio-
stachys, are actually — Ve abramsi Moldenke, Gonzdlez Quintero 2868
is in part V. elegans H.B.K. and in part v. elegans var. asperata
Perry, Cusick 1812, Getty 1, Hoover 9570, R. E. Nelson 158, Purer
5698 & 6755, and J. P. Tracy 18090 are V. lasiostachys var. sep-
tentrionalis Moldenke, and Cleveland s.n. [Sweetwater Valley] is
V. robusta Greene.
~~ Additional citations: CALIFORNIA: Los Angeles Co.: S. F. Blake
451 [Herb. Blake 1,03] (Ld), 523 {Herb. Blake 1,78] (La), ~ 70h
[Herb. Blake 1665] (Ld); Herb. James s.n. [1879] (W--260671) «
Monterey Co.: L. W. Reinecke s Son. [September 5, 1937] (Se--
14709); Youngberg | s.n. wn. [duly 1938] (Se--13637h) « San Bernardino
Co.: Munz 12313 (Se—129685). San Diego Co.: Gander 971 (Sd—
288h)); 1 Tie Flaw Wiggins 261) (Sd--l8715, Se—186887). San Mateo
Cos: Cummings & McCallum um 3726 (Sd--38891). Ventura Co.: Purer

1972 Mioldenke, Notes on Verbena 287

6703 (Sd——38892). County undetermined: M. Armstrong s.n. (Mi).
CULTIVATED: California: A. R. Moldenke 3396 (Ac, Z).

VERBENA LASIOSTACHYS f. ALBIFLORA Moldenke
Additional bibliography: Moldenke, Phytologia 13: 251. 1966;
Moldenke, Fifth Summ, 1: 65 (1971) and 2: 916. 1971.

VERBENA LASIOSTACHYS var. SCABRIDA Moldenke

Additional bibliography: Abrams, Illustr. Fl. Pacif. States,
pr. 1, 3: 611. 1951; Ferris in Abrams & Ferris, Illustr. Fl.
Pacif. States, pr. 1, : 730 (1960) and pr. 2, 4: 730. 1965; A-
brams, Illustr. Fl. Pacif. States, pr. 2, 3: 611. 1967; Moldenke,
Phytologia 16: 96. 1968; Munz & Keck, Calif. Fl. 688 & 1679.
1968; Moldenke, Fifth Summ. 1: 6) & 65 (1971) and 2: 679 & 916.
1971; Moldenke, Phytologia 22: 59. 1972.

Abrams (1951) reduces this taxon to synonymy under V. robusta
Greene, but it seems more probable to me that it may represent a
natural hybrid with that species.

VERBENA LASIOSTACHYS var. SEPTENTRIONALIS Moldenke

Additional bibliography: Abrams, Illustr. Fl. Pacif. States,
pr. 1, 3: 611. 1951; Ferris in Abrams & Ferris, Illustr. Fl. Pac-
if. States, pr. 1, 4: 730 (1960) and pr. 2, \: 730. 1965; Abrams,
Illustr. Fl. Pacif. States, pr. 2, 3: 611. 1967; Moldenke, Phy-
tologia 16: 96, 97, 188, & 200. 1968; Munz & Keck, Calif. Fl. 688
& 1679. 1968; Moldenke, Fifth Summ, 1: 64, 66, & 371 (1971) and
ee ee 685, 686, 691, & 916. 1971; Moldenke, Phytologia 22:

- 1972.

Recent collectors describe this plant as 8—18 inches tall,
growing on hillsides or in dry open ground with Hypericum perfor-
atum and Aira caryophyllea. The corollas are described as having
been "purple" on Dennis & Dennis 2255. Andrew R. Moldenke and I
saw this plant growing and flowering in Temaha County, California,
in July, 1968. A duplicate of the Cusick 812 collection, cited
below, was identified previously and cited by me in an earlier in-
stallment of these notes as typical V. lasiostachys Link. It
should be re-examined,

Additional citations: OREGON: Jackson Co.: Cusick 812 (Se—
17070); Dennis & Dennis 2255 (N). Josephine Co.: Steward & Mat-
thews 7188 (Se--202)55). CALIFORNIA: Glenn Co.: R. E. Nelson 159
(Se—161066). Humboldt Co.: J. P. Tracy 18090 (Se—203765). Kern
Co.: Jerabek s.n. [Hockett Place, July 1945) (Sd—36713). Mendo-
cino Co.: J. P, Tracy 5059 (Se--203766). San Diego Co.: Getty 1
(Sd--5007); Purer 5698 (Sd—38889). San Luis Obispo Co.: Hoover
9570 (Au--297791). Santa Barbara Co.: Tucker 1983 (Se—157877).
Trinity Co.: Moldenke & Moldenke 2)8)8 (Ac, Rf). Ventura Co.:
Purer 6755 (Sd--38893) .

XVERBENA LECOCQI Moldenke
Additional bibliography: Moldenke, Phytologia 9: 73. 1964; Mol-

288 P RTS LroteT a Vol. 23, no. 3
denke, Fifth Sum, 1: 371 (1971) and 2: 67h, 679, & 916. 1971.

VERBENA LILACINA Greene

Additional bibliography: Moldenke, Phytologia 16: 97. 1968;
Moldenke, Fifth Summ. 1: 76 & 78 (1971) and 2: 672 & 916. 1971.

Recent collectors describe this plant as a small suffrutescent
perennial or a bush 2 feet tall, growing on salty flats or on the
open slopes of peaks, occasionally on arroyo banks in canyons.
Harbison found it to be "very abundant", but Moran saw only "a
few". It has been collected in anthesis in February and in June,
and in fruit in June (in addition to the months previously re-
ported by me). The corollas are described as "pinkish-lavender"
on Wiggins & Thomas 187 and "blue or light-blue" on C. H. Muller
1078). Material has been misidentified and distributed in some
herbaria under the name V. gooddingii var. gooddingii Briq.

Additional citations: MEXICO: Baja California: C. F. Harbison

sen. [October 18, 1956] (Sd--l62)5); R. V. Moran 8195 - (Sd——54.632) 5

Wiggins & Thomas 187 (W--2521929). MEXICAN OCEA: OCEANIC ISLANDS:
Cedros: Haines & Hale s.n. [9 March 1939] (Sd—-5253); R. V. Mo-
ran 10669 (Sd=-51,0)3) , 10698 (Sd—5)0h1), 15162 (Sd—-67635) 5 ce
H. H. Muller 1078) (Sd--“150h) «

VERBENA LILLOANA Moldenke

Additional bibliography: Moldenke, Phytologia 9: 7l—75.
1964; Moldenke, Fifth Summ. 1: 201 (1971) and 2: 916. 1971.

The corollas on Olea 66 are described as having been "blue",
The species has been collected in anthesis in July and in fruit
in May, July, and December (in addition to the months previously
reported by me).

Additional citations: ARGENTINA: Jujuy: Reales 617 (N), 632

(N). Tucum4n: Olea 66 (N); F. Ortiz s.n. [30/5/45] (N).

VERBENA LINDBERGI Moldenke
Additional bibliography: Moldenke, Phytologia 9: 75-76.
196; Moldenke, Fifth Summ, 1: 178 (i971) and 2: 916. 1971.

VERBENA LINDMANII Briq.
Additional bibliography: Angely, Fl. Anal. Paran., eds, Lo Si2e
1965; Moldenke, Phytologia 16: 97. 1968; Moldenke, Fifth Summ.
1: 178 & 201 (i971) and 2: 680 & 916. 1971.
The corollas are described as "violet" on Ge Je Schwarz 1078.
Additional citations: ARGENTINA: Misiones: WG. ate Schwarz ; 1078

(N).

VERBENA LIPOZYGIOIDES Walp.

Additional bibliography: Reiche & Phil., Fl. Chil. 5: 289 &
293. 1910; Moldenke, Phytologia 1): 287. 1967; Moldenke, Fifth
Summ. 1: 193 (1971) and 2: 680 & 916. 1971.

VERBENA LITORALIS H.B.K.
Additional & emended synonymy: Verbena litoralis Humb. & Bonpl.

1972 Moldenke, Notes on Verbena 289

ex Steud., Nom. Bot. Phan., ed. 1, 873. 1821. Verbena littoralis
H.B.K. apud Walp., Repert. Bot. Syst. 4: 20, in syn. 1815; Hook.
f., Trans. Linn. Soc. Lond. Bot. 20: 195. 1647. Verbena bonari-
ensis sensu H. Mann, sensu Hillebr., sensu Heller, sensu Forbes,
sensu Rechinger, sensu Frear, and sensu St. John & Hosaka apud
Degener, Fl. Hawaii. 315: Verbena: Litoralis, in syn. 1960 [not V.
bonariensis Dill., 1938, nor L., 1753, nor Rendle, 1904, nor
Schau., 1960, nor Vell., 1959]. Verbena nudiflora "Nutt. ex Turez.!
ex Moldenke, Résumé Suppl. 18: 1), in syn. 1969.

Additional & emended bibliography: Steud., Nom. Bot. Phan., ed.
1, 873. 1821; Speg., Anal. Soc. Ci. Argent. 9: 17h. 1880; Brig.
in Chod. & Wilczek, Bull. Herb. Boiss., sér. 2, 2: 543. 1902; T.
Peckolt, Bericht. Deutsch. Pharm. Gesell. 1): L66. 1904; Speg.,
Anal. Mus. Nac. Buenos Aires 19: 323. 1909; Reiche & Phil., Fl.
Chil. 5: 283. 1910; Sacc. & Trott, Syll. Fung. 21: 775. 1912;
Speg., Rev. Argent. Bot. 1: 95 & 102. 1925; H. S. Jacks., Mycolo-
gia 2h: 62. 1932; Cheymol, Bull. Soc. Chim. Biol. 19: 1671653.
1937; Noack, Biol. Zentralbl. 57: 38) & 386, fig. 10. 1937; Anon.,
Chem. Abstr. 32: 2977. 1938; Baez, Mus. Entre Rfos Cart. Herb.
Paran. 3. 1938; Abrams, Illustr. Fl. Pacif. States, pr. 1, 3:
610. 1951; Cabrera, Man. Fl. Alred. Buenos Aires 395 & 397. 1953;
Darlington & Wylie, Chrom. Atl., pr. 1, 323. 1956; Cuatrecasas,
Revist. Acad. Colomb. Cienc. 10: 259. 1958; Schnack, Fehleisen, &
Cocucci, Revist. Fac. Agron. La Plata 35: 9, [5h], & 55, fig. 3.
1959; J. F. Macbr., Field Mus. Publ. Bot. 13 (5): 613, 615, 617,
621, 62h, & 628. 1960; Ferris in Abrams & Ferris, Ijlustr. Fl.
Pacif. States, pr. 1, 4: 730. 1960; Darlington & Wylie, Chron.
Atl., pr. 2, 323. 1961; MartinezCrovetto, Bonplandia 1: 112 &
132. 1962; Troncoso in Bocher, Hjerting, & Rahn, Dansk Bot. Arkiv
22 (1): 109. 1963; Angely, Fl. Anal. Paran., ed. 1, 572. 1965;
Iuynh, Denkschr. Schweiz. Naturforsch. Gesel. [Mém. Soc. Helv.
Sci. Nat.] 85: 100. 1965; Ferris in Abrams & Ferris, Illustr. Fl.
Pacif. States, pr. 2, 4: 730. 1965; Troncoso in Cabrera, Fl. Prov.
Buenos Aires 5: 128 & 132. 1965; Hirata, Host Range & Geogr. Dis-
trib. Powd, Mild. 277. 1966; Meyer & Weyrauch, Inst. Mig. Lill.
Misc. 23: 64 & 123. 1966; Rzedowski & McVaugh, Contrib. Univ.
Mich. Herb. 9: 76 & 107. 1966; Abrams, Illustr. Fl. Pacif. States,
pr. 2, 3: 610. 1967; Mueller-Dombois & Lamoureux, Pacif. Sci. 21:
298. 1967; Ornduff, Reg. Veg. 50: 86. 1967; Alzate, Pl. Medic.,
oT. 26h—265. 1968; Pollak-Oltz, Anal. Anthropol. Gesell. Wien
98: 51--58. 1968; Kunkel, Willdenowia ): 351. 1968; Moldenke,
Phytologia 16: 87, 97—99, 101, 102, & 342. 1968; Moldenke, Résu-
mé Suppl. 16: , 6, 11, & 26 (1968) and 17: 3 & 7. 1968; Munz,
Suppl. Calif. Fl. 101. 1968; Munz & Keck, Calif. Fl. 686, 687, &
1679. 1968; W. T. Pope, Man. Wayside Pl. 192, pl. 110. 1968;
Bolkh., Grif, Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 717.
1969; Farnsworth, Blomster, Quimby, & Schermerh., Lynn Index 6:
267. 1969; H. L. Mason, Fl. Marshes Calif., pr. 2, 676 & 877.
1969; A. L. Moldenke, Phytologia 18: 126. 1969; Moldenke, Biol.
Abstr. 50: 18. 1969; Rickett, Wild Fls. U. S. 3 (2): 36h & [367],
pl. 111. 1969; Angely, Fl. Anal. Fitogeogr. Est. S. Paulo, ed. 1,

290 P) Hate T Oy Le: Gx Eek Vol. 23, nenw

: 839, 840, & xix, map 139). 1970; Demnis, Kew Bull. Addit. Ser.
33 272; 1970; Gibson, Fieldiana Bot. 2h (95: 230 & 232—-233. 1970;
Moldenke, Phytologia 20: 80. 1970; Moldenke in Correll & Johnston,
Man. Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:] 131h&
1318. 1970; Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex.
Res. Found. Bot. 6:] 1876 & 1877. 1970; Oberwinkler, Pterid. &
Sperm. Venez. 19, 20, & 78. 1970; Anon., Biol. Abstr. 52. (3)e) Bene
Sak Ca Seclis 1971; Farnsworth, Pharmacog. Titles 6 (9): xii &
title 1628. 1971; Moldenke, Biol. Abstr. 52: 1316. 1971; Molden-
ke, Excerpt. Bot. 18.A: Lye 1971; Moldenke in Wiggins & Porter,
Fl. Gala4p. Isls. 504--508, fig. 133. 1971; Moldenke, Fifth Sumn.
UE aire We ui - 6h, 66, 16,024 82), Bl--86, 89, $1, 120,)/228,
LOT 138, 14h, , 18h, 188, 190, 193, 19h, 201, 205° 257, 312,
349-351, 353, nan (1971) and 2: 558; 559, 649, 652, 65h, 655,
oe 67h, 679--682, 686, 688, 708, & 916. 1971; Rickett, Wild Fls.

oh? (2)'s" (ESS, 456, & 665, pl. 152. 1971; Wiggins & Porter,
rh roan Isls. 997. 1971; Moldenke, Phytologia 22: 78 & 89
(1972) and 23: 182, 185, 225, & 233. *1972.

Additional illustrations: Noack, Biol. Zentralbl. 57: 386,
fig. 10. 1937; Schnack, Fehleisen, & Cocucci, Revist. Fac. Agron.
La Plata 35: [5h], fig. 3. 1959; We T. Pope, Man. Wayside Pl.

192. pl." 110" fas. Vs bonariensis| . 1968; Moldenke in Wiggins &
Porter, Fl. Gal4p. Isls. [507], fig. 133. 1971.

Recent collectors describe this plant as an herb to 1.5 m.
tall and have found it growing in low-lying grasslands near riv-
ers, in fields, in limestone sinks with steep walls in oak woods,
in dense woodlands, cloud forests, open fields, and second-growth,
in potreros and cultivated areas, on valley floors, grassy valley
floors, wet land, and high ridges, along roadsides and wet road-
sides in the open sun, streamsides, and village fencerows, by
small ponds in Pinus—Liquidambar forests, at the edge of ditches,
and among scrubby growth on hilltops.

Pennington claims that V. litoralis is an "excellent pasturage
plant". Mears found it to be "fairly common in open sunlight" in
San Luis Potos{, Mexico, while the Andersons call it an "abundant
roadside weed" in Guerrero. Taylor describes it as a "weed at
edge of forest adjacent to old fields in volcanic very moist
gray-black soil" in Costa Rica and Tyson refers to it as "common"
in Chiriquf, Panama. Duke found it growing in areas receiving a-
bout 80 inches of annual rainfall. Mueller-Dombois & Lamoureux
(1967) state that it is found in two of the "kipukas" in Hawaii
Volcanoes National Park, these being older areas on the slopes
of volcanic mountains that have been surrounded by more recent
lava flows, serving as vegetation islands providing seed sources
for the re-invasion of the surrounding new volcanic material.

The corollas are described as having been "blue" on Anderson &
Anderson 928, Breedlove 1429, Contreras Sell, J. A. Duke 9072,
Ebinger SLs e Laughlin in 652, Lew. Lewis, Burch, Dwyer, Elias, Escobar,
Oliver, & Ljnsberieen 3 339, Willians, “Molina R., Williams, & Gibson

2936L, a Woodson, Allen, & Seibert 858, pale-blue" on . Breed—
love & Thorne 17993, "bluish" on Herb. Inst. Miguel Lillo 7 36,

1972 Moldenke, Notes on Verbena 291

"dark-blue" on P. H. Allon 1412, "bright lavender—blue" on Seibert
169, "lilac" on E. Contreras 3 6152, Molina R. 22258, and Molina R.
& & Molina 2265, “magenta” on Tyson, Dwyer, & Blum 1,300, "lavender"
on - Bunting & L & Licht 976, Croat - 10Lbh, R. McVaugh 16919, C. W. Pal-
mer 105, Tyson 5627 m 5627 & 5675, a and E. H. “Walker 8127, "pale-lavender"
on Camp 233 and E. H. Walker 8103, “Mp ink" on | Gentle 6481, "pink-
ish" on Gentle 7119, “"purple" on on Blum, Olson, & Rasmussen 2409,
F. R. Fosberg 27947 279L7, Gutiérrez R R. 58, and Wartinez—Calderén 1h71,
"light-purple" on on J. W We. Le Robinson n 8, "pale-purple" on Molina R R.,
Burger, & Wallenta 16234, and "pale-violet" on Asplund 18329.

Schnack and his associates (1959) report this species to be
apomictic in its natural method of reproduction, The vernacular
names, "cotacd4n", "mountain verbina", "verbena de montafia", and
"verbena fina", have been recorded for this species (in addition
to the names previously reported by me). Peckolt (190) records
the vernacular name "herva de pai S. Caetano", which he trans-
lates as "Cajetans Vaterkraut", He describes’ the plant as "Meter-
hohe Pflanze mit sitzenden, lanzlettlichen Blattern. Blititen blau
in kurzen, zylindrischen Khren. Wird allgemein als Ersatz der
Verbena officinalis L. benutzt." Judging from his brief descrip-
tion, it seems that he has misapplied the specific name to V.
brasiliensis Vell., which has sessile leaves. He is not alone in
making this misidentification! The "Verbena litoralis" of liason
(1969) is certainly also actually V. brasiliensis, and the illus-
tration given by Rickett (1969) and labeled as depicting V. bonar-
iensis and "V. littoralis" actually shows V. bonariensis (in the
foreground) and V. brasiliensis (in the rear).

Kunkel (1968) is of the opinion that the "V. hispida Ruiz &
Pav." of Reiche [Anal. Mus, Nac. Chile, 1903] is actually V. lit-
oralis -- he cites Kunkel R.70 from La Mocha Island, Chile. In
regard to New Zealand mat material, Healy comments that "this plant
appears to have been included under V. officinalis [in previous
literature], but is markedly different from southern [New Zea-
land] plants".

Alzate (1968) confirms Peckolt's statement that V. litoralis
can be used as a substitute for V. officinalis in South American
folk medicine. Cheymol (1937) reports the presence of verbenal-
oside in V. litoralis. Ornduff (1967) and Munz (1968) report
its chromosome number as 2n = 56. Cuatrecasas (1958) reports
the species from Valle del Cauca, Colombia, and Briquet (1902)
cites Wilczek 56 from southern Brazil.

A letter received by me from Julia F. Morton, dated June 2,
1969, is of interest: "The USDA Plant Quarantine people have en-
countered ([V. litoralis] three times in the past month in the
possession of travelers coming in from Guatemala. All were
raving about the wonderful effect of the 'tea' [made from this
plant] in lowering blood pressure. It was first brought to my

292 P HeYnOp Liv 0 "Gy tek Vol. 23, no. 3

attention by a Fort Lauderdale doctor who confirmed that her
patient's blood pressure went down after taking this remedy. It
coincided with the first time the USDA people saw a bundle of the
dried plant being brought in by a 70-year-old man,"

Troncoso (1965) gives the overall distribution of the species
as "América cAlida y templada. Comin en campos secos y altos,
invasore de cultivos", citing Cabrera 1710, Rodriguez 93; and.
Troncoso s.n. [Herb. Sais Isidro 193] from Buenos Aires, Argen-
tina. She reduces V. brasiliensis Vell. to synonymy andee Vv. lit-
oralis, but the two s species are abundantly distinct. Munz (1968)
comments under V. brasiliensis "Confused with V. litoralis HBK,
but the infl. more condensed, the pubescence of rachis, bracts
and calyx more spreading". fie then separates them as follows:

V. litoralis -- inflorescence lax, elongate; flowers distant;
~~ pubescence on rachis, bractlets, and calyx very minute,
closely appressed.

V. brasiliensis -- inflorescence dense, contracted; flowers most—
ly congested; pubescence on rachis, bractlets, and calyx
spreading.

Macbride (1960) notes that "Hooker and Schauer added a [second]
't! to the name; the former remarked [Hook. Bot. Misc. 1: 166.
1830]: seems to be a variety [of V. bonariensis] with shorter
spikes than usual" -- but here, again, the plant actually referred
to is V. brasiliensis. Macbride continues with the remarkable
statement that "Actually, most material referred here seems doubt—
fully distinct from V. carolina L.", but the Mexican and Central
American V. carolina really has very little resemblance to V.
litoralis. He cites the following Peruvian specimens of V. lit-
oralis: Amazonas: Williams 7534. Arequipa: Guenther & Buchtien
132. Cajamarca: Bonpland s.n. [Truxillo], Osgood & Anderson 36.
Cuzco: Herrera 1505, 166, & 3362. Hudnuco: Macbride 1705,
Woytkowski 34175. Junin: Schunke 172). Lima: Killip & Smith
21541, Macbride 5h. Loreto: Williams 1 1297, 4367, & 789). San
Martin: Williams 5911.

Macbride notes that Humboldt, Bonpland, & Kunth (1817) "gave

Truxillo, Santa and Lima specimens as type collections" -—- ac-=
tually cotype collections -- but he designates the Truxillo one
as the "type", which, under the present edition of the Internat-
ional Rules of Botanic Nomenclature, he is entitled to do. In
this connection it should be noted that the photograph cited be-
low of Nuttall s.n. [Wahoo] indicates that the type of V. mudi-
flora Nutt. is deposited in the British Museum herbarium.
Hitherto the poor specimen in the herbarium of the Academy of
Natural Sciences at Philadelphia has been regarded by the curators
there as the type, "too valuable" to lend even for examination by
a monographer.

Jackson (192) reports the fungus, Aecidium verbenae Speg., as

attacking V. litoralis. He regards A. verbenicola Speg. [not A.

1972 Moldenke, Notes on Verbena 293

verbenicola Ellis & Kellerman, 188], A. spegazzinianum Sacc. &
Trott., A. elongatum Speg., and A. verbeniphilum Speg. as synonyms
of A. verbenae, but admits that "There is some doubt whether or
not all the names listed above belong to one species. Spegazzini
evidently considered that there were at least two forms. His A.
Verbenae he thought to be the aecial stage of Puccinia elongata
Speg. The latter, however, seems, from the description, to be a
short cycled form." He cites He 7 Jackson 1180, 1272, & 1454
from Rio de Janeiro and 1380 & 1479 from Minas Gerais ais and S80
Paulo, Brazil. Hirata (1966) adds Oidium sp. from South America,
while Dennis (1970) adds Septoria verbenae Rob. & Desm.

Material of V. litoralis has been misidentified and distribu-
ted in some herbaria as V. carolina L., V. ehrenbergiana Schau.,
V. recta H.B.K., and V. townsendi Svenson. On the other hand,
the Nicora 663, distributed as V. litoralis, is actually V. bon
ariensis L., Cory 50840 is V. brasiliensis Vell., Sagdstegui inks
7192 is V. glabrata H.B Baka. "Rambo 45339 is V. minutiflora Briq.
{not Ke montevidensis as previously reported | by me], Nicora 377
and Troncoso 357 are V. montevidensis Spreng., Edwin & & Schunke ~
V. 3746 and Lépez—Palacios 2552 are V. parwila Hayek, "and R. V.
Moran an 5793 & 5817 are V. sphaerocarpa _ Perry. The Alvarez 951 a and
Archer 4,788, previously y cited by me as representing the typical
form of Verbena litoralis, seem, rather, to represent its var.
caracasana (H.B.K.) Briq.

Additional citations: MEXICO: Chiapas: Breedlove 129 (Mi, N);
Laughlin 652 (Mi). Chihuahua: Pennington 55 (Au—26]053). Guer-
rero: Anderson & Anderson 4928 (Mi). Hidalgo: Gonzd4lez Quintero
B2 (Au—-2),9)83, Ip). Jalisco: Diaz Luna 60 (Mi); R. McVaugh
16949 (N). México: Chavez s.n. [5-V-1963] (Ip); M. E. Diaz 18
(Ae); Mitastein 2) (Ip), 7 72 (Ip). Michoac4n: Hinton 12156 (Se--
97689), 12520 (Se--117:23) « Oaxaca: Camp 233 (N). Puebla:
Gutiérrez R. | R. 58 (Ac). San Luis Potos{: Mears ars 92a (Au). Sinaloa:
Breedlove & Thorne 17993 (Ac). Veracruz: Martinez Calderén 1471
(Mi, N); Rosas R. 512 (Rf). GUATEMALA: El Petén: E. Contreras
6152 (W—2558713); C. L. Lundell 16386 (N). El Quiché: E. Con
treras 52h (Au-—-27857h). Solol4: Kol Molina R., Burger, & Wallenta
16234 (N, N, N). Department undetermined: Collector undesignated
S.n. (Jm). BRITISH HONDURAS: Gentle 6481 (Au—239635, S), 7119
(Mi). HONDURAS: Distrito Central: Gillis 9065 (Ft, Ft, Go); Ne Nel-
son & Barkley 39458 (Ac). Intibucé: Bark Barkley & & Barkley 0349 (Ac).
La Paz: Molina R. & Molina 2265 (N). Ocotepeque: A. me: A. Molina R.
22258 (N). NICARAGUA: Carazo: F. C. Seymour 570 (vt). Matagal-
pa: pa: Bunting & Licht 976 (N); Zelaya | M. 232) (N). COSTA RICA:
Alajuela: Molina R. Ras “Williams, Burger, & & Wallenta 17513 (N). Car-
tago: R. J. Taylor 208 (N). Puntarenas: C. W. Palmer 105 (N).

294 P PASE PO "eV rs Vol. 23, no. 3

San José: Williams, Molina R., Williams, & Gibson 2938) (N).
PANAMA: Chiriqui: P. H. Allen 112 (E--1190850);_ Croat 10k (N);
J. A. Duke 9072 (E--18)179h); Dwyer & Hayden 7738 (E—1925953);
Ebinger r 611 ~(W--2561792); Lewis, Burch, Dwyer, Elias, Escobar,
Oliver, & & Robertson 359 (E—-168198h, W-25891,52) ; Seibert 169° (E—
111029); E. L. Tyson 5627 (E—1980019), 5675 (E--1980017) ; Wood-
son, Allen, & Seibert 858 (E—-1171088). Panamé: Blun, Olson, & &
Rasmussen ssen 2109 (E--1974177) ; Tyson, Dwyer, & Blum 1,300 (c=
1889209). COLOMBIA: Cundinamarca: Barkley & Wrigley 38838 (Rf);
Humbert, Fernandez, Idrobo, & Jaramillo 27159 (N, S). Valle del
Cauca: is Wists Robinson nson 6 (W--2563571) « Department undetermined:
Mutis 3688 (W--1562719). VENEZUELA: Mérida: Lépez—Palacios 2035
(Ft). ECUADOR: Chimborazo: Asplund 15501 (N). El Oro: Asplund
15820 (N). Guayas: Valverde L79 (Ws). Loja: D. H. Knight 781
(Ws). Los Rfos: Jativa & Epling 55 (N). Napo-Pastaza: za: Asplund
18329 (N). GALAPAGOS ISLANDS: Indefatigable: L. A. Fournier mier 97
(Rf). PERU: Arequipa: Vargas Calderén 8)62 (Ac), "18250 (Ac).
Cajamarca: Ugent & Ugent % 5512 (WW——2558166) . Cuzco: Var Vargas Cal-
derén 1,580 (Ac), 1 17076 (Ac). Lambayeque: F. R. Fosberg 279L7
(Rf). BRAZIL: Rio Grande do Sul: Rambo 5117 (Au--122320). BO-
LIVIA: Santa Cruz: I. Peredo s.n. [23-IV-1946] (Se--130300).
PARAGUAY: Woolston 107 (N). ARGENTINA: Buenos Aires: R. Alvarez
31) (N). Catamarca: Brizuela 696 (N); Luna Risso 1058 -(N). For-
mosa: I. Morel 2810 (N), 1369 (Ny), 4862 (N) (N), “4916 (N). Jujuy:
Garolera & Romero s.n. [1]-I-197] (N). Misiones: G. J. Schwarz
5383 (N). San Luis: Varela 739 (N). Santiago del Estero: P.
Garcia s.n. [Herb. Inst. Miguel Lillo 738] (N). RYUKYU ISLAND
ARCHIPELAGO: Okinawa: E. H. Walker 8103 (Ww), 8127 (Rf, W—
2619387). NEW ZEALAND: North: Healy 50/215 (Nz—70251), 50/218
(Nz--70250). HAWAIIAN ISLANDS: Oahu: Nuttall SMe (Wahoo; F. G.
Mey. photo 3129] (N--photo). AUSTRAL ISLANDS: Raivavae: Whitney
Exped. 258 (N). == rae

VERBENA LITORALIS var. ALBIFLORA Moldenke
Additional bibliography: Moldenke, Phytologia 13: 206. 1966;
Moldenke, Fifth Summ, 1: 76, 120, & ib (1971) and 2: 916. 1971.

VERBENA LITORALIS var. CARACASANA (H.B.K.) Briq.

Additional & emended synonymy: Verbena caracasana H.B.K., Nov.
Gen. & Sp. Pl., ed. folio, 2: 223. 1817. Verbena caracassana
Humb, & Bonpl. ex Steud., Nom. Bot. Phan., ed. 1, 872. 1821.

Additional & emended bibliography: Steud., Nom. Bot. Phan.,
ed. 1, 873. 1821; Barnhart, Bull. Torrey Bot. Club 29: 590.

1902; *Moldenke, Phytologia "16: 99. 1968; Moldenke, Résumé Suppl.
16: 7 & 28 (1968) and 17: 3 & 7. 1968; Moldenke, Fifth Summ. 1:
120, 128, 184, 188, 201, & 350 (1971) and 2: 660, 680, 916, &
967. 1971.

1972 Moldenke, Notes on Verbena 295

Recent: collectors describe this plant as an upright herb, 0.3—
1m, tall, the calyx green-ochre and the corolla blue (R. Fe
Steinbach "699) or white (Nevling & Gémez-Pompa 1033), growing at
altitudes of 700 to 2020 m. Steinbach describes it as "common'
in rocky soil along roadsides in Bolivia, while in New Zealand
it is said by Healy to be "established on roadsides and waste
land". The latter collector notes that "this plant appears to
have been included under V. officinalis {in previous lists of New
Zealand plants], but is markedly different from southern [New
Zealand] plants."

The R. Alvarez 951 and W. A. Archer 1,788, cited below, were
previously cited by m me as typical V. litoralis H.B Ke, bat they
certainly are not representative of f the typical form of that ad-
mittedly very variable taxon. They have the appearance of large-
spiked V. montevidensis Spreng. and are placed here in V. litor-
alis var, caracasana tentatively until the true status of this
and related taxa can be more fully investigated.

If the following collections actually do represent this vari-
ety, then it has been collected in anthesis and fruit in January,
February, April, June, and December, it is known in the vernacular
as "vervena", and is used as a general remedy for coughs in folk
medicine.

Additional & emended citations: MEXICO: Veracruz: Nevli
Gémez—Pompa 1033 (Ac). BOLIVIA: Cochabamba: R. F. Steinbach 259
(W—-2533500, Ws). PARAGUAY: W. A. Archer 4788 (N). ARGENTINA: |
Cérdoba: Rufz Huidobro s.n. (San Martin] (N); Timmermann 71123
(Au—-3024),9). Formosa: I. Morel 2188 (Rf). Jujuy: O'Donell
2914 (N). Santa Fé: R. Alvarez arez 951 (N). Tucum4n: T. Meyer 15123
(Au--122322). NEW ZEALAND: North: Healy 50/74 (Nz—70252,

Z), 50/215 (Nz—9382)) .

VERBENA LITORALIS var. CONGESTA Moldenke

Bibliography: Moldenke, Phytologia 20: 80. 1970; Moldenke,
Biol. Abstr. 52: 1316. 1971; Anon., Biol. Abstr. 52 (3): B.A.S.I.
C. S.2h7. 1971; Moldenke, Excerpt. Bot. 18.A: 45. 1971; Molden-
ke, Fifth Sum. i: 76 (1971) and 2: 916 & 973. 1971.

’ Breedlove & Thorne found this plant growing on steep moist
north-facing slopes with Clethra, Cornus, Molinadendron, Ostrya,
Pinus, and Quercus, flowering and fruiting in September, and de-
scribe the color of the corollas as "blue",

Citations: MEXICO: Sinaloa: Breedlove & Kawahara 16735 (Z—
type); Breedlove & Thorne 18277 (Mi).

VERBENA LITORALIS var. MELANOPOTAMICA Hauman-lierck
Additional bibliography: Moldenke, Phytologia 10: 78—79.
1964; Moldenke, Fifth Sum. 1: 201 (i971) and 2: 916 & 973. 1971.

VERBENA LOBATA Vell.
Additional bibliography: Barroso, Rodriguésia 32: [69] & 70.

296 P HAY.1.0. L0G" rs Vol... 23,, nos 3

1957; Cabrera, Bol. Soc, Argent. Bot. 6: 272. 1957; Angely, Fl.
Anal. Paran., ed. 1, 572. 1965; Moldenke, Phytologia 16: 99.
1968; Moldenke, Résumé Suppl. 16: 6. 1968; Angely, Fl. Anal. &
Fitogeogr. Est. S. Paulo, ed. 1, 1: x1i.(1969) and : 839, 80,
& xix, map 139. 1970; Moldenke in Menninger, Flow. Vines 338.
1970; Moldenke, Fifth Summ, 1: 178, 190, & 201 (1971) and 2:
657, 664, 681, 682, & 916. 1971.

Additional citations: BRAZIL: Rio Grande do Sul: Palacios &
Cuezzo 895 (N).

VERBENA LOBATA var. GLABRATA Moldenke
Additional bibliography: Moldenke, Phytologia 12: 207. 1966;
Moldenke, Fifth Summ, 1: 178 & 190 (1971) and 2: 916. 1971.

VERBENA LOBATA var. HIRSUTA Moldenke

Additional bibliography: Moldenke, Phytologia 16: 99. 1968;
Moldenke, Résumé Suppl. 16: 6. 1968; Moldenke, Fifth Summ. 1:
178 & 188 (1971) and 2: 916. 1971.

The corollas on Hatschbach 14898 are described as having been
"lilac" in color when fresh.

Additional citations: BRAZIL: ParanA: Hatschbach 14898 (W—
256,580) .

VERBENA LOBATA var. SESSILIS Moldenke

Additional bibliography: Cabrera, Bol. Soc. Argent. Bot. 6:
272. 1957; Angely, Fl. Anal. Paran., ed. 1, 572. 1965; Moldenke,
Phytologia 16: 99. 1968; Moldenke, Fifth Summ, 1: 178 (1971) and
Pace Keys tale iale

VERBENA LONGIFOLIA Mart. & Gal.

Additional bibliography: Hocking, Excerpt. Bot. A.11l: 50h.
1967; Moldenke, Phytologia 1): 288. 1967; Moldenke, Biol. Abstr.
49: 199. 1968; Moldenke, Fifth Summ, 1: 76 (1971) and 2: 660 &
916. 1971.

Dieterle found this plant growing in flat pastured red clay
land with shallow pools in an area of low ground with shallow
surface water, describing the color of the corollas as "dark-
purple". In addition to the months previously reported by me,
the species has been collected in flower and fruit in June,

—_— ———- ———-

VERBENA LONGIFOLIA f. ALBIFLORA Moldenke
Additional bibliography: Moldenke, Phytologia 13: 207. 1966;
Moldenke, Fifth Summ. 1: 76 (1971) and 2: 916. 1971.

VERBENA LONGIFOLIA var. PUBESCENS Moldenke

Additional bibliography: Hocking, Excerpt. Bot. A.11: 50h.
1967; Moldenke, Phytologia 1): 288. 1967; Moldenke, Biol. Abstr.
9: hag99. 1968; Moldenke, Fifth Summ. 1: 76 (1971) and 2: 916.
1971.

1972 Moldenke, Notes on Verbena 297

VERBENA LUCANENSIS Moldenke

Additional bibliography: J. F. Macbr., Field Mus. Publ. Bot.
13 (5): 613, 614, & 62h--625. 1960; Moldenke, Phytologia 1): 288.
1967; Moldenke, Fifth Summ. 1: 1) (1971) and 2: 916. 1971.

VERBENA LUDOVICIANA Hirata
Bibliography: Hirata, Host Range & Geogr. Distrib. Powd. Mild.
277. 1966; Moldenke, Fifth Summ. 1: 375 (1971) and 2: 916. 1971.
Nothing is known to me of this plant except that it is listed
by Hirata (1966) as a host for the ubiquitous fungus, Erysiphe
cichoracearum P. DC. in the United States.

VERBENA MACDOUGALII Heller

Additional & emended synonymy: Verbena macdougali Heller ex
Moldenke, Suppl. List Invalid Names 9, in syn. 191; Hirata, Host
Range & Geogr. Distrib. Powd. Mild. 277. 1966. Verbena macdougalli
Shinn, Univ. Kans, Sci. Bull. 6: 881. 1967.

Additional & emended bibliography: Rydb., Fi. Rocky Mtns., ed.
2, pr. 1, 739 & 740. 1922; Tidestr., Contrib. U. S. Nat. Herb. 25
[Fl. Utah & Nev.], pr. 1, 469. 1925; Wyman & Harris, Navajo Ind.
Ethnobot. (Univ. N. M. Bull. 366 (Anthrop. Ser. 3, 5):] 32 & 45.
1941; Rydb., Fl. Rocky Mtns., ed. 2, pr. 2, 739 & 740. 1954; How-
ell & McClintock in Kearney & Peebles, Ariz. Fl., ed. 2, 725--
727. 1960; Lewis & Oliv., Am. Journ. Bot. 48: [639]--641, fig. 17.
1961; Hocking, Excerpt. Bot. A.6: 91. 1963; Hirata, Host Range &
Geogr. Distrib. Powd. Mild. 277. 1966; Shinn, Univ. Kans. Sci.
Bull. 6: 881. 1967; Burlage, Ind. Pl. Tex. 18), 206, 223, & 22.
1968; Moldenke, Phytologia 16: 188--189. 1968; Bolkh., Grif,
Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 717. 1969; A. L. Mol-
denke, Phytologia 18: 126. 1969; Rydb., Fl. Rocky Mtns., ed. 2,
pr. 3, 739 & 740. 1969; Tidestr., Contrib. U. S. Nat. Herb. 25
[Fl. Utah & Nev.], pr. 2, 469. 1969; Rickett, Wild Fls. J. S. 3
(2): 365 (1969) and h (33: 5hO, [SiiL], & 799, pl. 176. 1970;
Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex. Res. Found.
Bot. 6:] 1876 & 1877. 1970; Moldenke in Correll & Johnston, Man.
Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:] 1315 & 1320.
1970; Moldenke, Fifth Sum. 1: 50, 51, 59, 62, 63, & 371 (1971)
and 2: 682 & 916. 1971.

Additional & emended illustrations: Lewis & Oliv., Am. Journ.
Bot. 48: 640, fig. 17. 1961; Rickett, Wild Fls. U. S. (3):
[541], pl. 176 [in color]. 1970.

Recent collectors have found this plant growing in forests and
open woodlands, in subalpine meadows, on rocky hillsides with
Pinus ponderosa and Abies concolor, and along subalpine roadsides,
at altitudes up to 10,000 feet. Howell & McClintock (1960) state
that in Arizona it is found mostly in pine forests from 6000 to
7000 feet altitude, blooming from June to September, while Dem-
aree reports it as "common in rocky open wooded bottoms" in New
Mexico. Andrew and Alison Moldenke have reported to me that "V.
macdougalii is just as common, if not more so, at the Grand Can-
yon and to/from Flagstaff [Arizona], but only in weedy areas. We

298 P HY TsO LOG Vol. 23, no. 3

haven't seen a single plant here in non-disturbed areas — none
at all in 500 miles of N. Arizona and S. Utah."

Common names for the species, in addition to those previously
reported by me, are "dormillon" and "Macdougal verbena". Burlage
(1968) reports that "a tea of this is drunk as a diuretic [in Tex-
as] and the green leaves are mashed and used for toothache". Hir-
ata (1966) records the fungus, Erysiphe cichoracearum P. DC., as
attacking this plant. The H. R. Bennett 8055, distributed as V.
macdougalii, is actually mut. rosella Cockerell, while Johnston | &
Muller 877 is V. scabra Vahl.

Additional citations: COLORADO: Conejos Co.: W. A. Weber 7865a
(Se--1)5178). NEW MEXICO: Colfax Co.: Mahler 861 (Au——2))8858) .
Lincoln Co,: W. Hess 260 (Se—-226055); Tucker 3209 (Se--18)025) ;
Wooton & Standley y 47hs2- (Ws). McKinley Co.: “Co.: Baad aad 1076 (Se--
236110). Otero Co.: +: Baad 798 (Se--236953); Demaree 5099 (Ac),
60788 (Rf); Iwen & Iwen en 202 2 (Ws). San Miguel Co.: | Co.: R. S. Ferris
11521 (Se--189506); Rowlett 2 (Au--25),348); Studhalter & Marr S.
1836 (Lk). Taos Co.: Kempers s oh (Sd--6),213). ARIZONA: Coconino ~
Co.: G. N. Jones 2100 . (Se——199522) .

VERBENA MACDOUGALII f. ALBIFLORA Moldenke
Additional bibliography: Moldenke, Phytologia 10: 117--119.
1964; Moldenke, Fifth Summ, 1: 51 & 62 (1971) and 2: 916. 1971.

VERBENA MACDOUGALII mut. ROSELLA Cockerell

Additional bibliography: Moldenke, Phytologia 10: 118. 196);
Moldenke, Fifth Summ, 1: 62 (1971) and 2: 916. 1971.

Bennett encountered this plant at 7300 feet altitude in the
Transition Zone, flowering in July. He describes the corolla
color as "rose". It has been misidentified and distributed in
some herbaria as typical V. macdougalii Heller,

Additional citations: NEW MEXICO: San biiguel Co.: H. R. Ben-
nett 8055 (Go, W—-2)6278) .

VERBENA MACRODONTA Perry
Additional bibliography: Moldenke, Rat hate 10: 118--119.
1964; Moldenke, Fifth Summ. 1: 76 (1971) and 2: 916. 1971.

VERBENA MACROSPERMA Speg.
Additional bibliography: Moldenke, Phytologia 16: 100. 1968;
Moldenke, Fifth Summ, 1: 201 (1971) and 2: 916. 1971.

VERBENA MALMIT Moldenke

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 572.
1965; Moldenke, Phytologia 13: 208. 1966; Moldenke, Fifth Summ.
L276 CLG TR) amd 24682 1907 971.

VERBENA MARITIMA Small
Additional bibliography: J. A. Clark, Card Ind. Gen. Sp. Var.
issue 11, 1933; Greene & Blomquist, Fls. South 109. 1953; Molden-

1972 Moldenke, Notes on Verbena 299

ke, Phytologia 16: 100. 1968; Anon., Torrey Bot. Club Ind. Am,
Bot. Lit. 3: 308. 1969; Moldenke in Menninger, Flow. Vines 338,
ph. 28). 1970; Long & Lakela, Fl. Trop. Fla. 71 & 961. 1971; Mol-
denke, Fifth Summ, 1: 10 & 30 (1971) and 2: 521, 525, 653, & 917.
a a

Additional illustrations: Greene & Blomquist, Fls. South 109.
1953; Moldenke in Menninger, Flow. Vines ph. 28h. 1970.

O'Neill encountered this species in low pinelands. Greene &
Blomquist (1953) call it "seaside-verbena" and describe it as
"another flat-topped verbena, the only species often frequenting
coastal situations of dunes, dry pinelands, and hammocks. Flowers
range from rose to bright-purple. Plants spreading to widely
creeping. S. peninsular of Fla."

Additional citations: FLORIDA: Brevard Co.: P, 0. Schallert
20869 (Se--200558); B. Sharp s.n. [9.IV.68] (Ws). Dade Co.: O!
Neill 7596 (Mi). Flagler Co.: J. K. Small s.n, [Ocean City, Ap-
ril 18, 1922] (Se--162929). Indian River Co.: J. K. Small s.n.
{Near Sebastian, June 1928] (Ca--1216765, N). Jupiter Island:
Small, Mosier, & DeWinkeler 10891 (Se--19805).

VERBENA MARRUBIOIDES Cham,

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 572.
1965; Moldenke, Phytologia 16: 100. 1968; Moldenke, Fifth Summ.
1: ihn, 178, 190, & 201 (1971) and 2: 662, 675, & 917. 1971.

The corollas on Hatschbach 1283 are described as having been
"lilac" when fresh. This same distinguished collector, on the
label of his no. 17120, notes "repente, fl6r lilaz, intenso; cam-
po séco, prefere campo esgotado e também barrancos".

Additional citations: BRAZIL: ParanA: Hatschbach 128)3 (W——

2564873), 17120 (Ft, N, W—2536531).

XVERBENA MATRITENSIS Moldenke
Additional bibliography: Moldenke, Phytologia 1h: 289. 1967;
Moldenke, Fifth Summ. 1: 372 (1971) and 2: 660, 673, & 917. 1971.

VERBENA MEGAPOTAMICA Spreng.

Additional synonymy: Glandularia megapotdmica Schnack & Covas,
Revist. Argent. Agron. 12: 22h. 195.

Additional & emended bibliography: J. A. Clark, Card Ind. Gen.
Sp. Var. issue 185. 19; Schnack & Covas, Revist. Argent. Agron.
12: 222-229, fic. 1 B& C, 2A—C, 3 DC, & pl. 12 A, B, E, &
H. 1945; Cabrera, Man. Fl. Alred. Buenos Aires 397 & 398. 1953;
Michalowski, Serv. Tecn. Interam. Coop. Agr. Bol. 169 & 175.
1954; Darlington & Wylie, Chrom, Atl., pr. 1, 323. 1956; Schnack,
Fehleisen, & Cocucci, Revist. Argent. Agron. 2h: 129--135, pl. 1,
A, B, E, & F, & fig. 1. 1957; Darlington & Wylie, Chrom. Atl.,
pr. 2, 323. 1961; Angely, Fl. Anal. Paran., ed. 1, 572. 1965;
Troncoso in Cabrera, Fl. Prov. Buenos Aires 5: 133, 136, & 137,
fig. 6 A & B. 1965; Moldenke, Phytologia 16: 189 & 196. 1968;
Solbrig, Passani, & Glass, Am. Journ. Bot. 55: 1239. 1968; Bolkh.,
Grif, Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 715. 1969;

300 P HY 0. L410 (G7 EA Vol. 23, no. 3

Schnack & Rubens, Bol. Soc. Argent. Bot. 13: 205. 1970; Solbrig,
Princ. & Meth. Pl. Biosystem. 76. 1970; Moldenke, Fifth Summ, 1:
178, 188, 190, 201, & 371 (1971) and 2: 521, 683, 689, 690, 69h,
700, & 917. 1971; Troncoso, Darwiniana 16: (613]--616° & é2i, fig.
1. 1971; Moldenke, Phytologia 23: 239. 1972.

Additional & emended illustrations: Schnack & Covas, Revist.
Argent.,Agron., 2: 225, tice) B eC. 220, fie. 2 A--c; fig. 3 D--
G, -& pls te AO Be Bee As 195; Schnack, Fehleisen, & Cocucci,
Revist. Argent. heron. 2iie 130 fig. 1) & 133, pl. 1 A, Bee
1957; Troncoso in Cabrera, Fl. Prov. Buenos Aires 5: 136, fig. 6
A & B. 1965; Troncoso, Darwiniana 16: 615, fig. 3. 1971.

Schnack & Covas (195) and Schnack & Gonzalez (195) record
the chromosome number of this taxon as n = 10.

Troncoso (1965) cites from Buenos Aires only Burkart 7077 and
Cabrera 301 and gives the distribution of the species as "Sur

del Brasil, Uruguay y litoral argentino. Habita en albardones

altos y matorrales del Delta y Punta Lara." Schnack & Rubens

(1970) record it also from Corrientes, Argentina. The corollas

on Boffa s.n. [Herb. Inst. Miguel Lillo 18] are said to have

been "lilac" in color. The T. Rojas 2111 and Herb. Inst. Miguel

Lillo 107893, distributed as V. “megapotamica, are are actually Vv.

incisa Hook.

Troncoso (1971), in a recent very splendid paper, reviews the
history of this species nomenclaturally and critically compares
three closely related taxa, separating them as follows:

"A. Hierbas o sufritices subglabros y lisos, con pelitos estri-
gosos en tallos y hojas. CAliz ralamente estrigoso, pelitos
antrorsos apretados a la superficie y por lo general con
gl4ndulas subsésiles, pateriformes, antocidncias, sobre las
costillas. Brdcteas oval-oblongas o lanceolado-subuladas,
ciliadas, de 2-5 mm long.........2++esee..G. megapotamica.

A', Hierbas o sufriitices Aspero-hirstitulos, con pubescencia
variada simple y glandulosa en tallos y hojas. CAliz ya
sea con pelos largos patentes y pelitos rigidos mds breves,
o bien densamente cano-retrorso-pubescente, en ambos casos
por lo general con pelitos glandulares estipitados entre-
mezclados. Brd&cteas ovales o lanceoladas, pubescentes o
pestafiosas.

B. Brdcteas lanceoladas, de unos 5 mm long., pestafiosas, por

lo dem4s subglabras. Tubo corolar de 16--18 mm long.
Hojas oval-oblongas, brevemente cuneadas en la base, hi-
pofilo cano-hfspido sobre los nervios, reticulado hirst-
tulo por breves pelitos derechos, rigidos. CAliz con
pelos largos patentes, pelitos subrigidos breves y peli-
tos glandulares estipitados. Mericarpios de 3,5—-\,5
MM. LONG..-eeseesseeesecccccceccscccseccoGe phlogiflora.
B'. Brdcteas ovales u oval-lanceoladas de 1--2 mm long.,
hispido-pubescentes principalmente en la base y bordes.
Tubo corolar de 19--22 mm long. Hojas oblongo o trian-
gular-lanceoladas, subtruncadas o brevemente cuneadas en
la base, hipofilo hirsitulo. CdAliz densamente cano-pubes-

1972 Moldenke, Notes on Verbena 301

cente, pubescencia por lo general retrorso, y pocos peli-

tos glandulares estipitados entremezclados. Mericarpios

de 5--6 mm long....ssccccsccccccccseeeceeeGe Guaranitica."
She gives its distribution as "Sur del Brasil, Uruguay y NE de la
Argentina hasta las barrancas del Rfo de la Plata. Habita en
pajonales y albardones altos de las islas del Delta." She cites
the following specimens: BRAZIL: Rio Grande do Sul: Isabelle s.n.
[1835] (K); Sellow s.n. (K, V). URUGUAY: Osten 5389 (Si) ; Sellow
3162 (K). ARGENTINA: Buenos Aires: Boffa 148 148 (Si); . A. Burkart
7019 (Si), 2203 (Si—983, Si--16063) ; Lanfranchi 496— (Si). Entre
Rfos: A. Burkart 5127 (Si), 8293 (Si), 1513 (Si), 27055 (Si);
Boelke 915; Hung: Hunziker 4625.

Additional citations: ARGENTINA: Buenos Aires: Boffa s.n.

(Herb. Inst, Miguel Lillo 148] (N); A. Burkart 2203 (W—2567980) .

VERBENA MENDOCINA R. A. Phil.

Additional & emended bibliography: Reiche & Phil., Fl. Chil.
5: 295 & 63. 1910; J. A. Clark, Card Ind. Gen. Sp. Var. issue
18). 1944; Darlington & Wylie, Chrom., Atl., pr. 1, 323. 1956; J. F.
Macbr., Field Mus. Publ. Bot. 13 (5): 629. 1960; Darlington & Wy-
lie, Chrom. Atl., pr. 2, 323. 1961; Troncoso in Bocher, Hjerting,
& Rahn, Dansk Bot. Arkiv 22 (1): 109. 1963; Hocking, Excerpt. Bot.
A.10: 270. 1966; Moldenke, Phytologia 16: 189. 1968; Bolkh., Grif,
Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 715 & 717. 1969; Molden-
ke, Fifth Summ. 1: 190, 201, & 371 (1971) and 2: 521, 683, 688, &
917. 1971.

Troncoso (1963) cites Bocher, Hjerting, & Rahn 2070, collected
at an altitude of 1650 meters. Beetle (1943) cla classifies this spe-
cies in the Cruciferae, doubtless through some stenographic error.
He reports that Macloskie is of the opinion that Patagonia depends
on constant replenishment of its flora from the Andean mountain
chain; Neger suggests that the river courses are probably the mig-
ratory lanes for the entrance of such plants as Verbena mendocina.

VERBENA MENTHAEFOLIA Benth,

Additional bibliography: Abrams, Illustr. Fl. Pacif. States,
pr. 1, 3: 610, 611, & 616, fig. 4343. 1951; Howell & McClintock in
Kearney & Peebles, "ariz. Fl., ed. 2, 726 & 728. 1960; Ferris in
Abrams & Ferris, Tllustr. Fl. Pacif. States, pr. 1, fh: 651 & 730
(1960) and pr. 2. h: 651 & 730. 1965; Abrams, Tlustr. Fl. Pacif.
States, pr. 2, 3: 610, 611, & 616, fig. h313. 1967; Moldenke, Phy-
tologia 16: 100-101. "1968; Munz & Keck, Calif. Fl. 686, 687, &
1679. 1968; Rickett, Wild Fls. U. S. 3 (2): 364 (1969) and (3):
540 & 799. *1970; Gibson, Fieldiana Bot. 2 (9): 230 & 233. 1970;
Correll & Johnston, Man, Vasc. Pl. Tex. [Contrib. Tex. Res. Found.
Bot. 6:] 1876 & 1877. 1970; Moldenke in Correll & Johnston, Man.
Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:] 131) & 1318.
1970; Moldenke, Fifth Summ. 1: 59, 63, 66, 76, 81, 206, & 371 (1971)
and 2: 67h, 682 683, 686, 687, 695, 697, *& 917. 1971; *voldenke,
Phytologia 22: 473 &’99 (1972) and 23: 184 & 188. 1972.

302 P BLY BO E:01G; ek Vol. 23, no. 3

Additional illustrations: Abrams, Illustr. Fl. Pacif. States,
pr. 1, 3: 616, fig. 4343 (1951) and pr. 2, 3: 616, fig. 4343. 1967.
Recent collectors have found this plant growing in pastizal,
matorral, Opuntia matorral, woods, oak or pine and oak woods, cul-
tivated ground a nd old cultivated fields, on slopes with pine-oak

woods, on grassy banks and low banks near water, and at the edge
of lakes, flowering (in addition to the months previously reported
by me) and fruiting in November, Gonzdlez Quintero found it in
shrubby matorral of Juniperus in Hidalgo, Mexico, while in the
same state Mears reports it growing in association with Cassia,
Cuphea, Indigofera, Juniperus, Mimosa, Phoradendron, Quercus, and
Teucrium. In Zacatecas it was “Saute in Saeed soil in
full sun on a level plateau with microphyll desert scrub of the
Prosopis~Acacia association by Detling. Breedlove describes it as
a suffrutescent perennial growing under shrubs on tablelands
heavily covered with shrubs in Sonora, while he and Kawahara found
it on slopes with Arbutus xalapensis, "Quercus epileuca, Q. urbani,
Pinus ayacahuite, and P. and P. lumholzii in Sinaloa. Cruz C. and areas
N. came upon it on "ladera bas4ltica con vegetacién de matorral

de Acacia y Opuntia" in México and in "terrenos fangosos" in Hid-
algo. In the state of México, also, Morales Didz found it on
"ladera andesftica con vegetacién secundaria" and Rico A. in "en-
cinar abierto y pastizales". An additional vernacular name var=-
iant is "telran".

The corolla is described as having been "very faint purple"
on Hinton 11991, "red-purple" on Lagunas & Castillo 17, "purple"
on Rico A. 31, J. Rzedowski 1088, 20302, & 22205a, "violet" on
Franco R. 8 sen. and Garcfa Saucedo Sone, "pale-lavender" on Die-
terle 357h, "blue" on Breedlove dlove 1198, : and "pale-blue" on Breed-
love | & . Kawahara 16815.

Pennington reports that in Chihuahua and Sonora the entire
plant is boiled in preparing a tea for use by the natives against
stomach disorders and to stop vomiting, and also that goats browse
upon the plant.

Howell & McClintock (1960) cite Jones s.n. [near Yuma, 1906].
Material has been misidentified and distributed in some herbaria
under the names V. bipinnatifida var. latilobata Perry, "V.
neomexiczna (A. Gray) Small", and V. pinetorum Moldenke. “The Je
Rzedowski 22205a, cited below, is a mixture with V. ciliata Benth.,
while Franco R. s.n. [20.VIII.1967], Garcfa Romero | son. [11/Vile
1968], and M4rquez Soe [23/VI1/1962] are m are mixtures with V. caro-
lina L. The R. Pearce ree 2276, distributed as V. menthaefolia, is is
actually V. halei Small, wh while Pinkava, Keil, | & Lehto 14553 is V.
neomexicana var. ape, epleens Perry.

Additional citations: MEXICO: Baja California: Mrs. F. Wylie

s.n. [April 8, 1950] (Sd--)3915). Chihuahua: Kruckeberg g 14928
(Se--207L95) ; Pennington 3 (Au--287715); Stuessy 990 (Au-—
257h6h). Federal District: Huerta M. 2 (Ip); Marquez sen. [23/

1972 Moldenke, Notes on Verbena 303

VII/1962] (Ip); J. Rzedowski 1088 (Au—2)1215, Ip); Villegas Ds
80 (Ip), 550 (Ip). Hidalgo: Chavez 0. sn. (i, .vIrI Be (Ip

Cruz C. Lov" (Ac); Garcfa Saucedo s.n. ‘s [10.1X.1966] (Ip); eee
Quintero 257 (Ip), 304 (Ip), 931 (Ip), 994 (Ac), 1265 (Ip), 2590
(Ip), 2882 (Ip, Mi); Lo Lagunas & & Castillo 17 at (Ip); Mears 259a (Au--
255097), 326a (Au--254967). Jalisco: J. V. A. Dieterle 3574 (Mi).
México: Cruz ruz C. 564 (Mi); Franco R. s.n. [20.VIII.1967] (Ip);
Morales Didz 3. gn. . [22/VII/1962] (Ip); Rebolledo Vélez s sen. [20.
VIII.1967] (Ip); | Rico A. 31 (Rf); J. Rzedowski 20302 (RE), 22205a,
in part (Ip), 222078 (Ip), , 24,086 (Ip); Vargas N. s.n. [2h.1V.
1966] (Ip). Michoac4n: Garcfa R Romero s.n. on. [11.VIII 1968] (Ip);
Hinton 11991 (Mi, Se--117\42). “Sinaloa: Breedlove = Kawahara
16815 (Rf). Sonora: Breedlove 1498 (Au—232106); Pennington 66 nington 66
(Au—2876h2) , 98 (Au--287612) . “Veracruz: Dodds “g oF Zaca—
tecas: Detling - 9436 1/2 (Ip). State undetermined: J. Rzedowski
1133 [La aaertrry (Ip).

XVERBENA MERETRIX Moldenke
Additional bibliography: Moldenke, ae ne Ids: 472. 19655
Moldenke, Fifth Sum, 1: 371 (1971) and 2: 67h, 686, & 917. 1971.

VERBENA MICROPHYLLA H.B.K.

Emended synonymy: Verbena microphylla Humb. & Bonpl. ex Steud.,
Nom, Bot. Phan., ed. 1, 873. 1821. Glandularia microphylla
(H.B.K.) Cabrera, Revist. Invest. Agric. 11: 398. 1958.

Additional bibliography: Cabrera, Revist. Invest. Agric. ll:
332 & 398. 1957; Cabrera, Bol. Soc. "argent. Bots "72 150. 19565 Js
A. Clark, Card Ind. Gen. Sp. Var. issue 228. 1958; J. F. Macbr.,
Field Mus. Publ. Bot. 13 (5): 614 & 625--626. 1960; Troncoso in
Bocher, Hjerting, & Rahn, Dansk Bot. Arkiv 22 (1): 109. 1963;
Meyer & Weyrauch, Inst. Mig. Lill. Misc. 23: 33 & 123. 1966; G.
Taylor, Ind. Kew. Suppl. 13: 61. 1966; Moldenke, Phytologia "16:
101 & 206. 1968; Schnack & Rubens, Bol. Soc. Argent. Bot. 13: 205.
1970; Moldenke, Fifth Sum. 1: 37, Uy, 184, 190, 193, & 201
(1971) and 2: 521, Su1, 678, 68h, & 917. 1971; Moldenke, Phytolo-
gia 23: 199. 1972.

Recent collectors have encountered this plant on grazed grassy
paramos. Macbride (1960) comments that the species is "Scarcely
more than a part of V. laciniata (L.) Briq. but perhaps recogniz-
able and apparently a a high Andean state; listed by Ball as 'V.
diffusa Willc.'" He gives its extra-Peruvian distribution as "To
Uruguay and Ecuador" and cites the following collections from Pe-
ru: Ancash: Macbride i7l, Weberbauer 302. Ayacucho: Ferreyra
5509. Cuzco: Vargas 9827. Lima: Ball 3. s.n., Ferreyra 3531, Mac-
bride 169, Weberbauer 226. Puno: Pennell 11 13359, J. Sharpe 58,
Soukup 70 & 870. Troncoso (1963) cites Kurtz rtz 108.

The Cabrera (1957) reference cited above is sometimes cited as
"1958", but is plainly dated "1957" in the original.

[to be contimed]

A NEW COMBINATION IN THE GENUS EUTHAMIA

David J. Sieren
Department of Biology
University of North Carolina at Wilmington

Euthamia hirtipes (Fernald) Sieren comb. nov. was first

named by M. L. Fernald (1946) from specimens collected in
Sussex Co., Virginia. Fernald included the new species in the
genus Solidago and proposed it to be the result of hybridization
between S. graminifolia (L.) Salisb. var. nuttallii (Greene)
Fernald and S. microcephala (Greene) Bush inasmuch as the
specimens were morphologically intermediate between the two and
the ranges of the proposed parents would allow it. Since then,
several specimens have been collected from along the coasts of
the Carolinas which bear a strong resemblance to the photo and
ae ae of the type in Fernald's paper. Because the name

. hirtipes has generally been overlooked, however, the more
eee collections have usually been referred to either S. lepto-
cephala T. & G. or S. tenuifolia Pursh, both of which are closely
allied to S. hirtipes. S. leptocephala is a species of the lower
Mississippi valley extending from western Tennessee and Kentucky
west to Texas while S. tenuifolia [including S. microcephala
(Greene) Bush] is a narrower-leaved species also found along the
coast, but extending from Louisiana to Florida north to New
England. The species group (Solidago section Euthamia) to which
S. hirtipes belongs has often been considered to constitute a
separate genus Euthamia (Nuttall, 1840; Greene, 1902; Shinners,
1951). That treatment is accepted by this author as the more
natural one and for annotation purposes therefore the new combin-
ation becomes necessary.

Literature Cited

Fernald, M. L. 1946. Technical studies on North American
plants. Rhodora 48: 65-66.

Greene, E. L. 1902. A study of Euthamia. Pittonia 5: 72-80.

Nuttall, T. 1840. Descriptions of new species and genera of
plants in the natural order of the Compositae. In
Trans. Amerneeehtl Soc. ln, 0/ 1) 325-520)

Shinners, L. H. 1951. The Texas species of Euthamia (Com-
positae). Field and Lab. 19: 137-138.

30h,

STUDIES IN THE EUPATORIEAE (ASTERACEAE). LXVII.

GRAZIELIA NOM. NOV. FOR DIMORPHOLEPIS

R. M. King and H. Robinson
Smithsonian Institution, Washington, D.C. 20560

A second example of a homonym among the recently
named genera of the Eupatorieae has been called to
our attention by Dr. Max Gray of Australia. The name
Dimorpholepis (Barroso) R.M.King & H.Robinson (1971a)
is preoccupied at the generic level by Dimorpholepis
A. Gray an Australian plant now placed in the genus
Helipterum DC. The two Later homonyms that we pro-
En at nearly the same time, Neobartlettia and
Dimorpholepis, are particularly regrettable because
of the ease with which they could seemingly have been
avoided. Almost every other phase of the present
series of studies is more difficult including the
search for an increasing number of new and appropriate
names. Here is not so much the failure to check as
the failure to make it a practice to check many times.
Both of these names we did think we had checked while
obviously we had not.

The new name that we propose for Dimorpholepis
(Barroso) R.M.King & H.Robinson is Grazielia, honor-
ing again Dr. Graziela Maciel Barroso, the leading
authority on Brazilian Compositae.

Grazielia R.M.King & H.Robinson, nom nov. for

Dimorpholepis (Barroso) R.M.King & H.Robinson,
Bist cloci= 5D: LUB=1202"2971%

Type species: Eupatorium dimorpholepis Baker.

Our studies of the genus indicate that it
contains the following nine species.

Grazielia anethifolia (A.P.Decandolle) R.M.King &

eRobinson, comb. nov. Eupatorium anethifolium
A.P.Decandolle, Prodr. 5: 182. 1836. Brazil.

Grazielia coriacea (Scheele) R.M.King & H.Robinson,
comb. nov. Eupatorium coriaceum Scheele, Linnaea
18: 457. 1844. Brazil.

305

306 Poo Yt 10 LeOsGer ks Vol. 23, no. 3
Grazielia dimorpholepis (Baker) R.M.King & H.Robinson,

comb. nov. Eupatorium dimorpholepis Baker, Mart.
FL. Brasi. 6T: 55s: te7o¢ eeaeri.

Grazielia gaudichaudeana (A.P.Decandolle) R.M.King &
H.Robinson, comb. nov. Eupatorium gaudichaudeanum
A.P.Decandolle, Prodr. 5: 148. 1838. Brazil.

Grazielia intermedia (A.P.Decandolle) R.M.King &

H.Robinson, comb. nov. Eupatorium intermedium
A.P.Decandolle, Prodr. 5: 148. 1836. Brazil.

Grazielia mollissima (Sch.-Bip. ex Baker) R.M.King &
H.Robinson, comb. nov. Eupatorium mollissimum
Sch.-Bip. ex Baker, Mart. FL. Bras. 6(2): 331.
S762 Brazil.

Grazielia multifida (A.P.Decandolle) R.M.King & H.
Robinson, comb. nov. Eupatorium multifidum
A.P.Decandolle, Prodr. ES 182. 1836. Brazil.

Grazielia numnularia (Hook. & Arn.) R.M.King & H.
Robinson, comb. nov. Eupatorium numnuLlaria
Hook. & Arn., Hook. Comp. Bot. Mag. L: 241.
ESs5.oBrazit,.

Grazielia serrata (Spreng.) R.M.King & H.Robinson,

comb. nov. Eupatorium serratum Spreng., Syst.
3: 415. 1826. Argentina, Brazil, Uruguay.

Reference
King, R.M. & H.Robinson 1971. Studies in the
Eupatorieae (Asteraceae). LV. The genus,
Dimorpholepis. Phytologia 22: 118-120.
Acknowledgement
This study was supported in part by the National

Science Foundation Grant GB- 20502 to the senior
author.

STUDIES IN THE EUPATORIEAE (ASTERACEAE). LXVIII.

A NEW GENUS, CONOCLINIOPS IS

R. M. King and H. Robinson
Smithsonian Institution, Washington, D.C. 20560

The genus we name here as Conocliniopsis is
based on a single species distributed widely in South
America from Colombia to southern Brazil. The species
is most closely related to the genus Barrosoa but
differs most prominently by the densely setiferous
achene and the extremely high conical receptacle. In
addition to the technical differences between
Conocliniopsis and Barrosoa, the former single species
has a distribution which includes the entire range
of all the species of the Latter and there is every
reason to believe that the two genera have a Long
distinct evolutionary history.

The species that we place here in Conocliniopsis
is very distinct and well known to all students =;
the Asteraceae of South America, but the name
Eupatorium ballotaefolium H.B.K. usually applied to
the species is incorrect. In a note on Eupatorium
ballotaefolium, B. L. Robinson (1918) mentioned some
phylLlary characters of typical material and he named
a new variety. The two varieties recognized by
B. L. Robinson actually differ by far more than
phyllary characters. Typical E. ballotaefolium has
phyllaries more blunt and less pubescent, the backs
of the corolla lobes covered with short hairs, the
achenes nearly glabrous and extremely constricted
below the pappus, the carpopodium stopper shaped
with small cells and without setae on the upper rim,
and the receptacle is plain. True Eupatorium ballot-
aefolium, in fact, is a Lourteigia which differs from
the description of that genus ting & Robinson, 1971)
only by the larger number of flowers per head (up to
40). The Eupatorium ballotaefolium var. caucense
B. L. Robinson with phyllaries more pointed and
hirsute, with backs of the corolla Lobes Lacking
hairs, with the achenes densely setiferous without
a constriction above, with a carpopodium broad
with setae on the upper rim and a high conical
receptacle; is the same plant previously described
as Conoclinium prasiifolium A.P.Decandolle and which
we Place here in ConocLiniopsis. The South American

307

308 Puy 7 1.0.L 0,4 0 A Vol. 23, nO. 3

material recognized in present treatments under the
name Eupatorium ballotaefolium H.B.K, should be placed
in the following two taxa.

Lourteigia ballotaefolia (H.B.K.) R.M.King & H.Robin-
son, comb, nov. Eupatorium ballotaefolium H.B.K.,
Nove) Gen. set Sp. 7ase95, LS8l8. Eds Folzoz

Colombia.

Conocliniopsis R.M.King and H.Robinson, genus
novum Asteracearum (Eupatorieae). Plantae herbaceae
erectae pauce ramosae. Caules teretes. Folia opposita
vel ex parte alterna distincte petiolata, Laminis
ovatis margine crenulatis. Inflorescentiae dense
corymbosae paniculatae vel cymosae. Involucri squamae
eximbricatae 12-16 subaequilongae hirsutae et gland-
uliferae; receptacula alte conica glabra. Flores
20-30 in capitulo; corollae anguste infundibulares
5-lobatae extus glanduliferae intus glabrae, cellulis
angustis parietibus sinuosis, lobis aequilateraliter
triangularibus vel longioribus extus ad apicem
papillosis intus mamillosis vel sublaevibus; fila-
menta antherarum in parte superiore angusta, cellulis
plerumque elongatis, parietibus annulate ornatis,
cellulis exothecialibus subquadratis, appendicibus
antherarum ovatis; styli inferne non incrassati
glabri, appendicibus lLinearibus breviter papillosis;
achaenia prismatica 4-5-costata valde setifera;
carpopodia valde distincta brevia, superne valde
setifera, cellulis subquadratis inflatis, parietibus
parum incrassatis; pappus setiformis uniseriatus,
setis 30-35 scabris ad.apicem tenuibus, cellulis
apicalibus acutis vel subacutis.

Species typica: Conoclinium prasiifolium A.P.
Decandolle.

The genus is monotypic.

Conocliniopsis prasiifolia (A.P.Decandolle) R.M.King
& H-Robinson, comb. nov. Conoclinium prasiifolium
A.P.Decandolle, Deless. Ic. Sel 4: t TS L840.

Eupatorium ballotaefolium H.B.K. var. caucense
Beli Robanson, (Contr. Gray’ Herb. ‘ntis. SSsrZore
1918. Colombia, Venezuela, Brazil.

References

King, R.M. & H.Robinson. 1971. Studies in the Eupat-
orieae (Compositae). XXXV. A new genus,

1972 King & Robinson, A new genus, Conocliniopsis 309
Lourteigia. Phytologia 21:28-30.

Robinson, B.L. 1918. Diagnoses and notes relating to
tropical American Eupatorieae. Contr. Gray Herb.
Dos 235-263.

Acknowledgement
This study was supported in part by the National

Science Foundation Grant GB- 20502 to the senior
author.

STUDIES IN THE EUPATORIEAE (ASTERACEAE). LXIX.

A NEW GENUS, GYPTIDIUM.

R. M. King and H. Robinson
Smithsonian Institution, Washington, D.C. 20560

Among the Gyptoid species in eastern South
America are three with the corollas extremely long
and narrow in the basal part. One of these species
we have already covered in our treatment of the genus
Urolepis (King & Robinson, 1971). The other two
species treated here seem related to Urolepis by the
form of the corolla and by the blunt cells at the
tips of the pappus setae, among other features.

These species lack two features that are very prom-
inent in Urolepis, however, the extremely long
papillae on the stylar appendage and the large carpo-
podium with large thin-walled cells. The corolla
lobes of the two species are also slightly to strong-
ly papillose while those of Urolepis are essentially
smooth. A final distinction of oe is the
greatly expanded receptacles bearing LOO-150 flowers.

On the basis of the characters given above we
recognize here the genus Gyptidium with two species.
Still, the two species have notable differences.
Gyptidium militare has a paleaceous receptacle and
rather poorly developed apical cells on the pappus
setae, while G. trichobasis has only hairs on the
receptacle as in Urolepis and has very enlarged cells
on the tips of the pappus setae.

As presently known the genus Gyptidium is
restricted to southern Brazil and adjacent Argentina.

Gyptidium R.M.King & H.Robinson, genus novum
Asteracearum (Eupatorieae). Plantae herbaceae erectae
raro ramosae. Caules teretes striati. Folia opposita
vel ex parte alterna distincte petiolata, lLaminis
ovatis vel lanceolatis basi truncatis margine cren-
ulatis. Inflorescentiae corymboso-paniculatae vel
cymosae. Involucri squamae subimbricatae vel
eximbricatae ca. 25 subaequilongae alte glanduliferae;
receptacula conica hirsuta vel ubique anguste pale-
ifera. Flores 50-80 in capitulo; corollae inferne
perangustatae superne anguste campanulatae 5-lobatae
extus glanduliferae intus glabrae, cellulis angustis
310

1972 King & Robinson, A new genus, Gyptidium 311

parietibus sinuosis, lLobis aequilateraliter triangul-
aribus vel Longioribus extus ad apicem papillosis
intus sublaevibus vel papillosis; filamenta anther-
arum in parte superiore angusta, cellulis infernis
plerumque quadratis, parietibus annulate ornatis,
cellulis exothecialibus subquadratis, appendicibus
antherarum ovatis; styli inferne non incrassati
glabri, appendicibus Linearibus valde dense papillosis;
achaenia prismatica 4-5-costata glandulifera; carpo-
podia obsoleta; pappus setiformis uniseriatus, setis
25-35 scabris cellulis apicalibus obtusis plerumque
dilatatis.

Species typica: Eupatorium militare B.L.Robinson

Our studies of the genus indicate that it con-
tains two species.

Gyptidium militare (B.L.Robinson) R.M.King & H.Rob-
inson, comb. nov. Eupatorium militare B.L.Rob-
inson, Contr. Gray Herb. n.s. 65: 50. 1922.
Argentina.

Gyptidium trichobasis (Baker) R.M.King & H.Robinson,,
comb. nov. Eupatorium trichobasis Baker, Mart.
Pl. Bras. 6C): 364. 1876. Brazil.

Reference

King, R.M. & H.Robinson. 1971 Studies in the Eupator=
ieae (Asteraceae). XL. The genus, Urolepis.
Phytologia 21: 304-305. ;

Acknowledgement

This study was supported in part by the National

Science Foundation Grant GB- 20502 to the senior
author.

STUDIES IN THE EUPATORIEAE (ASTERACEAE). LXX.

A NEW GENUS, BAHIANTHUS.

R. M. King and H. Robinson
Smithsonian Institution, Washington, D.C. 20560

The state of Bahia in northeastern Brazil is
noted for its distinctive flora. Of the many endemic
Eupatorieae, certain species of Agrianthus and Stylo-
trichium are perhaps the best known. Careful invest-
lgation has shown that another eupatorian endemic
represents a distinct genus which we name here after
the state of Bahia.

The single species Bahianthus viscosus has a
synonymy unusually representative of the generic chaos
that has prevailed in the Eupatorieae. The species
has been placed in Mikania, Kuhnia, Gyptis and
Symphyopappus, only recently being placed in
Eupatorium, (Steyermark, 1953) the genus where its
closest relatives have long resided. The species is
strikingly similar in vegatative aspect to the genus
Symphyopappus and a relationship is impossible to
ignore, but Bahianthus has an irregular flower number
ranging from 15-22, papillose corolla lobes, and has
a conical receptacle, three characteristics not found
in the Disynaphioid complex to which Symphyopappus
belongs. Bahianthus actually seems most closely
related to Barrosoa in the Gyptoid complex which also
has the achenes with glands but no setae on the sides
and the carpopodia with Large thin-walled cells.

The new genus differs from Barrosoa in its vegatative
appearance with glabrous obovate Leaves and also in
the details of the style branches which are dilated
at the tips and densely papillose. Bahianthus can
also be distinguished by the very thick base on the
corolla, the large cells of the corolla lobes, the
slightly Larger than usual pollen with unusually

long spines and the coarser more irregular pappus
setae.

Bahianthus R.M.King and H.Robinson, genus novum
Asteracearum (Eupatorieae). Plantae herbaceae vel
frutescentes erectae pauce ramosae. Caules teretes
striati. Folia alterna distincte petiolata, glabra,
laminis coriaceis obovatis basi cuneatis ad apicem
obtusis serrulatis. Inflorescentiae corymboso-panic-

312

1972 King & Robinson, A new genus, Bahianthus 313

ulatae; pedicelli glabri striati. Involucri squamae
18-20 subimbricata fere glabra; receptacula conica
glabra. Flores 15-22 in capitulo; corollae tubulares
carnosae 5-lobatae extus inferne glabrae intus
glabrae, cellulis oblongis parietibus interioribus
sinuosis, lobis aequilateraliter triangularibus vel
longioribus extus glanduliferis superne papillosis
intus mamillosis; filamenta antherarum in parte
superiore lata, cellulis plerumque quadratis pariet-
ibus annulate vel intricate ornatis, cellulis exothec-
ialibus subquadratis, appendicibus antherarum ovatis;
Styli inferne non incrassati glabri, appendicibus
elongatis ad apicem parum dilatatis dense erecte
papillosis; achaenia prismatica 4-5-costata parce
glandulifera, punctis saepe in seriebus transversis,
costis in parte inferiore callosis; carpopodia
distincte incrassata, cellulis quadratis inflatis,
parietibus tenuibus; pappus setiformis uniseriatus,
setis ca. 30 breviter ciliato-dentatis, dentibus
inferioribus irregulariter dispositis superioribus
congestis, cellulis apicalibus angustis obtusis.

Species typica: Mikania viscosa Spreng.

The genus is monotypic.

Bahianthus viscosus (Spreng. ) R.M.King & H.Robinson,
comb. nov. Mikania viscosa Spreng. Neue Entdeck..
lL: 277. 1820. Kuhnia baccharoides A.P.Decandolle,
Prodr. 5: 128. 1836. Gyptis baccharoides Schultz-
Bip. Mss in herb. Sonphyaaanpas viscosus
Schultz-Bip. ex Baker, Mart. Fl. Bras. 6(2):366.

1876, Eupatorium harvardianum Steyermark, Field-
iana: Botany 28:636. 1953.
Reference
Steyermark, J.A. 1953. Botanical Exploration in
Venezeula. Fieldiana: Botany 28(3):636.
Acknowledgement
This study was supported in part by the National

Science Foundation Grant GB-20502 to the senior
author.

FIVE MORE VERBENACEOUS NOVELTIES

Harold N. Moldenke

AEGIPHILA GLORIOSA var. PARAENSIS Moldenke, var. nov.

Haec varietas a forma typica speciei pilis ubique multo brevi-
oribus minore divaricatis recedit.

This variety differs from the typical form of the species in
having its pubescence throughout much shorter and far less con-
spicuously divaricate-spreading.

The type of the variety was collected by M. Silva and R. Souza
(nee 2253) in woods on terra firme from Ramal to Novo Imperio, at

35 on the road to Palh&o, Santamém, Para, Brazil, on August
il, 1969, and is deposited in my personal herbarium at Plainfield,
New Jersey. The collectors describe the plant as a shrub, 3
meters tall, with yellow flowers. It has much the overall appear-
ance of A. racemosa Vell.

AEGIPHILA MICROCALYCINA Moldenke, sp. nov.

Frutex scandens; ramulis junioribus gracillimis densissime
adpresseque puberulentibus, pilis antrorso-strigosis; internodiis
elongatis; foliis decussato-oppositis; petiolis gracillimis brev-
issimis densissime adpresseque puberulentibus; laminis foliorum
membranaceis ellipticis vel lanceolatis supra glaberrimis niti-
disque, subtus in reticulo venarum adpresso—puberulentibus,
caetero subglabratis punctulatis, ad apicem acuminatis, ad basin
abrupte acutis; inflorescentiis axillaribus terminalibusque multi-
floris laxis, ramulis densissime adpresso=-puberulis; calyce parvo
patente-campanulato parce strigilloso margine )-dentato; corollae
tubo pergracillimo glaberrimo.

Woody vine, the uppermost branchlets apparently very slender,
very densely appressed—puberulent with strigose russet—brownish
subantrorsely oriented hairs; principal uppermost internodes much
elongated, 2.5--6 cm. long, the nodes flattened and somewhat am-
pliate; leaves decussate-opposite, numerous, decreasing in size
toward the tips of the branchlets and among the inflorescences;
petioles very slender, 3—8 mm. long, very densely appressed-
puberulent with russet-brown antrorsely strigose very short hair;
leaf-blades elliptic or somewhat lanceolate, 6--12 cm. long, 2—-5
cm. wide, acuminate at the apex, abruptly acute at the base, com-
pletely glabrous and very shiny above, glabrous or subglabrate
and shiny beneath except for the densely strigose-puberulent lar-
ger venation, densely punctulate between the veins, the midrib
and the approximately 6 pairs of arcuate-ascending secondaries
very slender, the latter arcuately joined in many loops at the
margins; inflorescence abundant, axillary and terminal, at the
tips of the youngest branchlets, more or less pyramidal, rather
foliose, the peduncles, sympodia, and pedicels very slender and
very densely strigose-puberulent with antrorsely appressed russet-
brown hairs; bracts rather few, the lowermost foliaceous, the up-

1972 Moldenke, Verbenaceous novelties 315

per linear-spatulate and densely appressed-strigose; calyx broad-
ly campanulate, relatively small, 1 mm. long, 2 mm. wide, extendirg
far beyond the corolla-tube on both sides, very sparsely scatter-
ed-pilose, apparently herbaceous, the rim plainly l-dentate; cor-
olla cream-colored, very distinctly hypocrateriform, the tube very
slender, about 5 mm. long, less than 0.5 mm. wide except at the
ampliate apex, completely glabrous and shiny on the outer surface,
the limb completely glabrous, the lobes , oblong-lingulate, wide-
spreading in anthesis, about 2 mm. long and 1 mm. wide, subacute
or obtuse at the apex; filaments long-exserted, white, projecting
6--8 mm. beyond the corolla-tube,

The type of this very distinct species was collected by G. T.
Prance, W. C. Steward, E. P. Harter, J. F. Ramos, W. S. Pinheiro,
and 0, *P, Monteiro (no. 10907) at the margin of the Uaicd air-
strip, Rio Uraricoeira, at 3°33! N., 63°11' W., Roraima, Brazil,
on March 12, 1971, and. is deposited in my personal herbarium at
Plainfield, ” New Jersey. Superficially this species greatly re-
sembles A. ’ vitelliniflora Klotzsch, but its calyx characters at

once distinguish it with ease.

CLERODENDRUM KLEMMEI var. PUBERULUM Moldenke, var. nov.

Haec varietas a forma typica speciei inflorescentibus dense
puberulentibus recedit.

This variety differs from the typical form of the species in
having the inflorescences, including the peduncles, sympodia,
pedicels, calyxes during anthesis, and outer surface of the
corolla-tubes, densely puberulent.

The type of the variety was collected by Maximo Ramos [Herb,
Philip. Bur. Sci. 7251] in the province of Abra, Luzon, Philip-
pines, in January or February of 1909, and is deposited in the
United States National Herbarium at Washington. The general ap—
pearance of the inflorescence is much like that seen in C. quadri-
loculare (Blanco) Merr. or C. mindorense Merr., but the calyx-
lobes are quite different,

CLERODENDRUM MINAHASSAE var. GRANDICALYX Moldenke, var. nov.

Haec varietas a forma typica speciei calycis floriferis cm.
longis 8--10 mm, latis et tubo corollae 11 cm. longo recedit.

This variety differs from the typical form of the species in
having its calyx during anthesis ) cm. long and only 8--10 m,
wide and the corolla-tube 11 cm. long.

The type of the variety was collected by Chester A. Wenzel
(no. 261) on the island of Leyte, Philippines, sometime in 1913,
and is deposited in the United States National Herbarium at
Washington.

VITEX TRIFLORA var. HIRSUTA Moldenke, var. nov.

Haec varietas a forma typica speciei pedicellis tuboque caly-
cis lobisque calycis patente hirsutis recedit.

This variety differs from the typical form of the species in
having its pedicels, calyx-tubes, and calyx-lobes conspicuously

316 PAY eT O L-OnGel ah Vol. 23, no. 3

rather long-hirsute with brown and distinctly wide-spreading
hairs.

The type of the variety was collected by P. J. M. Maas, K. Ku-
bitzki, W. C. Steward, J. F. Ramos, W. S. Pinheiro, and J. F.
Lima (no. P.13120) in a forest on terra firme at Aldeota, between
Porangaba and Papagaio, Rio Jurud-Mirim, Cruzeiro do Sul, along
the Rio Jurud and/or Rio Moa, Acre, Brazil, on May 18, 1971, and
is deposited in my personal herbarium at Plainfield, New Jersey.
The collector describes the plant as a tree, | meters tall, with
pale-blue flowers.

BOOK REVIEWS

Alma L. Moldenke

"INSECT RESISTANCE IN CROP PLANTS" by Reginald H. Painter, 2nd
edition, xi & 520 pp., illus., University of Kansas Press,
London & Lawrence, Kansas 66044. 1968. 0 sh. or $4.75
paperback.

The first edition of this very fine work appeared in 1951 and
recently has become scarce as the proverbial "hens' teeth". Yet
this field of study has only grown in importance because of in-
creasing food needs of our increasing population, because of in-
creasing insecticide immunity, and because of the growing aware-
ness of the serious wide-spread havoc upon the ecosystem caused
by the excessive use of non-biodegradable insecticides.

The format and content are basically the same. They deal
with the mechanisms of resistance, the factors affecting the ex-
pression or the permanence of resistance, the resistance to in-
sects in wheat, corn, cotton, sorghums and potatoes, and the
methods and problems in breeding for resistance. There is a top-
ically arranged bibliography which was one of the treasures of
the first edition. Little has been added to this new edition to
bring it up to date.

The preface to this new paper—bound edition closes with this
orientation: "In contrast to the use of insecticides, where re-
sults are sudden and there is decreasing effectiveness until re-
applied, insect-resistant varieties are more permanent and cumu-
lative in effectiveness. This is especially true of low levels
of insect resistance, which have not received the attention they
deserve. The use of insect-resistant varieties should be of in-
creasing value around the world in the coming development of in-
tegrated insect control."

"HAWAIIAN HERBS OF MEDICINAL VALUE" - Found Among the Mountains
and Elsewhere in the Hawaiian Islands, and Known to the

1972 Moldenke, Book reviews 317

Hawaiians to Possess Curative and Palliative Properties

Most Effective in Removing Physical Ailments, by H. F. Berg-
man and the Territorial Board of Health, 7 pp., facsimile
reprint by Charles E. Tuttle Co., Tokyo, Japan & Rutland,
Vermont 05701. 1972. $3.00 paperback.

This work was first published in 1922, has long been out of
print, and is now reprinted "in the interest of cultural research"
Almost 200 plants are listed alphabetically by the Hawaiian names,
are then described as to their curative effects and localities,
and are given their scientific names where known,

Since this is a facsimile reprinting the original spelling er-
rors persist for Chenopodium, Cibotium, Pittosporum, Colocasia
and Ipomoea. For manawanawa the correct (oldest) scientific
name in the varietal category is Vitex trifolia var. simplicifo-
lia Cham,
~~ The material makes interesting reading.

"DRAWINGS OF BRITISH PLANTS" Being Illustrations of the Species
of Flowering Plants Growing Naturally in the British Isles,
by Stella Ross-Craig, 109 pp., illus., G. Bell & Sons, Ltd.,
London W.C.2. 1971. £1.00.

This is Part XXVIII of the series, comprising 53 beautifully
accurate plates illustrating members of the Hydrocharitaceae and
the Orchidaceae. There is an index of these plants included.

"COMMON WEEDS OF THE UNITED STATES" prepared by the Agricultural
Research Service of the United States Department of Agri-
culture, 63 pp., illus., Dover Publications, Inc., New
York, N. Y. 10014. 1971. $4.50.

This is an unaltered replication of "Selected Weeds of the
United States" originally published by the Govermment Printing
Office for the United States Department of Agriculture in 1970.
The clear text and the fine geographic distribution maps are the
careful work of the botanist, Clyde F. Reed, and the excellently
accurate and lifelike drawings are the skilled work of the scien-
tific illustrator, Regina 0. Hughes.

It is good to have this helpful and attractive book easily
available at a very reasonable price,

The printing is easily legible. Only the varietal name,
pinnatifidum, on p. 389 was noticed as being misspelled,

Comparison with H. N. Moldenke's recent (1971) "Fifth Sum-
mary of the Verbenaceae, Avicenniaceae, Stilbaceae, Dicrastyli-
daceae, Symphoremaceae, Nyctanthaceae, and Eriocaulaceae of the
World as to Valid Taxa, Geographic Distribution, and Synonymy"
reveals the fact that the states of Maine, Vermont, Massachusetts
and Idaho can be added to the states in which Verbena bracteata
Lag. & Rodr. has been found, albeit in only very scattere
localities, while V. stricta has been found in Arizona and Utah.

318 PH Y THO LO Gt A Vol. 23, mos 3

On the other hand, the extent and boundaries of geographic dis-
tribution indicated for most of the species in the Verbenaceae
herein treated appear to be largely assumed since thus far they
are not substantiated by actual herbarium specimens.

"FLORA ANALITICA E FITOGEOGRAPHICA DO ESTADO DE SAO PAULO" by
Joo Angely, Volume 3, 1--lx & )57--68) & 17 pp., illus.
1970. Volume , i--xii & 685--892 & 19 pp., illus. 1971.
Volume 5, i--xii & 893—108) pp. & 17 pp., illus. 1971.
PHYTON, Universidade de S80 Paulo, S&o Paulo, Brazil.

The first two volumes of this valuable flora were favorably
reviewed in this journal about a year ago. Now it is time to
draw attention to these three continuing volumes of prodigious
botanical work covering in all 6,326 species in 1,25 genera and
157 families. There are 1,692 geographic distribution maps and
18,472 plant names indexed.

In reference to the material on the Verbenaceae in Volume
a few items noted are: Petraea violacea in the index instead of
Petunia violacea; Lippia gehrtii, Erimus, Petrea, officinalis,
and Timotocia misspelled; Verbena melindre treated as a scientif-
ic rather than a vernacular name; the incorrect accredition of
Verbena peruviana (L.) Britton; the omission from the index of
the following Verbena species treated quite properly in the
text - V. gracilescens, V. hirta, V. hispida, V. hybrida and V.
incisa; “and the inaccurate EE NEE of Verbena eé: ephedroides as
"sphedroides". These, of course, are merely unintentional slips
in an otherwise highly valuable work.

"FLOWERING PLANTS OF JANAICA" by C. D. Adams, 848 pp., University
of the West Indies Press, Mona, Jamaica. 1972.

This beautiful tropical isle of some ,)00 square miles has
along with its crops, adventive weeds, pantropics with showy
flowers, a very fascinating and diversified native flora and un=
til now no comprehensive available botanical guide to it. The
estimated total of known native and fully naturalized flora here
is 2,888 species in 996 genera and 173 families.

Since there are no drawings included, a greater burden is
placed upon the text, but it hold up well under it.

Checking the Verbenaceae against H. N. Moldenke's recent
(1971) "Fifth Summary of the Verbenaceae, Avicenniaceae, Stilba-
ceae, Dicrastylidaceae, Symphoremaceae, iyetenene eres and Eee Erio=
caulaceae of the World as to Valid Taxa, Geographic Distribution,
and Synonymy", I find a few taxa oeenicd by him from Jamaica but
not accounted for in this flora, viz., Aegiphila martinicensis, A.
nervosa, Bouchea prismatica var. longirostra, Callicarpa cubensis,
Citharexylum pentandrum, C. spinosum, C. x perkinsi, C. x jamaicen-
se, C. fruticosum var. subvillosum, C. ~fruticosum var. -, villosum,
Glerodendrimn aculeatum var. eracile, uC angustifolium, C. indicun,

1972 Moldenke, Book reviews 319

Cornutia coerulea, C. pyramidata, Duranta repens var. alba, D.
repens var, serrata, Gmelina elliptica, Lantana arida, L. camara
var. aculeata, L. camara var, mista, L, camara var. ar. nivea, L.
camara f. parvifolia, Le L. trifolia f. hirsuta, _ Petrea volubilis var.
pubescens, Phyla nodiflora var. reptans, P._ P. strigulosa var. sericea.
Aegiphila oligoneura is treated by Moldenke as only a variety of
the widespread A. martinicensis, while Clerodendrum speciosum is
actually nothing more than a gn of C. umbellatum Poir. extreme-
ly close to the typical variety, if not actually the same; cer-
tainly not a hybrid! Lantana angustifolia is not endemic to
Jamaica because it has also been collected in Oriente, Cuba. Also,
it is very likely that Lantana involucrata does not occur on the
Gald&pagos, the plants so identified there all being L. peduncula-
ris Anderss. The Clerodendrum incisum in cultivation in Jamaica
is not the typical form, but is var. macrosiphon. The following
binomials have apparently had their orthography or accredition
"corrected" from the original: Lantana urticaefolia, Cornutia
thyrsoidea, Lantana brittoni and Clerodendrum aaa ehane Letters
are inadvertently shifted in "the year" on p. 633.

This book represents years of diligent work and will be tre-
mendously useful to botanists for years to come. Our congratula-
tions to the author!

"PHOTOBIOLOGY OF MICROORGANISMS" - Responses in the Visible Spec-
tral Region, edited by Per Halldal, x & 79 pp., illus.,
Wiley-Interscience, London, sydney, Toronto & New York, N.Y.
10016. 1970. $19.50.

This book consists of 1) papers, each in similar format start-
ing with an introductory outline and ending with a thorough bibli-
ography. At the end there are author, taxonomic and subject in-
dexes.

Photosynthesis, which is "probably the most elaborate process
for absorption and transformation of light energy" is treated as
the main subject in only two chapters because so much recent mater-
ial is already in print on it.

The topics considered are: photobiological principles and meth-
ods, photosynthetic apparatus of microalgae and its adaptation to
environmental factors, photosynthetic bacteria, organic photometab-
olism, light effects on ion fluxes and carbohydrate and protein
metabolism in algae, synchronous cultures, microorganismic photo-
motion, phototaxis and photokinesis in flagellates, light-oriented
chloroplast movements, fungal photo-responses, photobiology of
circadian rhythms in protozoa, etc., and bioluminescence in bac-
teria and fungi and marine protozoa.

There is a wealth of carefully organized and documented mater-
ial between the covers of this book!

"MICROBIAL ECOLOGY" by Martin Alexander, x & 511 pp., illus., John
Wiley & Sons, Inc., London, Sydney, Toronto and New York, N.Y.

320 PHY T’Orh OG "ik Vol. 23, no. 3
TOOTS, 197 L ete ae

"This book, as an introduction, should open the door to both
new and important problems in applied (pollution, medicine,
public health, plant pathology, soil science, food production,
limnology, oceanography, etc.) or basic (microbiology, bacteriolo-
gy, mycology, phycology, protozoology, virology, etc.) fields
creating a foundation for the reader to delve [into] more thor-
oughly or comprehend more fully those aspects of greatest interest
to him."

The book is divided into three parts. The first is about the
community and its development, considering dispersal, coloniza-
tion, succession, climax, mutrition, tolerance range, geography,
microniches and natural selection. The second is about interspe-
cific relationships involved in homeostasis, commensalism, proto-
cooperation, symbiosis, competition, amensalism, parasitism and
predation. The third part deals with the effect of microorganisms
on their surroundings in terms of biogeochemistry, animals, and
plants.

At the end the microorganisms mentioned in the text are all
listed taxonomically. There is a useful index. At the end of
each chapter pertinent literature is listed as reviews of the
topic or as other literature cited.

The exposition is in clear, simple, accurate language.

"ENVIRONMENT, POWER, AND SOCIETY" by Howard T. Odum, ix & 331 pp.,
illus., Wiley-Interscience, London, Sydney, Toronto & New
York! N.Y. 0016, “1971. > ‘$9295\.

"Intended for the general reader, this account also attempts to
introduce ecology through the energy language. Hopefully, it may
be useful to the widespread efforts under way in undergraduate
colleges to develop courses in human ecology." With other readings
this choice would be stimulating and useful. "Energy diagraming
helps us consider the great problems of power, pollution, popula-
tion, food and war free from the fetters of indoctrination." Copi-
ous, excellent, obvious diagrammatic illustrations are used
throughout. They and the text are closely integrated.

The book describes the world system with anc without highly in-
dustrialized man; the basic energy laws operating in our biosphere,
including man's high energy requirements for his machines; power
in ecological systems, for man, for order and evolution, for eco-
nomics, for politics, for a religious basis whose "God is in expo-
nential growth", electrical simulation of energy networks like
computers; and partnership with nature as our only chance for sur-
vival on "spaceship earth -—- the urban sector, the agricultural
sector, and the wilderness life-support sector" using "energy val-
ues for evaluation rather than money that belongs to only part of
the system."

Bibliography, index and an appendix with energy module formulas
are provided.

oe

; PHYTOLOGIA
. Designed to expedite botanical publication
%
ol. 23 May, 1972 No. 4

CONTENTS

_ RUDD, V. E., Reduction of Balisaea to Aeschynomene

NIINITRCEAI SRAC AEN Ch dc: nga a Seo ws en) Liles ac eee a Ra 321
"BARKLEY, F. A., & BOGHDAN, K. S., An unusual
Begonia SMR 25,2. Ata Sete t Ak seca www ie ole e 323
BOGHDAN, K. S., & BARKLEY, F. A., Stomatal patterns in the
RR CEM DD iS aia’: Uae Blain te J Mew inre: (aie my, & oa rch 327

GIBSON, D. N., Studies in American plants, IV

aoa ef MORSE, AR BL 334
: LE, M. E., Jr., Six new species of Parmelia (Lichenes)
4 MN OR Cake a acer oe ee Pam o's %5h ai ocd are ah 343
CUATRECASAS, J., Miscellaneous notes on neotropical flora, IT... .. 350
N OLDENKE, H. N., Additional notes on the genus Verbena. XIIl ..... 366

ROBINSON, H., Cheilothela vaginata and Syrrhopodon steyermarkii two
new moss species from Venezuela .............4.-+45: 390

K NG, R. M., & ROBINSON, H., Studies in the Eupatorieae (Asteraceae).

LXXL A new genus, Hatschbachiella ...............4.. 393
KING, R. M., & ROBINSON, H., Studies in the Eupatorieae (Asteraceae) .
LXXIL Notes on the genus Koanophyllon” ............... 395
KING, R. M., & ROBINSON, H., Studies in the Eupatorieae (Asteraceae).
f RAMI The genus, Ophryosporus. ©. 56.2. ee ek Fe bea ee 397
YORK

CAL Published by Harold N. Moldenke and Alma L. Moldenke
. 303 Parkside Road
| 5 1972 Plainfield, New Jersey 07060
U.S.A.

: Price of this number, $1; per volume, $7.50, in advance,
i or $8, at close of volume
2
=

REDUCTION OF BALISARA TO AESCHYNOMENE (LEGUMINOSAE)
Velva E. Rudd
Smithsonian Institution

The monotypic genus Balisaea Taubert recently caught my
attention. Upon checking the original description and the excel-
lent illustration I found the type species, B. genistoides Taub-
ert, to be identical with my Aeschynomene irwinii.

Taubert recognized that his Balisaea was related to Cyclo-
lobium and Aeschynomene but did not have fruit available to place
it in either of those genera. Since my original opinion stands,
that this taxon is referable to Aeschynomene, a new combination
is in order:

AESCHYNOMENE GENISTOLDES (Taubert) Rudd, comb. nov.

Balisaea genistoides Taubert, Bot. Jahrb. 21: 43, tab. 2,
fig. A. 1895. Type: E. Ule 2837, Brazil, Goias, "in campis
montium Serra de Balisa,"” September 1892, in flower. Holo-
type: B ? destroyed ? Isotypes ?

Aeschynomene irwinii Rudd, Phytologia 15: 117, fig. 1. 1967.
Type: H. S. “Irwin, et al. 9367, Brazil, Goias, Chapada dos
Veadeiros, ca. 13 km. N. W. of Veadeiros, 20 October 1965,
in fruit. Holotype: US. Isotype NY.

To my knowledge this unique species with its needle-like
leaflets is known only from the two collections cited above.

Taubert's original figure is here reproduced.

321

322 Pu Y £0 L0G. 1 A Vol. 23, no.

Lingler, Bot. Jahrb. XXT Bd. Tat 1.

Fig. A. Balisuea genistoides Taub Fig.B. Goyazia rupicola Taub.

J Pohi del Meisenbach Riffarth 8C° Berlin
Verlag vWilh Engelmann Leipzig

An Unusual Begonia Inflorescence

Fred A. Barkley and Kalil S. Boghdan
Northeastern University Botanical Institute and Greenhouses
35 Cambridge Street, Woburn, Massachusetts 01801

It was rather startling to run across a plant with superior
carpels in the Begonia collection. One that was remindful of the
flowers of Paeonia. It was ona plant whose stems were covered
with miniature adventitious embryos, a cultivar with the name
"Fairy" (see Plate I).

There were four flowers in the inflorescence. Two normal
pistillate flowers (see Plate II), and two teratological flowers
with separate hypogynous carpels (see Plate III).

This seemed sufficiently strange and unusual to be worthy of
recording. The family Begoniaceae has inferior ovaries, although
Hillebrandia has its perianth sufficiently far from the style
base as to have the ovary only partially inferior, but even so
the carpels are firmly united.

The only other references to comparable conditions in
Begonia seen by the writers were to a flower with several stamens
and two separate superior carpels and with a wing-like growth on
the pedicel, which was mentioned and illustrated in an editorial
in the Gardeners' Chronicle ("Hybrid Begonias", Gardeners'
Chronicle, Series 3, 2: 560. 1887.), and in a recent article by
George Kalmbacher ("Begonia Flowers", Begonian 36: 32-3%4. 1971.).

J. D. Hooker (Curtis's Botanical Magazine 122: t. 7457.
1896.) remarks that the male flowers of Begonia umbraculifera
Hooker f. are polygamous with several carpels at the center of
the ring of stamens in those flowers. His illustration shows this
condition,

323

32h Vee leh oNG AE (Ole ay (0) (6 DE .\ Vol. 23, Nae u

-—— = lem

PLATE 1. Begonia plant bearing two normal and two teratological flowers in its
inflorescence.

1972 Barkley & Boghdan, Begonia inflorescence 325

PLATE II. View of a teratological flower (in front) showing absence of
ovary, and normal flower (rear right) showing inferior ovary.

326 Posey Th O.LeOuGs Tk Vol. 23, mama

PLATE III. Face view of teratological flower showing separate superior
carpels.

Stomatal Patterns in the genus Begonia‘,

Kalil S. Boghdan and Fred A. Barkley
Northeastern University Botanical Institute and Greenhouses
35 Cambridge Street, Woburn, Massachusetts 01801

In the study of leaves of several species of Begonia it has
been observed that on the lower epidermis various stomatal
patterns exist inthe genus. This pan-tropic genus is large,
consisting of approximately a thousand species distributed among
some seventy sections of the genus.

Figure 1. Cross sections of Begonia
leaves showing the various types
of epidermal layers in the genus
and the stomatal chambers (sc).
la. B. evansiana Andrews.

lb. B. paulensis A. DC.

le. B. acetosa Vellozo.

Scale ,__;.1 m.

ee ENaEE=aSeRGPoT
1 The Begonia specimens used in this study were contributed by the
Kartuz Greenhouses, Wilmington, Massachusetts.

327

328 PHY T OLBOG I & Vol. 23, no.

As Fellerer (1892), Barkley & Boghdan (1968, 1969), and
Barkley & Hozid (1971) have recorded, the morphology and anatomy of
the Begonia leaf is quite interesting, especially regarding the
various types of epidermal layers which occur. The epidermis,
depending on the species, can be uniseriate (single upper and lower
epidermis), biseriate (double upper and lower epidermis), or
multiseriate (with three or more layers of upper and lower
epidermis) (Fig. 1).

Another interesting feature of the Begonia leaf morphology, is
the various types of stomatal patterns found associated with the
lower epidermis. Fellerer (1892) first described the stomata of
Begonia and its ontogeny. Payne (1970) described and classified
the mesogenous stoma in Begonia as helicocytic (Fig. 2a) and
described the ontogeny of this type of stoma. The stomata in
Begonia have been observed by the authors to occur in several types

vi fs all

in
re ;
'

ec ew
HH: ys” Mi) e c

~ : “4 G

i J
na (a

‘ W/ lf er
Pst 7 ot 4

Win, |
Ne
SC Rms

2a

Ww

Figure 2. Cellulose acetate
impressions of the lower
epidermis and stomata of
Begonia leaves.

2a. 3B. purpurea Sweet
showing a single stoma (st),
guard cells (gc), subsidiary
cells (sc) and epidermal
cells (ec). xX 400.

2b. B. purpurea Sweet
showing single stomata
separated by epidermal
cells. X 100.

2c. 3B. paulensis A. DC.
showing clusters of stomata
separated by epidermal
cells, X 100.

1972 Boghdan & Barkley, Stomatal patterns in Begonia 329

of patterns. One pattern, the simplest is that with each stoma
appearing by itself surrounded by subsidiary cells and separated
from other stomata by epidermal cells (Figs. la-c). If the stomata
occur in clusters (Fig. 2c), the stomatal chamber is as wide as the
cluster and as deep as the layers of epidermal cells.

Depending on the species, the leaf may have only stomata
occuring singly as illustrated in figures 2a and 2b, or in some
species there may be single stomata as well as clustered, compound
stomata, or there may be only compound stomata in clusters. Where
compound stomata occur, the number of stomata in a cluster may vary
somewhat from cluster to cluster on the same leaf.

Table 1 lists the Begonia studied, the taxonomic section of
that species, the area where the species is native, its type of
epidermis, and the number of stomata per cluster found on the lower
surface. As mentioned previously, where stomata are compound, the
number of stomata are not constant from cluster to cluster, even in
the same species. The table gives the number of stomata observed
in clusters for a species, with the predominant type of cluster
underlined. In the species observed, not a single one had stomata
on the upper surface. Plates 1 ami 2 refer to species in Table l.

Among the Begonia species studied by the authors, there
appears to be eeieelatian of stomatal patterns, number of stomata
in accluster, and the type of leaf epidermis of each particular
species. Where the upper and lower epidermis is uniseriate,

the stomata (with the exception of Begonia goegoensis N. E. Brown)
occur as single stoma, while those Begonia with a milticellular
epidermis, have the stomata compound in clusters.

330 PHY POL 06 2A Vol./23, ‘mosau
REFERENCES

Barkley, Fred A. 1972. Begomiaceae: the genera, sections and
known species of each. 26 pp. Northeastern University, Boston.

Barkley, Fred A. 1972. A Tentative Key to the Sections of the
Begoniaceae. 6 pp. Northeastern University, Boston.

Barkley, Fred A. and Kalil S. Boghdan. 1968. Begonia leaves. The
Begonian 35: 64-67.

Barkley, Fred A. and Beverly Hozid. 1971. Leaf Anatomy of
Begonia. The Begonian 38: 135-142.

Boghdan, Kalil S. and Fred A. Barkley. 1969. Begonia trichomes.
The Begonian 36: 130-132.

Fellerer, Carl. 1892. Beitrage zur anatomie und systematik der
Begoniaceen. (Dissertation.) Munich.

Payne, Willard W. 1970. Helicocytic and Allelocytic Stomata:
unrecognized patterns in the dicotyledons. Amer. J. Bot. 57: 140-
147.

331

Boghdan & Barkley, Stomatal patterns in Begonia

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ee

POH Yat) OP ONGarrs Vol. 23,5

of stomata from the lower

PLATE I. Cellulose acetate impressions
nia species in which their upper and lower epidermis are
B.

epidermis of Bego
uniseriate. (A) B. cathayana Hemsley; (B) B. rex Putzeys; (C)
(E) B. goeogensis N. E.

pustulata Liebmann; (D) B. liebmannii A. DC.;
Brown; (F) B. masoniana Irmscher. ae

333

Boghdan & Barkley, Stomatal patterns in Begonia

1972

on

Yo
we
v's

“OD

ae re ae.
een eee ee en
Uh arg cae
- ‘i ee ee
laa a .
7 CL 4 F <a ,
=e le fe
(> . ‘6 Bw 17
ad rad fi =
\ ) ¥ (f
Aw / \ | *
. ”\ ~;
ple
}
CLP» Ba A
VR ="

BE.
B.

from the lower
ower epidermis
(A)
(C) Bi
Vellozo; (F

’
B. acetosa

i Hooker f.;

iseriate epidermis (D - F).
(E

mpressions of stomata
biseriate upper and 1

ca

FOuK

and having mult
(B)
Beddome ;

nia species having
hubert ;

loccifera

Cellulose acetate i
Hooker f.

os

Bef

PLATE II.
Skan ex

epidermis of Be

miae Smith & Se

(A -C; figure 2c

olso
Lindley; (D)

venosa

STUDIES IN AMERICAN PLANTS, IV

Dorothy N. Gibson
Field Museum of Natural History

Studies in the family Gesneriaceae for the Flora of Guate-
mala resulted in the discovery of a few new species and, per-—
haps of more importance, led to a new evaluation of some close
generic relationships, requiring the transfer of some species
to other genera.

I wish to thank the curators of the U. S. National
Herbarium and the New York Botanic Garden for their kindness
in lending specimens, and especially to express my appreciation
to Mr. Conrad V. Morton, of the U. S. National Herbarium, for
his continued interest, cooperation, and helpful comments.

The work for this paper and the preparation of manuscript
for the Flora of Guatemala has been assisted by continuing
grants GB19071, GB7254, and GB27385 from the National Science
Foundation to Dr. Louis O. Williams, principal investigator.

ALLOPLECTUS Mart.

The difficulties in separating certain species of
Alloplectus and Columnea have been discussed by various
authors, and in 1969, Dr. Wm. T. Stearn, in his paper, The
Jamaican species of Coluninea and Alloplectus (Gesneriaceae),
Bull. Brit. Mus. Bot. 4: 181-236, took an important step
toward clarifying this situation by defining the two genera
much more narrowly than has been done before. By restricting
the corolla characters of Columnea to those species with "a
distinctly bilabiate red or yellow corolla, with the four
upper (posterior) lobes fused and extended into a galea, the
lower (anterior) lobe narrow and bent downwards...", and by
placing in Alloplectus those species with the corolla limb
“regular or almost so with five short rounded lobes" [Sect.
Stygnanthe (Hanst.) Hanst. and Sect. Collandra (Lem.) Hanst. |
he has provided us, at least for the Guatemalan representatives
of these genera, with a practical solution to an old and
complicated problem. It seems to me not only convenient but
highly desirable to follow his suggestions instead of clinging
to previous vague generic descriptions, especially since the
fruits of many species remain unknown. In my forthcoming
treatment of the Gesneriaceae for the Flora of Guatemala, I
am reinstating Alloplectus calochlamys Donn.-Sm. (Columnea
calochlamys Morton), and propose the following new combina-
tions:

ALLOPLECTUS VINACEUS (Morton) D. Gibson, comb. nov.
Columnea vinacea Morton, Contr. U. S. Nat. Herb. 29: 38. 1944
(type from Solola, Guatemala, Steyermark 47408).

33h

1972 Gibson, Studies in American plants 335

ALLOPLECTUS VIRIDIS (Morton) D. Gibson, comb. nov.
Columnea viridis Morton, Contr. U. S. Nat. Herb. 29: 38. 1944
(type from Alta Verapaz, Guatemala, Goll 187).

DRYMONIA Mart.
DRYMONIA PSILA D. Gibson, sp. nov.

Frutices epiphyticae interdum repentes, rami glabri;
folia late elliptica vel oblanceolata breviter acuminata
integra glabra, 9-25 cm. longa, basi cuneata obliqua; pedi-
celli 1-2, glabri, O.5-l cm. longi; bracteae parvae, incon-
spicuae, caducae; calyx glaber, lobis 5, oblongo-ovatis vel
lanceolatis acutis 1-2 cm. longis; corolla alba vel cremori-
color, interdum purpureo-maculata, glabra, 3-3.5 cm. longa,
limbus obliquus, lobis 5, crenatis, lobus anticus plus minusve
fimbriatus; antherae oblongae cohaerentes inter se, basaliter
dehiscentes; ovarium superum glabrum; stigma stomatomorphum;
capsula ignota.

Shrubby, sometimes scandent epiphytes, glabrous throughout,
the stems more or less quadrangular, often red or purplish;
leaves on petioles 1-3.5 cm. long, those of a pair subequal or
unequal, the blades coriaceous, broadly elliptic to oblanceo-
late, mostly 9-25 cm. long, short-acuminate, cuneate and often
oblique at the base, glabrous on both surfaces, paler beneath,
margins entire, lateral veins 5-7 pairs; pedicels 1-3, glabrous,
0.5-1 cm. long; bracts small and inconspicuous, linear-lanceolate,
caducous; calyx glabrous, accrescent, the 5 segments unequal,
one considerably smaller than the others, oblong-ovate to
lanceolate, acute, 1-2 cm. long; corolla white or cream-colored,
sometimes marked with purple, glabrous, 3-3.5 cm. long, the tube
a little saccate at the base, ampliate above, the limb oblique,
the 5 lobes more or less rounded, crenate, the anterior one
more or less fimbriate; anthers oblong-linear, about 3 mm. long,
united face to face, dehiscent basally, the pores uppermost
until the pollen is shed; ovary superior, glabrous; stigma
stomatomorphic; disk gland solitary; fruits unknown.

GUATEMALA: Izabal, Cerro San Gil, uppermost ridges and
summit, 1,200-1,300 meters, Steyermark 41980, type (F), (US);
Alta Verapaz, near the Finca Sepacuite, Cook & Griggs 469 (US).

Three species formerly placed in Alloplectus were found
to have anthers typical of Drymonia, appearing sagittate at
the base, and united by their inner faces with the pores (at
the base of the thecae) uppermost until the pollen is shed;
in addition, the corolla tubes of all of these species are
ampliate upward, not contracted in the throat, and the corolla
lobes are spreading, the limb oblique and more or less
bilabiate. The following new combinations are therefore
proposed:

336 POH ey OO) GL ak Vol. 23, nosey

DRYMONIA GUATEMALENSIS (Morton) D. Gibson, comb. nov.
Alloplectus guatemalensis Morton, Contr. U. S. Nat. Herb. 29:
37. 1944 (type from San Marcos, Guatemala, Steyermark 37403).

DRYMONIA MACRANTHA (Donn.-Sm.) D. Gibson, comb. nove
Alloplectus macranthus Donn.-Sm. Bot. Gaz. 31: 117. 1901 (type
from Cubilguitz, Alta Verapaz, Guatemala, Tuerckheim 7642).

DRYMONIA OINOCHROPHYLLA (Donn.-Sm.) D. Gibson, comb. nove

Alloplectus oinochrophyllus Donn.-Sm. Bot. Gaz. 54: 239. 1912
(type from Pansamala, Alta Verapaz, Guatemala, Tuerckheim 1080).

KOHLERIA Regel
KOHLERIA SKUTCHII Morton and Gibson, Spe nove

Sect. Moussonia. Frutices graciles usque ad 2-3 m. alti;
folia oblongo-ovata vel elliptico-oblonga, acuminata, serrata
vel crenata, 12-25 cm. longa, basi rotundata aut subcordata;
inflorescentiae axillares, pedunculo elongato, 7-9 cm. longo;
bracteae obovatae, foliiformes, 10-15 mm. longae; calyx
velutinus, lobis late ovatis vel obovatis acutis vel obtusis
4-5 mm. longis; corolla cinnabarina, lobis suborbicularibus
pectinato—denticulatis; stigma stomatomorphum; capsula ignota.

Slender shrubs 2-3 m. tall, the stems and branches
velutinous with multiseptate, reddish hairs; leaves opposite,
on velutinous petioles 3-7 cm. long, the blades oblong-—ovate
to elliptic-oblong, mostly 12-28 cm. long, 5-13 cm. wide,
acuminate, oblique and rounded to subcordate at base, densely
velutinous to tomentose above, velutinous below, the margins
serrate to crenate, the lateral veins 9-10 pairs; inflores-
cences axillary, the common peduncle elongate, 7-9 cm. long,
velutinous, the pedicels usually 4, sometimes 3, velutinous,
3-4 cm. long, subtended by a pair of obovate, leaflike bracts
10-15 mm. long; calyx densely velutinous, the lobes broadly
ovate to obovate, acute or obtuse, 4-5 mm. long; corolla
erect in the calyx, 3-5-4 cm. long, the tube orange-red,
velutinous outside, slightly dilated at the base but not
saccate, more or less ventricose above, 10-15 mm. wide, the
lobes yellowish, marked with red, 7-9 mm. long, suborbicular,
the margins more or less pectinate-denticulate; stamens short—
exserted, the filaments inserted on the base of the corolla
tube, pubescent, dilated at base, the anthers initially
connate, quadrate in outline; ovary densely villous; style
pubescent; stigma stomatomorphic; disk annular, the glands
much reduced, irregular and inconspicuous; capsule unknown.

GUATEMALA: Quezaltenango, 2,400 meters, Fuentes Georgi-
nas, near Zunil, Skutch 906, type (US). MEXICO: Chiapas,
2,585 meters, Pinabeto, Motozintla, Matuda 15430.

Probably most closely related to K. deppeana (Schlecht.

1972 Gibson, Studies in American plants 337

and Cham.) Fritsch and to K. elegans (Dcne.) Loesener, which
differ in their smaller leaves (the blades mostly 6-15 cm. long),
in their bracts (linear to very narrowly oblanceolate) and in
their calyx lobes (triangular and acute to subulate at the apex
in K. deppeana, linear to linear-lanceolate and attenuate to the
apex or subulate in K. elegans).

NIPHAEA Lindley

NIPHAEA SAXICOLA (Brandegee) D. Gibson, comb. nov.
Napeanthus saxicola Brandegee, Univ. Calif. Publ. Bot. 6: 65.
1914 (type from Chiapas, Mexico, Purpus 6701).

The half-inferior ovary and the subglobose anthers with
distinct thecae not confluent at the apex are characteristic
of Niphaea. The small corollas, about 6 mm. in diameter, and
the calyx lobes with more or less crenulate or denticulate
margins near the apex distinguish the species.

SOLENOPHORA Benth.
SOLENOPHORA CHIAPASENSIS D. Gibson, spe. nove

Frutices subrepentes vel ascendentes, rami juniores plus
minusve villosi; folia opposita, subaequalia vel disparia,
petiolis 1-4 cm. longis sparse villosis, laminis ellipticis
vel oblanceolatis 3-9 cm. longis, acutis vel abrupte acumina-
tis, serratis, supra sparse pubescentibus vel scabellis, infra
glabratis, praeter costas et nervos brevillosis, nervorum 7-8
paribus praesentibus; inflorescentia axillaris, pedunculus
communis obsoletus vel brevissimus et inconspicuus 3-8 mm.
longus, pedicelli solitarii vel geminati, sparse brevivillosi,
2-3-5 cm. longi; bracteae lineares inconspicuae; calyx ruber,
tubuloso-campanulatus, 8-15 mm. longus, lobis triangularibus,
4-5 mm. longi, acuti, denticulati; corolla aurantiaca, 3 cm.
longa, sparse villosa, lobis fimbriato-denticulatis; glans
disci solitaria plus minusve bilobata, apex pubens; capsula
8 mm. longa, inclusa in calyce accrescente.

Weak shrubs, subrepent or ascending to perhaps a meter
tall, branching, the older stems glabrate, the younger stems
and branches more or less villous with very short, reddish,
multiseptate, spreading hairs; leaves of a pair subequal to
unequal, on petioles 1-4 cm. long, the blades elliptic to
oblanceolate, mostly 3-9 cm. long, 1.5-4.5 cm. wide, acute or
abruptly short-—acuminate, cuneate or attenuate to the base
and often oblique, the margins serrate, sparsely pubescent to
somewhat scabrous above, glabrous below except on costae and
veins which are more or less short—villous, often red or
reddish, the lateral veins 7-8 pairs; inflorescences axillary,
the pedicels usually solitary, sometimes geminate, sparsely
short-villous, 2-3.5 cm. long, subtended by a pair of small,
inconspicuous, linear bracts, the common peduncle obsolete or
if present, inconspicuous, 3-8 mm. long; calyx rose-red or

338 PLY 20-2011 & Vol. 23, no.

PLATE

Solenophora chiapasensis. A, habit; B, pedicellate
flower; C, corolla opened to show stamens and staminodes;

D, calyx opened to show bilobate disk gland and style.

1972 Gibson, Studies in American plants 339

reddish, tubular-campanulate, 8-15 mm. long, almost glabrous or
thinly villosulous, more densely so near the base, the lobes
triangular, mostly 4-5 mm. long, acute, denticulate; corolla
orange, about 3 cm. long, sparsely villous or almost glabrous,
the limb to 1.5 cm. wide, the lobes semiorbicular, 3-5 mm. long,
the margins fimbriate-pectinate, usually conspicuously so, and
often more or less ciliate; stamens about equalling the corolla
or slightly exserted, the anthers 1.5-2 mm. long, united by
their tips; disk gland solitary, large, very shallowly and
irregularly bifid or bilobate, more or less pubescent, at least
near the apex; style pubescent; stigma stomatomorphic; capsule
about 8 mm. long, enclosed in the accrescent calyx; seeds
broadly fusiform.

MEXICO: Chiapas, Municipio of Tenejapa, 9,100 feet,
Breedlove 15187, type (F); Shilom Ton 1768; 6,600 feet,
Breedlove 10741; 7,600 feet, Breedlove 6996.

Previously identified as S. tuerckheimii Donn.-Sm. which
differs in its larger leaves (mostly 8-15 cm. long with 8-10
pairs of lateral veins), its subumbelliform to cymose inflor-
escences with peduncles mostly 3-8 cm. long, and its short
calyx lobes (mostly 1-2 mm. long, very rarely to 3 mm.) that
are very broadly and irregularly subtriangular and obtuse to
nearly rounded rather than acute.

SOLENOPHORA MACULATA D. Gibson, spe. nov.

Frutices vel herbae suffruticosae; folia late elliptica
vel ovato-elliptica 11-25 cm. longa acuminata, duplicato—
serrata, supra sparse villosa, infra glabrata; pedunculus
communis 1-2 cm. longus, pedicelli 2-3 cm. longi, bracteati;
calyx 2-2.5 cm. longus, lobis triangularibus denticulatis;
corolla lutea vel aurantiaca intus purpureo-maculata 2.5-3 cm.
longa, tubo villoso; glans disci solitaria, plus minusve
bilobata.

Shrubs or suffruticose herbs, simple or sparsely
branched, the young stems reddish, usually villous, later
more sparsely villous or glabrate; leaves of a pair subequal,
on glabrous petioles 5-10 cm. long, the blades broadly
elliptic or ovate-elliptic, mostly 11-25 cm. long, 8-14 cm.
wide, short-acuminate, oblique at base, sparsely short-
villous above, glabrate beneath, the costae and veins rose-
red, the margins doubly serrate, the lateral veins 9-12
pairs; inflorescence axillary, the common peduncle 1-2 cm.
long, sparsely pubescent or glabrate; bracts oblong-—lanceo-
late or oblong-elliptic, O.5-1 cm. long; pedicels glabrous,
2-3 cm. long; calyx suffused with rose or maroon, sparsely
villous near the base, glabrous above, 2-2.5 cm. long, the
triangular lobes unequal, 2-4 mm. long, more or less denticu-
late; corolla 2.5-3 cm. long, deep yellow or orange-yellow,
spotted or striped with maroon within, the tube erect,

340 Pony TOM! Ged 1k Vol. 23, now w

ampliate above, about 1 cm. broad below the lobes, not con—
stricted, villous outside, the lobes about 4 mm. long, irregu-
larly undulate; stamens exserted, the anthers about 2 mm. long,
united by their tips; style exserted; disk gland solitary, more
or less bilobate.

GUATEMALA: Between Todos Santos and Finca El Porvenir,
middle slopes of Volcan Tajumulco, San Marcos, 1,300-3,000

meters, Steyermark 36992, type (F).

This species may be distinguished from the: other large-
leaved species in our area by its much shorter corollas. It
is similar in many ways to S. pirana Morton, but that species
has longer peduncles (commonly 5-10 cm. long), and the corollas
of S. pirana are not only longer (4-5 cm.) but are not spotted
nor marked within.

SOLENOPHORA OBLIQUA Denham and Gibson, spe nov.

Frutices vel arbores debiles, usque ad 6 m. altae, rami
sparse pubescentes vel glabrati; folia opposita, petiolis
glabris, 3-12 cm. longis, laminis 14-30 cm. longis, oblongo-—
ovatis vel elliptico-oblongis acuminatis vel acutis duplicatis,
serratis, supra sparse pilosis, infra glabratis vel glabris;
inflorescentia cymosa, pedunculus communis ad 2 cm. longus,
pedicelli O.5-3 cm. longi, bracteati; calyx glaber, 3.5-4 cm.
longus, plus minusve bilabiatus urceolatus, lobis denticulatis;
corolla 6.5-7.5 cm. longa, glabra aurantiaca, limbus ca. 3 cm.
latus, lobi intus purpureo-maculati, fimbriato-denticulati;
glans disci solitaria bilobata, pubens; calyx multo longior
quam capsulae

Shrubs or weak trees to 6 m. tall, the stems and branches
glabrate or sparsely, bifariously pubescent with multiseptate
hairs; leaves opposite, subequal, the petioles essentially
glabrous, mostly 3-12 cm. long, the blades oblong-—ovate to
elliptic-oblong, acuminate or acute, cuneate or rounded at
the base and usually very oblique, mostly 14-30 cm. long, 7-
17 cm. wide, sparsely pilose above, nearly glabrous beneath,
the margins doubly serrate, the lateral veins 10-14 pairs;
inflorescences axillary, cymose, the common peduncle 1-2 cm.
long, glabrous (sometimes the flowers solitary, the peduncle
then much reduced and inconspicuous), pedicels 0.5—3 cm. long,
glabrous, bibracteate, the bracts leaflike, mostly 2-3 cm.
long, sometimes caducous in age; calyx glabrous, brownish or
green with orange-brown stripes, 3.5-4 cm. long, accrescent,
3-4 times longer than the ovary, conspicuously urceolate in
bud, the limb narrowly contracted, oblique, more or less bi-
labiate, the 5 lobes irregular, short, 3-8 mm. long,
triangular to lanceolate, appearing irregularly denticulate
but the lobules actually rounded and obtuse, sometimes
minutely papillate within, the tube in anthesis becoming
deeply cleft on one side for about 2/3 of its length, the

1972 Gibson, Studies in American plants 341

calyx then appearing narrowly tubular-campanulate; corolla 6.5-
7.5 cm. long, glabrous or nearly so, erect in the calyx, yellow-
ish-orange to reddish-orange outside, deep yellow to orange and
sometimes marked with maroon within, the tube gradually ampliate,
the limb somewhat oblique, about 3 cm. wide, the 5 lobes broadly
suborbicular, spotted with purple or maroon within, the margins
rounded and more or less fimbriate-pectinate; stamens 4, didyna-
mous, short-exserted, the glabrous filaments inserted on the
base of the corolla tube, the anthers about 3 mm. long, initially
united by their tips, the thecae oblong, fully dehiscent; a
staminode usually present; ovary inferior; style densely pubes-—
cent; stigma stomatomorphic; disk gland solitary and usually
deeply bilobate, or 2 glands connate at the base, pubescent;
capsule far surpassed by the calyx tube, rupturing irregularly
near the middle when mature, containing numerous, minute,
reddish-brown, shining, ellipsoidal to fusiform seeds.

GUATEMALA: Quezaltenango, western slopes of Volcan Zunil,
opposite Santa Maria de Jesus, alt. 1,500 m., Steyermark 35182,
type (F); between San Martin Chili Verde and Colomba, alt.
1,800 m., Standley 85478 and 85523; El Pocito, south of San
Martin Chili verde, on road to Colomba, alt. about 2,200 m.,
Standley 85049; between Quebrada Chicharro and Montana Chi-
charro, on southeast-facing slope of Volcan Santa Maria, alt.
1,300-1,400 m., Steyermark 34329. Chimaltenango, Johnson 1785.
San Marcos, wet mountain forest near Aldea Fraternidad, west—
facing slope of the Sierra Madre Mountains, alt. 1,800-2,400 m.,
Williams, Molina, & Williams 26209 and 26251.

Differs from S. purpusii Brandegee in its longer, more
narrow, urceolate calyx.

SOLENOPHORA TOUCANA Denham and Gibson, sp. nove

Frutices usque ad 4 m. altae, rami et petioli plus minusve
pilosi; folia opposita oblongo-ovata vel oblongo-elliptica,
16-30 cm. longa acuminata, duplicato-serrata, supra sparse vel
dense pubescentia vel pilosa, infra pilosa non nisi in costis
et nervis tantum; inflorescentia cymosa, pedunculus communis
ad 5 mm. longus, pedicelli 5-15 mm. longi, bracteati; calyx
campanulatus, 1.5-2.5 cm. longus, pilosus, fere truncatus,
lobi denticulati; corolla aurantiaca 5.5-7 cm. longa, pilosa,
limbus 3-3.5 cm. latus, lobi intus rubro-maculati fimbriato-
denticulati; glans disci solitaria bilobata pubens; capsula
ignota.

Shrubs 2-4 m. tall, the stems and branches more or less
pilose with multiseptate hairs; leaves opposite, subequal, on
pilose petioles mostly 5-12 cm. long, the blades oblong-ovate
to broadly oblong-elliptic, mostly 16-30 cm. long, usually
acuminate, rarely acute, obliquely cuneate to nearly rounded
at the base, sparsely or densely pubescent to pilose above,

32 PeeyY TOL 0G Tuk Vol. 23, no.

pilose beneath on costae and veins, otherwise pubescent or
glabrate, the margins doubly serrate, the lateral veins about
10 pairs; inflorescences axillary, cymose, the common peduncle
seldom more than 5 mm. long, more or less pilose, the pedicels
5-15 mm. long, pilose, bibracteate, the bracts linear, ciliate;
calyx urn-shaped in bud, becoming campanulate and 1.5-2.5 cm.
long at anthesis, about 3 times longer than the ovary,
accrescent, more or less pilose, densely so near the base,

very shallowly lobate and appearing almost truncate, the margins
unevenly denticulate; corolla orange outside, yellow within,
5-5-7.5 cm. long, erect in the calyx, the tube gradually
ampliate, pilose outside, the limb 3-3.5 cm. in diameter, the
lobes suborbicular, broadly rounded, spotted with red within,
the margins minutely fimbriate-denticulate; stamens 4, didyna-
mous, short-exserted, the filaments usually sparsely pubescent,
inserted on the base of the corolla tube, the anthers united
by their tips, the thecae fully dehiscent; staminode present;
ovary inferior; the style exserted, densely pubescent; stigma
stomatomorphic; disk gland solitary and usually deeply bilobate
or 2 glands connate at the base, pubescent; capsule unknown.

EL SALVADOR: Depto. Chalatenango, banks of Rio Sumpul
at boundary between El Salvador and Honduras, alt. ca. 2,250 m.,
Tucker 1172, type (UC); (F, NY, US, PH). HONDURAS: Depto. La
Paz, 2,100 m., Molina y Molina 14072; Depto. Ocotepeque,
1,800 m., Molina 22359. GUATEMALA: Depto. Guatemala, Palencia,
1,480 m., Morales 947.

The only large-leaved species in our area with a nearly
truncate calyx limb.

SIX NEW SPECIES OF PARMELIA (LICHENES) FROM AFRICA

Mason E. Hale Jr.
Smithsonian Institution, Washington, D.C. 20560

1. Parmelia chowoensis Hale, sp. nov. Fig

Thallus adnatus, corticola, coriaceus, albocinereus, 4-6 cm
latus, lobis subirregularibus, 2-2.5 mm latis, contiguis vel sub-
imbricatis, margine bulbato-ciliatis, parce inflatis basi, super-
ne planus, nitidus, emaculatus, sorediis isidiisque destitutis,
cortex superior 14-18 crassus, stratum gonidiale 15-10u crassum,
medulla alba, 70-90u crassa, cortex inferior 12u crassus, subtus
castaneus, modice rhizinosus, rhizinis castaneis, simplicibus.
Apothecia numerosa, subpedicellata, 2-4 mm diametro, amphithecio
crenato, disco imperforato, sporis octonis, 3-4 x 6-7.

Holotype: Zambia: Chowo Forest, Nyika Plateau, collected by
Memeika Jellicoe, no. 53, April 1969 (BM; isotype in US).

Chemistry: Cortex K+ yellow (Atranorin), medulla K-, P+
red, C- (protocetraric acid).

This small species in section Bicornuta is characterized by
the pale lower surface and the presence of protocetraric acid.
Only one other species, P. subinflata Hale from the Philippines,
produces this acid, and it too has weakly inflated bulbate cilia.
Another African species, P. hypocraea Vainio, is superficially
close but has distinct bulbae and salazinic acid.

2. Parmelia composita Hale, sp. nov. ride

Thallus expansus, corticola, coriaceus, laxe adnatus vel
suberectus, viridi-cinereus, 8-15 cm latus, lobis rotundatis,
5-15 mm latis, margine ciliatis, ciliis 1-3 mm longis, superne
planus, nitidus, valde albomaculatus, aetate rimosus, cortex su-
perior 18-20u crassus, stratum gonidiale 20u crassum, medulla al-
ba, 130-15Qu crassa, cortex inferior 15u crassus, subtus niger
vel marginem versus castaneus vel albo-variegatus, centro rhizi-
nosus, rhizinis sparsis, simplicibus vel furcatis, longis, mar-
gine nudus. Apothecia pedicellata, 3-15 mm diametro, disco per-
forato, sporis octonis, 10-12 x 22-24y, episporio 2u crasso.

343

34 PHY TOLOG IA Vol. 23, no. 3

Holotype: Tanzania: 2 km N of Kitoto Camp, Mt. Meru, E.
slope, Arusha Prov., elev. 2500 m, collected by R. Santesson, no.
22985a, 7 Jan. 1971 (UPS; isotype in US).

Chemistry: Cortex K+ yellow (atranorin), medulla K+ red, P+
orange, C+ rose, (gyrophoric and norstictic acids).

Additional specimens examined: Same locality: Santesson
21594a, 22850, 23069c (UPS); 21594a, 22886b (UPS, US).

This Amphigymnia species appears at first to be related to
Parmelia perforata (Jacq.) Ach.), and American species which also
has norstictic acid. P. composita, however, produces in addition
gyrophoric acid, the first instance of the joint occurrence of
these two acids, as far as I am aware, and has a darker mottled
margin below rather than the distinct pure white rim of P. per-
forata. It also has larger spores and the less dense but strong

maculation more typical of P. pseudonilgherrensis Asah.

3. Parmelia enormis Hale, sp. nov. Fig.03

Thallus expansus, enormis, 10-30 cm latus, coriaceus, saxi-
cola, cinereo-albidus, lobis sublinearibus, centro imbricatis,
5-8 mm latis, margine integris vel axille ciliatis, Superne plan-
us, nitidus, emaculatus, sorediis isidiisque destitutis, cortex
superior 20-22u crassus, stratum gonidiale 30-40y crassum, con-
tinuum, medulla alba, 130-150u crassa, cortex inferior 16-20u
crassus, subtus pallide castaneus, dense rhizinosus, rhizinis
pallidis, elongatis, simplicibus. Apothecia numerosa, subpedi-
cellata, 3-8 mm diametro, disco imperforato, sporis octonis, 6-7
x 8-1]p.

Holotype: Zambia: On granite rocks, Zambia Rest House, Nyi-
ka Plateau, elev. 7600 ft., collected by Memeika Jellicoe, s.n.,
September 1968 (BM; isotypes in TNS, US).

Chemistry: Cortex K+ yellow (atranorin), medulla K+ red,
P+ orange, C- (salazinic acid).

The presence of cilia in the axils of this most remarkable
lichen places it in section Imbricaria, although superficially,
because of the linear lobes, it seems to be a Hypotrachyna species.
P. usambarensis Stein. & Zahlbr., another very large linear-
lobed, saxicolous species in Africa, differs in having isidia and

1972 Hale, New species of Parmelia 345

a black lower surface.

4. Parmelia pachydactyla Hale, nom. et stat. nov. Fig. 4

Parmelia caperata var. isidiophora Steiner, Stiz. K. Akad.
Wiss. Wien Math.-Naturw. Cl. 106:215. 1897.

Parmelia steineri Dodge, Ann. Mo. Bot. Garden 46:125. 1959.
Based on P. caperata var. isidiophora Steiner [not P.
steineri Gyelnik, Ann. Mycol. 36:289. 1938 (=P. molli-
scula Acharius)].

Holotype: Kenya: Athi Plains, Liechtenstein s.n. (WU).

Chemistry: Cortex K- (usnic acid), medulla K-, P+ red, C-,
(protocetraric acid).

Additional specimen examined: S. Rhodesia: Zimbabwe, Div.
Victoria, L Kofler, 27.6.1963 (LD,US).

The type of this saxicolous lichen is rather fragmentary but
the discovery of a better specimen by Kofler in Rhodesia has
provided good material for a fuller understanding of the species.
The lobes are quite adnate and rather narrow (1-2 mm) and the
"isidia" scattered and very thick (about .3 mm wide and .5 mm
high). It would appear to be related to the pantemperate P.
caperata (L.) Ach., a larger species with soredia. P. rudis
Harm. from Indochina has the same chemistry but the isidia are
much finer.

5. Parmelia subschimperi Hale, sp. nov. Figs. 5

Thallus laxe adnatus, corticola, 6-12 cm latus, viridi-cin-
ereus, lobis rotundatis, 10-18 mm latis, margine ciliatis, ciliis
1-2.5 mm longis, sorediatis, soraliis linearibus vel pro parte
irregularibus et submarginalibus, superne planus, nitidus, crasse
albo-maculatus, cortex superior 15-18u crassus, stratum gonidiale
20-25u crassum, medulla alba, 100-120 crassa, cortex inferior
12-14u crassus, subtus niger, sparse rhizinosus, rhizinis simpli-
cibus, ambitu nudus, castaneus. Apothecia ignota.

Holotype: Kenya: Bamboo zone, National Park Road, W. slope,
Mt. Kenya, elev. 2700-3100 m, Nanyuki Distr., Central Prov. col-
lected by R. Santesson, no. 22150 (UPS; isotype in US).

3h,6 PAE ide vO) Te O 1G eek VoL... 23, , newa3

Chemistry: Cortex K+ yellow (atranorin), medulla K-, P-,
C-, KC+ rose (norlobaridone).

Additional specimens examined: Kenya: Mt. Meru, Arusha,
Prov., Santesson 21583a (UPS), 22911b, 22912a, 22943 (UPS, US);
same locality as holotype, Santesson 22042b, 22173 (USP, US).

As the name implies, P. subschimperi (subg. Amphigymnia)
is related to P. schimperi Mull. Arg., which may in fact be con-

sidered its nonsorediate counterpart. P. schimperi has similar
strong white maculae and a dark lower surface, even toward the
marginal zone. Norlobaridone is now known to be rather common

in Amphigymnia (Kurokawa, Jour. Hattori Bot. Lab. 32:205-218.
1969). One other norlobaridone-containing species in Africa,

P. hababiana Gyelnik, could be confused with P. subschimperi, but
it has smaller more membranous lobes, uniformly short cilia,
smaller maculae, and a more or less distinct white or mottled
zone below at the margin. P. pseudonilgherrensis Asah. is very
close in overall morphology but contains alectoronic acid.

6. Parmelia xanthoparmelioides Hale, sp. nov. Fig. 6

Thallus laxe adnatus, saxicola, viridi-flavicans, 4-8 cm
latus, lobis linearibus, dichotome ramosis, 1.5-2.5 mm latis,
eciliatis, superne planus, nitidus, emaculatus, apicem versus
granulato-sorediatus, sorediis crassis, pro parte subfatiscenti-
bus, cortex superior 12-]l4u crassus, stratum gonidiale 20-25y
crassum, medulla alba, 100-130u crassa, cortex inferior 12-14u
crassus, subtus niger, modice vel sparse rhizinosus, rhizinis
tenuibus, dichotome divisis. Apothecia ignota.

Holotype: South Africa: Cliffs on hills NW of Woodhead Res-
ervoir, Table Mtn., Cape Prov., collected by N. S. Pillans 3857,
February 1920 (BM; isotypes in LD, US).

Chemistry: Cortex K- (Usnic acid), medulla K+ red, P+ orange,
C-(salazinic acid).

Were it not for the presence of dichotomously branched
rhizines, this species would appear to be in the P. stenophylla
group of subgenus Xanthoparmelia. As is the case with P. sinuosa
(Sm). Ach., another yellow species formerly placed in Xantho-
parmelia, the correct sectional disposition is better made accord-
ing to rhizine characters. A very similar rare species from

1972 Hale, New species of Parmelia 3,7

tropical America, P. meyeri Zahlbr., also has usnic and salazinic
acids but is smaller, closely adnate, and has more pustulate
soralia.

Fig. 1. Isotype of Parmelia chowoensis CUS}. Fig. 2:
Isotype of P. composita (US). Scale in mn.

348 PB Yat. E0iG yk Vol. 23, no. 4

Fig. 3. Isotype of Parmelia enormis (US). Fig. 4. Parmelia
pachydactyla (Kofler, US). Scale in mm.

1972 Hale, New species of Parmelia 39

WEE

Fig. 5. Isotype of Parmelia subschimperi (US). Fig. 6.
Isotype of P. xanthoparmelioides (US). Scale in mm.

MISCELLANEOUS NOTES ON NEOTROPICAL FLORA, II.

by Jose Cuatrecasas
Department of Botany, Smithsonian Institution

These notes are descriptions of new taxa extracted from a
paper which for a long time is in the hands of the publisher of
another journal. They are preliminary, partial studies to be
supplemented in forthcoming monographs now in preparation.
Genera involved in these notes are Diplostephium, Oritrophiun,
Vernonia and Espeletia (Compositae), all from Andean collections.
The work has been partially sponsored by the National Science
Foundation.

DIPLOSTEPHIUM RITTERBUSHIIT Cuatr. sp. nov.

Frutex parvus 30 cm ad 1 m altus, caule circa 8 mm diametro
a basi ramoso ramis robustis erectis tuberculato cicatricosis
parce ramulosis ramulis extremo dense foliosis glandulosis
glutinosis et hirsutulo lanuginosis, pilis rigidiusculis inaequa-
liter furcatis 1-3 mm longis intricatis denique deciduis tectis.

Folia alterna coriacea rigida sessilia subelliptico-oblonga
ad basim paulo angustata apice attenuata subacutaque margine
incrassati-revoluta, 12-18 mm longa 4-6 mm lata; supra glabra
viridia nitidula copiose minute impresse glanduloso-punctata
tantum costa paulo visibili; subtus densiuscule villoso-lanugio-
nosa pilis longis paulo flexuosis intricatis tecta sed costa
crassa prominenti glabraque basi magis dilatata bene conspicua.

Capitula quinque subterminalia congeste racemosa ex foliis
subtendentibus brevioribus angustioribusque axillaria. Pedicelli
ecrassiusculi hirsuto-sublanati usque ad 1 cm longi. Capitula
radiata tubuloso-campanulata circa 12 mm longa 6 mm diametentia,
circulo ligularum 14 mm diametro. Involucrum villoso-lanugino-
sum 8-9 mm altum, phyllariis 4eseriatis exterioribus lineari-
lanceolatis 5-5.5 x 1-1.2 mm extus longe piloso-lanuginosis et
sparse glandulosis, interioribus linearibus acutis dorso glandu-
losis margine ciliatis subapicem pilosis, 7.5 x 0.8-1 m.
Receptaculum 3 mm diam alveolatum marginibus alveolarum dentatis.

Flores radii feminei ligulati circa 23 in capitulo. Corolla
alba 8 mn, tubo 3-3-3 mm pilosulo pilis pluricellularibus sub-
elaviformibus ad 0.15 mm longis sursum densioribus, lamina
lineari 4.7-5 mm longa 0.8-1 mm lata 2-3 dentata glabra vel supra
sparsissimis pilis minutis adpressis. Stylus circa 5 mm longus.
Ovarium 2-2.2 mm longum obovoideo-oblongum sursum sparsis vel
sparsissimis minutis glandulis vel pilis praeditum. Pappus he
4.2 mm setis exterioribus tantum 0.5-2 (-3) m acutis, interior-
ibus sursum leviter dilatatis.

Flores disci pseudohermaphroditi circa 23 in capitulo.
Corolla lutea 5 mm longa tubo 2.5 mm pilis subclaviformibus

350

1972 Cuatrecasas, Notes on neotropical flora 351

minutis circa 0.1 mm longis copiosis ornato, limbo 2.4-2.5 mm
inferne contracto pilosoque dentibus 5 interdum 4 triangularibus
acutis conniventibus glabris 0.5 mm. Antherae 1.5 mm longae
basi minute auriculatae appendice oblonga obtusiuscula glandu-
loso=-punctata. Filamenta uniformia stricta glabra. Stylus
corollam excedens extremo sublanceolatus piloso-papillosus apice
breviter bifidus. Ovarium lineare 4 mm longum sparsis minutis
glandulis et raris minutis pilis munitum. Pappus 4 m longum
setis exterioribus 1-2 mm strictis acutis interioribus ad apicem
paulo dilatatis.

Typus: Colombia, Huila: West slope below Pico Norte of
Nevado del Huila, 13,800 feet altitude (range is 13,600-14,000),
very abundant, collect. Jan. 11, 1970 by Philip C. Ritterbush,
8.n. Holotypus, US.

Species named for Dr. Philip C. Ritterbush, scientific
historian and scholar, also alpinist and explorer who led two
expeditions to the abrupt Nevado del Huila. During his second
expedition in 1970 he escalated the highest peak, the Pico Mayor,
jointly with Steve Salva and Cary Ullin. See Ritterbush,
Conquest of Huila, rev. Americas 22(1):19-25 and frontispice,
Jan. 1970, and Report of the 1970 Expedition to the Nevado del
Huila, Central Andean Cordillera, Colombia. Occasional Publ.
Mus. Dep. Hist. Nat. Cali, Colombia, 1970.

DIPLOSTEPHIUM CAMARGOANUM Cuatr. sp. nov.

Frutex ramis foliosis teretibus dense adpresseque cinereo-
crispoianatis.

Folia alterna coriacea rigidula. Petiolus 6-12 mm longus
rigidulus adpresse crispo-lanatus supra sulcatus subtus teres
basi ampliatus amplectens. Lamina oblongo-elliptica utrinque
attenuata apice acuta vel subacuta basi cuneata margine integra
plana, 4-8 cm longa 1.5-3.2 cm lata; supra viridis nitens plus-
minusve viscosa, sparsis pilis malpighiaceis vel inaequaliter
furcatis, vel excepto costa impressa minute pilosa glabrata,
nervis secundariis filiformibus venulis reticulatis paulo promi-
nulis; subtus cinerea dense adpresseque crispo lanata, costa
eminenti, nervis secundariis minus prominentibus 7-9 utroque
latere subpatulis et arcuato-anastomosatis, venulis reticulo
parum visibili instructis.

Inflorescentiae corymboso-paniculatae subterminales folia
suprema valde excedentes, 8-12 cm longae 2-6 cm latae, axe
elongato modice tenui sed recto erectoque adpresse crispulo-
lanato cinereo nudo, tantum superne ramis ramulisque parcis
subfastigiatis cinereo-lanatis, ultimis capitula 2-3 breviter
pedicellata vel singula ferentibus instructo. Bracteae
bracteolaeque subtendentes ovatae subobtusae vel subacutae
dorso lanatae vel glabratae 1.5-0.5 mm longae. Pedicelli
ecrispo-lanati 1-10 mm longi tantum apice parcissimis bracteolis
minimis capitulum adjacentibus.

Capitula brevissime radiata circa 10 mm longa 2.5-3 mm
lata, 18-24 flores involucrum valde excedentes ferentia.

352 PHYTO LO GMA Vol. 23, noo l

Involucrum tubulosum 5.4-6 mm altum, phyllariis 5-6-seriatis
imbricatis straminosis rigidisque margine anguste hyalino-
membranaceis plus-minusve eroso-ciliatis extus parce sparseque
papillosis tantum infimis breviter pilosis dorso sursum
brunnescentibus ceterum stramineis, interioribus 3.6-4 x 0.8-
1.2 mm elliptico-oblongis obtusiusculis medianis ovato-oblongis
3-3.3 x 1.2-1.3 mm exterioribus ovatis 2-1.5 x 1.3 mm intermediis
gradatim transeuntibus. Receptaculum 1.4 mm diametro alveolatum
marginibus alveolorum membranaceis.

Flores radii feminei 7-10 in capitulo. Corolla 3.5-4 mm
longa minute ligulata, tubo recto crassiusculo supra medium
minute puberulo pilis antrorsis tenuibus circa 0.2 mm longis,
lamina 0.7-0.8 m longa 3-dentata dentibus triangularibus 0.3-
0.4 mm raro bidentata, saepe labio adaxiali lineari circa 0.5 mm
longo interdum bifido saepe minutum dentem reducto. Stylus 3.5
mm longus crassiusculus basim dilatatus, ramis 0.5 mm longis
lineari-subulatis marginato-stigmatiferis corollam non excedent-
ibus. Ovarium 2.2-2.5 mm longum fertile oblongo-obovoideum
obtuse tricostatum paulo compressum pubescente pilis geminis
antrorsis rectis acutis circa 0.3 mm longis. Pappus albidus
circa 3.5 mm longus setis biseriatis strigulosis acutis paulo
inaequalibus parcis brevioribus 2-2.5 mm sursum levissime
dilatatis.

Flores disci pseudohermaphroditi 11-14 in capitulo. Corolla
4-4.5 mm longa parte media dilute pubescens pilis tenuibus circa
0.2 mm longis, tubo 1.8-2 m longo, limbo anguste infundibuli-
formi lobis 5 oblongis acutatis marginatis 1 mm longis glabris
apice papilloso. Antherae 1.3 mm longae basi auriculatae apice
appendice triangulari-oblonga acuta O.4 mm longa. Stylus 4.5
mm longus basi incrassata cum annulo nectarifero cincta, ramis
anguste lanceolatis 0.8 mm longis extus dense antrorseque
papilloso-pilosis. Ovarium lineare 3 mm longum tricostatum
sterile vacuum, densiuscule pubescente pilis geminis antrorsis
circa 0.3 mm longis. Pappus 3.5 mm longus albidus, setis com-
planatis strigulosis subaequilongis sursum parum ampliatis
interdum parcis brevioribus.

Typus: Colombia, Boyaca: Arcabuco, abrededores de la
poblacion 2739-2850 m alt, 20-X-1965 col. G. Huertas & Luis A.
Camargo 6309. Holotypus US.

DIPLOSTEPHIUM ASPLUNDII Cuatr. sp. nov.

Frutex ramis terminalibus copiose ramulosis fusco-brunneis
argute striolulatis hirsutis, pilis ad circa 2 mm longis plus-
minusve flexuosis acutis ochraceis. Ramuli ad ramos racemose
dispositi dense foliosi et copiose indumento lanuginoso-
hirsutulo praediti, ad extremos vestimentum densius.

Folia alterna copiosa et patula in ramulis, modice cori-
acea. Petiolus circa 2 m longus fere planus subtus lanugino-
sus basi amplectens. Lamina 6-8 x 3-2.5 mm, elliptica vel
oblongo-elliptica utrinque attenuata apice acutata mucronulata-
que margine bene revoluta; supra viridis nitidula glabra laevis

1972 Cuatrecasas, Notes on neotropical flora 353

vel leviter undata tantum costa depressa notata; subtus fusco-
ochracea dense lanata costa incrassata tantum conspicua.

Capitula radiata solitaria sessilia ramulos foliosos
terminantia, 12-14 mm alta, 23-37 flores ferentia, circulo
ligularum 30-35 mm diametienti. Involucrum 9-9.5 mm altun,

5 mm diametro, phyllariis 5-seriatis papyraceis stramineis
lilacino-lineatis maculatisque, margine anguste scarioso, extus
breviter, albido-pubescentibus pilis flexuosis subadpressis,
interioribus lanceolato-linearibus acutis 7-8 x 1 m, exterior-
ibus lanceolatis 5-3.5 x 1-0.7 mm, magis pubescentibus.
Receptaculum 15 mm diametro marginibus alveolarum membranaceis.

Flores radii feminei ligulati 14-21 in capitulo. Corolla
violacea 19-20 mm longa, tubo 4 mm ad apicem parcis minutis
pilis, lamina lineari-oblonga obtusa minute 3-dentata 4-nervata.
Stylus 7 m, ramis linearibus strictis 2 mm longis. Ovarium
2.5 mm longum oblongum ad basim paulo attenuatum cum basi
callosa 5-costatum copiose glandulosum, glandulis lacrimiformi-
bus sessilibus vel minute pediculatis. Pappus 7.5-8 mm longus
rubescenti-stramineus setis inaequilongis strictis strigosis
exterioribus dimidio vel tertio brevioribus.

Flores disci hermaphroditi 9-16 in capitulo. Corolla
olivaceo-lutescens circa 8 mm longa; tubo 2.8 mm apice parcis
pilis flexuosis 0.2 mm; limbo tubuloso-ampliato basi parcis
minutis pilis, dentibus 1.4-1.5 mm longis oblongis acutis
incrassato-marginatis apice papilloso, extus sursum copiosis
glandulis ad extremo saepe parcis pilis sericeis tenuissimis
flexuosis 1-6 mm longis praeditis. Antherae 2.5 mm basi
sagittatae appendicibus apicalibus oblongis attenuatis O.7 mm
longis. Stylus 10-10.5 mm, ramis 1 mm lineari-lanceolatis extus
dense papilloso-hirsutis. Ovarium 4-3.6 mm 5-nervatum copiose
glandulosum, basi parcis vel parcissimis pilis tenuibus circa
0.5 mm. Pappus 8 mm longus, setis sursum leviter ampliatis,
exterioribus circa 1 mm brevis vel 2-3 mm.

Typus: Ecuador, Tungurahua, Cordillera de Llanganates near
Las Torres, 3500 m. alt, rays violet, disc corollas olivaceous-
yellow, coll. Erik Asplund 9831. Holotypus, S.

DIPLOSTEPHIUM SAGASTEGUII Cuatr. sp. nov.

Frutex valde ramosus, ramis robustis adpresse lanatis
denique gradatim glabratis, teretibus rugulosis, terminalibus
crasse densissimeque albo-crispo-lanatis, ramusculis floriferis
conferte foliosis tam crasse spisseque albo-lanatis 2-5 ecm
longis racemose dispositis numerosis congestis instructis.

Folia parva visu sessilia sed intra vestimentum bene
petiolata. Petiolus planus membranaceus saepe trinervatus
3-3-5 x 1-1.2 mm, basi cum vagina circa 0.5 mm elongatus,
adaxiale glaberrimus abaxiale dense crasseque lanatus. Lamina
coriacea marginibus revolutissimis argute conchata visu ovata
obtusa basi rotundata seu subcordata 2.5-3 x 1.8-2.2 mm, supra
viridis nitidissima enervata sparsis pilis verrucoso-conicis
subspinulosis minutissimis praedita, subtus cavitate infra

35h P Pee POLO Glick Vol. 23, nowy

margines densissima lana alba repleta, etiam petiolo ascendenti
cum vestimento crasso occulto.

Capitula radiata terminalia singula sessilia in extremis
ramusculorum, 40-50 flores ferentia, circa 12 mm alta circulo
ligularum 15 mm diametienti. Involucrum late tubulosum circa
9 x 5 mm, scariosum nitidum inferne lanuginosum, phyllariis
imbricatis 6-seriatis interioribus oblongis obtusis 8 x 1.6-2.2
mn medianis usque 2.8 m latis exterioribus ovatis 4-3 x 3 m,
omnibus nitentibus stramineis sed rubescenti-submarginatis et
cum margine late scarioso hyalino fimbriato instructis.
Receptaculum paulo convexum alveolatum 2 mm diametro.

Flores radii feminei ligulati 16-24 in capitulo. Corolla
pallide violacea glabra 10.5-11.5 mm longa, tubo circa 2 m,
lamina elliptico-oblonga obtusa minute 3-dentata 4-nervata 1.8-
2.2 mm lata. Stylus 5.5 mm ramis stricte linearibus obtusiuscu-
lis crasse stigmatico-marginatis 1.5 mm longis. Ovarium fertile
juvenile 1.5 mm longum basim versus attenuatum striolatum
praesertim sursum copiose glandulosum glandulis albicantibus
suglobosis vel lacrimiformibus interdum minute pediculatis.
Pappus 5.5 mm longus stramineus sed ad apicem purpurascens,
setis interioribus acutissimis complanatis strigosis sursum
leviter ampliatis exterioribus latioribus O.4-1 mm longis.

Flores radii pseudohermaphroditi 20-24 in capitulo.

Corolla 6-6.2 mm longa, tubulo 1.5-1.7 mm glabro, limbo infun-
dibuliformi tubuloso inferne sparsis pilis circa 0.3 m,
dentibus triangularibus acutis incrassato-marginatis apice
papilloso 1-1.2 mm longis. Antherae 2.5 mm basi attenuatae
breviter acuteque sagittatae appendicibus apicalibus oblongis
apice attenuatis subobtusis 0.5 mm longis. Stylus 6.5-7 mm
basi strangulatus cum minuto annulo nectarifero circumdatus,
ramis lanceolato-subulatis 2 mm longis dense papilloso-hirsutis.
Ovarium sterile lineare 5-nervatum exovulatum copiose glandulo-
sum. Pappus 5.5 mm stramineus sursum purpurascentibus setis
interioribus paulo inaequalibus sursum satis dilatatis exterior-
ibus brevioribus latiusculis 0.5-1.5 mm omnibus strigosis.

Typus: Peru, Libertad, Santiago de Chuco: Laguna La
Victoria, camino hacia Consugo, 4000 m alt, ladera rocosa,
arbusto con flores blanco-violaceas; col. 19-VII-1966,

A, Sagastegui, A. Aldave, Fernandez & M. Feekushima 6185.
Holotypus, US.

ORITROPHIUM FIGUEIRASII Cuatr. sp nov.

Herba rosulifolia scapifera rhizomate inclinato vel repente
breviter rosulifero-ramoso copiose radicifero.

Folia omnia rosulata crasse herbacea viridia 12-46 mm longa
obovato-oblonga vel oblongo-elliptica apice obtusa vel paulo
attenuata subacutata 3-5 dentata, deorsum in pseudopetiolum
latiusculum marginatum angustata, basi in vainam amplectentem
margine membranacea ad 3-6 mm latam ampliata, utrinque bene
rugulosa (in sicco) tantum costa subtus satis eminenti supra
leviter impressa vel inconspicua alteris nervis invisibilibus,

1972 Cuatrecasas, Notes on neotropical flora 355

subtus glabra, supra sparse pilosa marginibus inferne petioloque
parce ciliatis, pilis pluricellularibus acutis ascendentibus vel
patulis, rectis vel leviter flexuosis 0.3-0.7 mm longis praedita.
Limbus 7-28 x 4-12 mm, petiolus 5-18 mm longus.

Scapi monocephali 6-32 cm longi singuli vel 2-3 in roseta,
teneri recti erectique firmi fistulosi striolati violacei sparse
bracteati, copiose sursum dense glandulosi, glandulis globosis
minute pedicellatis 0.04-0.1 mm longis, interdum rarissimis pilis
acutis circa 0.2 mm longis muniti. Bracteae scaporum distantes
ovato-oblongae sessiles parce ciliatae virides vél violaceae
3-7 mm longae 1.5-3 mm latae supremae 2-3 capitulum adjacentes.

Capitula radiata erecta circulo ligularum circa 18 um
diametro in vivo (18-25 mm in sicco, compressa), 51-123 flores
ferentia. Involucrum conico-cupulatum 6-7 mm longum, phyllariis
keseriatis subherbaceo-scariosis firmulis pallide viridibus
extremo lilacinis margine hyalinis, interioribus 5-6 x 0.7-1 m
linearibus acutis uninervis margine sursum bene ciliatis,
exterioribus 3.5-4.5 x 1.5-1.1 mm oblongis subobtusis costa et
nervis lateralibus ascendentibus anastomasantibus notatis apice
minute eroso-ciliatis, intermediis gradatis. Receptaculum
alveolatum planum 2.2-3 mm diametro.

Flores radii feminei ligulati 28-66 in capitulo. Corolla
alba 8-6 mm longa, tubo 1.8-2 mm longo sursum leviter ampliatum
pilosiusculo, pilis flexuosis 0.1-0.3 mm, lamina lineari-
oblonga apice attenuata subacuta vel obtusa integra vel minute
2-3-dentata glabra 3-4 venis obsoletis 0.7-0.9 (-1) mm lata.
Stylus 3 mm ramis subulatis 0.5 mm longis. Achaenia ovoideo-
oblonga haud compressa apice paulo angustata basi attenuata
cuneata callosa 2.2-2.5 x 0.7 mm, 5-nervata copiose hispidula
pilis geminis antrorsis rigidis acutis 0.4 mm longis circa
apicem parcis glandulis intermixtis praedita. Semina albida
levissima oblongo-ovoidea. Pappus stramineus 2.5-3 mm setis
subuniseriatis strigosis acutis paulo inaequilongis.

Flores disci pseudohermaphroditi 23-57 in capitulo. Corolla
lutea vel luteo-viridula 3.3-3.5 mm longa, tubulo 1.2-1.4 m
tantum ad apicem sparsis pilis vel papillis, limbo infundibuli-
formi inferne sparse piloso pilis flexuosis circa 0.2 mm, denti-
bus crassis oblongo-triangularibus margine minute papillosis
extus subapicem sparsis pilis. Antherae 1.3 mm longae basi
breviter sagittatae appendicibus apicalibus oblongis subacutis.
Stylus 2.8-3 mm ramis erectis oblongis obtusiusculis extus
dense breviterque papilloso-pilosis. Ovaria 2 mm longa linearia
exovulata sterilia 5-nervata hispidula pilis antrorsis geminis
circa 0.4 mm, circa apicem parcis glandulis munita. Pappus
quam in floribus radii. ,

Typus: Venezuela, Merida: Llano Corredor, junto a la
Sierra de Guirigay, lindero con Trujillo, paramo 3300 m alt,
rosula, hojas crasas verde medio semimate haz, verdes o
violaceas enves; escapo violaceo monocefalo con bracteolas
verdes o verdoso violaceas. Capftulos extendidos 18 mm didmetro,
ifgulas blancas, flosculos amarillos o amarillo-verdosos, col.
J. Cuatrecasas, M. Lépez-Figueiras & L. Marcano-Berti no 28166.

356 P Hey pior nO Gia Vol. 23, no.

Holotypus, US; isotypus MER.

ORITROPHIUM GRANATUM Cuatr. sp. nov.

Herba rosulifolia scapifera rhizomate visu brevi repente
copiose fibroso-radicifero.

Folia omnia basilaria rosulata herbacea crassiuscula
viridia supra nitida 25-65 mm longa 9-20 mm lata. Lamina
obovato-oblonga apice subrotundata vel plus-minusve attenuata
interdum acuta basim versus attenuata in pseudopetiolum alatum
3-6 mm latum, basi ampliato-vaginata amplectenteque, sursum
(quarto vel tertio superiori) dentata dentibus 2-5 utroque latere
acutis antrorsis, marginibus glandulosis et ciliatis pilis
tenuibus acutis patulis 0.5-2 mm longis ornata, utrinque copiose
pediculato-glandulosa et sparse setifera, costa bene notata
basim versus dilatata nervis lateralibus 2-3 utroque latere
ascendentibus arcuato-anastomosatis vulgo inconspicuis.

Seapi monocephali axillares plures 6-30 cm longi argute
striati erecti vel leviter curvati dense glandulosi glandulis
globosis pediculatis 0.2-0.6 mm longis vestiti et sparse
bracteati. Bracteae distantes 15-5 mm longae 2-1 mm latae
sparse glandulosae et pilosae, ad apicem scapi 2-4 bracteolae
capitulum adjacentes.

Capitula radiata erecta solitaria ad terminationes scaporun,
110-140 flores ferentia, circulo ligularum 25-30 mm diametienti,
disco 8-10 m. Involucrum campanulatum subherbaceum 10-11 mm
altum, phyllariis heseriatis paulo inaequalibus stricte lineari-
lanceolatis acuminatisque acutis, interioribus 9-10.5 x 1-1.2
mm paleaceo-seariosis subhyalinis obscure uninerviis glabris
tantum sursum apiceque margine-ciliatis, exterioribus 6-8.5 x
1.2-1.3 mm margine anguste scarioso ceterum viridulo sursum
lilacinis intus saepe 3-nervis visibilibus, extus copiose
glandulosis glandulis globosis pediculatis 0.2-0.4 mm longis.
Receptaculum alveolatum 4-5 mm diametro. Flores radii feminei
ligulati biseriati fertilia interdum aliquis sterilia, 50-73 in
capitulo. Corolla alba vel lilacina 10.5-12.5 mm longa, tubo
2.(/-3 mm praecipue sursum sparse pilosum pilis circa 0.4 mm,
lamina lineari oblonga 1-1.3 mm lata glabra 4 nervis incon-
spicuis apice angustata subacuta vel acuta integra vel minute
2=-3-dentata. Stylus circa 3.5 mm longus vix exsertus ramis
brevibus acutis 0.5 mm. Achaenia 2.4 mm longa ellipsoideo-
oblonga haud compressa apice paulo contracta basi cuneata et
callosa, 5-6 nervata, subhispido pilosa, pilis geminis antrorsis
rectis circa 0.3 mm, ad apicem parcis glandulis fuscis minute
pediculatis etiam praedita. Pappus rubescens circa 4 mm longus
setis strigulosis uniseriatis inaequilongis.

Flores disci pseudohermaphroditi 60-70 in capitulo.

Corolla lutescens 4.5-5.4 mm longa, tubulo 1.8-2.2 mm tantum
apice parce pilosulo, limbo infundibuliformi tertio inferiori
sparse piloso, dentibus triangularibus acutis 1 mm longis
marginibus apiceque minute papillosis extus subapicem 2-44
minutis glandulis pediculatis munitis. Antherae 1.5 mm longae

1972 Cuatrecasas, Notes on neotropical flora 357

basi breviter sagittatae appendicibus apicalibus oblongis
acutatis. Stylus ramis 0.8-1 mm oblongis crassiusculis sub-
acutis extus dense papilloso-hispidulis. Ovarium 2.2 mm exovu-
latum interdum ovulatum sterile 5-6 nervatum dilute antrorso-
pilosi subapicem parcis glandulis. Pappus 4-4.5 mm cum femineus
similimus.

Typus: Venezuela, Merida: Sierra Nevada de Santo Domingo,
Paramo de Los Granates: Canada del Padre, 3370-3380 m alt; hoja
verde brillante haz, escapos mas o menos purpureos, lfgulas desde
blanco a lildceas, col. 12-X-1969. J. Cuatrecasas, L. Ruiz Teran
& M. Lopez-Figueiras 28088. Holotypus US; isotypus MERF.

VERNONIA NEOGLEASONIANA Cuatr. sp. nov.

Arbor circa 6 m alta, ramulis ultimis brunneis teretibus
copiose lenticellatis minute adpresseque tomentosis, denique
glabratis.

Folia alterna crassiuscule coriacea rigida. Petiolus
6-12 mm longus supra sulcatus ceterum breviter denseque tomen-
tosus. Lamina late oblongo-lanceolata basi obtusa vel obtuse
cuneata apice attenuata acutaque margine revoluta integra, 4-7 an
longa 1.8-2.6 cm lata; supra in sicco sordide brunnescenti-
viridis juvenilis puberula adulta glabra rugulosa costa valde
impressa nervis secundariis filiformibus et minoribus plus-
minusve impressis subtus viridi-ochracea dense lanuginoso-
tomentosa costa satis prominenti nervis secundariis prominenti-
bus 16-18 utroque latere patulis circa marginem arcuato-anasto-
mosatis, nervis minoribus etiam prominentibus reticulum minutum
elevatum formantibus.

Inflorescentiae terminales paniculatae subrotundatae con-
geste floriferae folia suprema vix attingentes, 6-10 cm amplae
ramis alternis interdum oppositis angulatis striatisque dense
hirsutulo-tomentosis viridi-ochraceis. Ramuli copiosi tenues
flexuosi angulati tantum ad extremos bracteis pedicellos sub-
tendentibus. Bracteae stricte lineares pedicellos subaequi-
longae. Pedicelli angulati tomentelli recti vel flexuosi 8-3 m
longi tantum ad apicem 1-3 bracteolis minutis juxta capitulum.

Capitula cylindracea basi attenuata, 13-14 mm longa 5-6 mm
lata. Involucrum 11-12 mm altum 28-30 phyllariis imbricatis
circa 5-seriatis subcoriaceis rigidulis, extus adpresse villosis,
intus ad apicem et marginibus plus-minusve villosis ceterum
glabris nitidisque, interioribus oblongis acutatis 8-8.5 x
2-1.8 mm, medianis oblongo-ovatis latioribus usque ad 2.7 mm
latis, externis ovatis 2-3 x 1.2-2 mm, ceteris gradatim transeun-
tibus. Flores 10-11 in capitulo; corolla pallide violacea 7.5-
8 mm longa, tubulo stricto 4 mm longo sursum sparsis pilis
minutis 0.1 mm longis, ad apicem parcis glandulis globosis,
limbo tubuloso paulo ampliato sed maturitate in 5 lobos fere
usque ad basim fisso, lobis linearibus 3-3.5 mm longis 0.5 mm
latis acutis incrassato-marginatis denique recurvatis apice
minute papilloso sursum extus sparsis pilis minutis antrorsis
et copiosis glandulis globosis albis praeditis. Antherae

358 PEED OmO-G Da Vol. 23, no.

purpuraceae 2-7 mm longae basi sagittatae apice appendice
oblonga sessili O.5 mm longa. Stylus extremo 2.5 mm longitu-
dine antrorso-papilloso maturitate ramis duobus subulatis 1.5

mm longis arcuato-divergentibus. Nectarium annulari-

cupulatum circa O.4 mm altum integrum. Ovarium oblongum basim
angustatum basi breviter callosum, 10-costatum, pilis sparsissi-
mis minutis saepe munitum, glandulis rotundis sesilibus deorsum
eopiose praeditum. Pappus stramineus 7 mm longus, setis longis
interioribus biseriatis minute strigosis sursum levissime ampli-
atis, setis exterioribus brevibus numerosis uniseriatis subcom-
planatis strigulosis O.3-1 mm longis.

Typus Ecuador, Imbabura: East of Cayambe, ridge just
south of Rio Clavadero, along trail to Rfo San Pedro, 10,300 ft.
alte (Colls27 dull 19h, Ira L. Wiggins no. 10483. Holotypus,
US. Another collection: Ecuador, Paramos de San Juan, arbusto
4m, flores blancas, Col. 26-II-1931, Benoist no. 3920.

Vernonia neogleasoniana belongs to section Critoniopsis.

It is dedicated, in his 90th anniversary, to honor the outstand-
ing botanist and specialist in Vernonia, Dr. Henry Allen Gleason,
former professor of Columbia University and The New York
Botanical Garden, now retired at Willowdale, Ontario.

ESPELETIA GARCIAE Cuatr. sp. nov.

Caulirosula, caudice simplice vel extremo pauciramoso usque
circa 6m alto 6 cm diametro tantum ad apices folia creba rosu-
lata pallide viridia ferenti. Caules juveniles densissime
barbati pilis circa 8 mm longis erectis ascendentibus flavescen-
tibus congestissimis tecti.

Folia coriacea, adulta rigida, breviter petiolata. Lamina
25-31 cm longa 7-12 cm lata, late lanceolata vel obovato-
lanceolata acuta vel subacuta interdum obtusata, circa basin in
petiolum brevem attenuata et in vaginam amplectentem producta,
margine integra angustissime revoluta; supra juvenili dense
villosa adulta scabrida subtomentosa, pilis robustis circa 1 mum
longis basim incrassatis patulis extremo tenuibus ascendenti-
curvatis, costa magis tomentosa nervis secundariis notatis;
subtus costa crassa elevata, nervis secundariis 20-25 utroque
latere prominentibus inaequaliter ascendentibus angulo 40°-60°
divergentibus 1-2.5 cm inter se distantibus, nervis tertiis
prominentibus in angulo acuto cum venulis reticulum prominentem
formatibus, alveolis minutissimis pilis crispis albis densis
tectis, reliqua subvilloso-tomentosa pilis ad 2.5 mm deorsum
robustis patulis antrorsum tenuibus acutis arcuatis ascendenti-
busque supra costam nervosque saepe subsericeos copiosioribus
densioribusque. Petiolus robustus 1.5-2.5(-5) em longus 0.5-

1 em latus. Vagina fere semiorbiculata late amplectens crasse
coriacea supra glabra extus dense tomentosa et inferne barbata,
14-22 mn alta 25-40 m lata. Folia valde juvenilia mollia
dense lutescenti subsericeo-villosa.

Inflorescentiae axillares folia attingentes vel paulo
excedentes, 30-40 mm longae. Axis mediocris erectus leviter

1972 Cuatrecasas, Notes on neotropical flora 359

striatus dense villosus vel tomentoso-villosus pilis curvato
ascendentibus 1-2 mm longis, nudus tantum ad apicem paniculi-
ferus. Paniculae corymbosae 8-15 cm longae 6-10 cm latae, ramis
alternis interdum suboppositis teneris sed rigidis erectis levi-
ter striatis dense tomentoso-villosis pilis patulo-ascendentibus
flavis, bracteis subtendentibus anguste linearibus basi semi-
amplectentibus, inferioribus usque ad 2.5-7 cm longis 2-3 mm
latis sursum gradatim brevioribus.

Capitula discoidea parvula subglobosa 5-8 mm alta 4-6.5 mm
lata, sessilia 3-6 ad extremos pedunculorum 3-10 m longorum
hirto-tomentosorum congeste glomerata rare singula. Bracteolae
lineares acutae pilosae 6-9 mm longae 1-2 mm latae. Involucrum
5-6 mm altum. Phyllaria sterilia 5 crassiuscule herbacea
rigidula 3.5-6 mm longa 2-4 mm lata oblongo-elliptica vel late
lanceolata acuta vel subacuta 5-9-nervata sursum ciliata extus
moderate patulo-pilosa pilis acutis 0.2-0.6 mm et sparsis
glandulis subsessilibus. Phyllaria fertilia (2) obovata sub-
acuta vel subobtusa basi callosa vetusta incrassatissima reliqua
parte membranacea tenuia 2.6-3.5 x 1.5-2.5 mm dorso praecipue
sursum margineque parce pilosa et glandulosa pilis 0.5 m.
Receptaculum conicum circa 2-2.5 mm altum 1-1.5 m diam glabrun.
Paleae disci membranaceae hyalinae 3-4 x 1.6-2.3 mm obovatae
obtusissimae tenuiter plurivenosae subcarinatae amplectentes
margine sursum parce ciliatae pilis obtusis sparsis et parcis
glandulis subseéssilibus praeditae.

Flores radii feminei 7-11 in capitulo 1-2-seriati.

Corolla brevissima 0.5-1(-2) mm alta cum annulum 0.2-0.5 m
altum densissime crasso-pilosum saepe reducta sed saepe minutum
appendiculum liguloideum abaxialem lineare vel ovatum integrum
vel fissum 0.2-1(-2) mm longum 2-5 nervatum producta; pili
crassi obtusi (raro acuti) 0.1-0.4(-0.5) mm. Stylus 2.5-3.2 mm
longus crassiusculus erectus ramis subulatis circa 1 m.

Ovaria obovato-oblonga triangulata dorso plano-convexa basi
acuta 1.8-2 x 0.8 mm. Achaenia nigricantia obtuse triangulata
vel fere laevia oblongo-obovata apice rotundato basi acuta

2 mm longa 1 m lata.

Flores disci masculi 10-21 in capitulo. Corolla lutea 4-
4.8 mm, tubulo 1.6-1.8 mm longo praecipue sursum pilis patulo-
ascendentibus acutis vel obtusis 0.2-0.8 mm longis et sparsis
glandulis pediculatis munito, limbo tubuloso campanulato basi
sparsis pilis et glandulis, dentibus triangularibus acutis 0.5-
0.6 m longis marginibus incrassatis longeque papillosis dorso
parcis pilis patulis et copiosis glandulis sessilibus vel
subsessilibus instructis. Antherae 1.6-1.8 mm basi breviter
sagittatae appendicibus apicalibus late ovatis subacutis 0.4 m
longis. Stylus circa 4.5 mm longus crassus apice conico-dilata-
to dense papilloso. Nectarium tubulosum obtuse angulatum 5-
denticulatum 0.7-0.8 mm longum.

Typus: Colombia. Boyaca: Leiva, vereda de Capilla,
encima del km 21, en el monte, 2640 m. alt.; tallo liso de 4m
alt x 6 cm diam que culmina en tres cortas ramificaciones,
flores amarillas, 2-XII-1970, Lorenzo Uribe Uribe 6491;

360 P HPL AT Orbs0: Go Tek Vol. 23, nosed

holotypus, US. Ibidem, vereda de Capilla, monte sobre kn. 21,
2640 m, mata 6 m, tallo muy duro bifurcado a 1 m sobre el

suelo, cada rama terminada en cortas ramificaciones apicales,
inflorescencias erguidas con flores amarillas. Crece en
matorral alto. 2>xII-1970, Lorenzo Uribe Uribe 64.92 ; para =
typus, US. Boyaca: Arcabuco, al NE de la poblacion, cerca
iimite con Santander, 2650 m; 4 m, erecta, tallo 5 cm diam,
nervios foliares color oro, pelos amarillos, flores amarillas;
cuando se corta se ramifica; 12-X-1966, H. Garefa Barriga 18764;
paratypus, US.

ESPELETIA JABONENSIS Cuatr. sp. nov.

Acaulirosula densissime foliosa 30-50 cm alta 45-65 cm
lata visu albo-cinereo-sericea nitens. Caudex lignosus sub-
tuberosus pyriformis 6-10 cm longus basi radice axonomorpha
crassa brevi cum ramulis radialis robustis instructa.

Lamina foliorum coriacea rigidula, 14-30 cm longa 3-7
(-10) mm lata, stricte linearis levissime oblanceolata acutaque
basin versus sine sensu attenuata margine valde revoluta, supra
pilis longis sericeis intricatis valde adpressis vestimentum
laevem nitidum argentatum formantibus tecta, subtus costa crassa
elevata dense adpresseque villoso-sericea lucidaque reliqua
superficie adpresse lanato-sericea. Vagina 2-2.5 cm longa 5-8
mn lata adaxiali facie sursum lanato-villosa reliqua glabra,
abaxiali dense longeque sericeo-villosa.

Inflorescentia terminalis thyrsiforme paniculata valde
floribunda 40-80 em alta 25-35 cm lata, axe robusto ad basin
20-30 mm diametro, striato purpurascenti sed spisse albido-
villoso-lanato, dense folioso foliis inferne linearibus acutis
sericeis rosulantibus similimis, sursum gradatim minoribus
(20-10 em x 8-4 mm) e basi copiosissime ramoso. Rami erecti
robustiusculi rigiduli aphylli dense albo lanato-sericei
extremo simplice vel duplicato corymbosi (3-) 6-15 capitula
ferentes, 10-30 cm longi superiores gradatim minores; folia
subtendentia linearia villoso-sericea quam rami valde breviora
sed circa apicem aequilonga. Pedunculi pepedicellive 0.5-

3.5 em longi erecti sericeo-villoso-lanati, bracteis subtendent-
ibus brevioribus, sub apicem parcis bracteolis alternis
linearibus acutis purpurascentibus et albo-villoso-lanatis 20-
12 mm longis 1-1.5 mm latis ad involucrum adjacentibus instruc-
tis.

Capitula radiata erecta vel reclinata circulo ligularum
25=-35(-40) mm disco 10-14 mm diametientibus, 184-250 flores
ferentia. Involucrum cupulatum 12-16 mm diametro dense longeque
villoso-lanatum albo-cinereum, phyllariis sterilibus 16-28
bi-triseriatis linearibus vel lineari-subulatis interioribus
magis lanceolatis acutis vel subacutis gradatim decrescentibus
11-6 x 1.2-2(-2.5) mm extus copiose longeque villoso-vellereis;
phyllariis interioribus fertilibus lanceolatis acutis (8-)6.5 x
1.5-2.5 mm extus marginibusque villosis, saepe etiam sparsis
minutis glandulis pediculatis praeditis. Receptaculum planum

1972 Cuatrecasas, Notes on neotropical flora 361

7-10 mm diametiente sparse pilosum pilis tenuibus erectis ad
0.8 mm longis. Paleae radii 5-4 x 1.1-2 mm late lanceolatae
vel oblongo-lanceolatae acutae subamplectentes, interiores
magis tenues amplectentesque costa signata marginibus hyalinae
et sursum pilosae. Paleae disci lanceolate oblongae acutatae
amplectentes saepe naviculares 4-5 x 1.5-2 mm marginibus late
hyalinis dorso firmo et extremo pilis copiosis antrorsis sub-
acutis vel subobtusis 0.3-0.5 mm longis et parcis glandulis
subglobosis pediculatis instructis.

Flores marginales feminei ligulati 52-86 in capitulo 3-
seriati. Corolla lutea vel luteolo-aurantiaca 8-13 mm longa
tubulo 1.22.2 mm longo stricto copiose piloso pilis patulo-
antrorsis hyalinis subobtusis vel acutis seu obtusis 0.3-0.4
(-0.8) mm longis interdum parcis glandulis pediculatis vel
subsessilibus praedito, apice saepe appendice adaxiali minuto
dentiformi vel elongato lineari curvato ad 1 mm longo raro
duplo; lamina lineari vel oblonga basi apperta apice 2-3-
dentata 1-1.8(-2) mm lata 4-8-nervata supra minute papillosa
velutina subtus sparsis vel sparsissimis pilis 0.2-0.3 mm
longis et interdum parcis minutis glandulis munita sed ad
basim semper pilis copiosioribus instructis. Stylus 3.5-5 mm
longus ramis 1.2-1.5 mm crassiuscule lanceolatis. Achaenia
atra exteriora obovoidea triquetra apice rotundata basi cuneata
breviter callosa dorso convexo 1.8-2 x 1.3-1.5 m, interiora
oblonga basim attenuata subquadrangulata dorso subcostato
ere 2) St AN Brailes

Flores disci pseudohermaphroditi 110-154 in capitulo.
Corolla lutea 4.5-5.5 mm longa, tubulo 2-2.3 mm hirtulo pilis
subpatulis 0.3-0.6 mm saepe parcis glandulis subglobosis pedi-
culatis vel subsessilibus munito, limbo tubuloso-campanulato
subglabro dentibus triangularibus 0.8-1.2 mm longis acutis
marginibus papillosis extus parcis pilis patulis 0.3-0.4 mm et
interdum raris glandulis subsessilibus praeditis. Antherae
oblongae 1.7-2 mm basi sagittatae apice appendice ovata acutata
0.35-0.4 mm. Stylus circa 5 mm. Nectarium 0.7-0.8 mm longum
erassiusculum tubulosum leviter quinquedentatunm.

Typus: Venezuela, Trujillo: Tres Pozos, sector del Paramo
del Turmal, 2800-3850 m, 14.4 km al E de Carache; acaulirrosula
50 em alto, 65 cm diam, hojas blanco-plateado-sericeas en ambas
caras; inflorescencia terminal 45 cm alto 28-35 cm diam,
capftulos 3-4.5 cm diam total, disco 14 mm diam; ifgulas y
flosculos amarillo a amarillo anaranjado claro; dominante en la
localidad pero contados individuos enetiilor, 8-VI-1971 colect.
Luis Ruiz-Terdn & Manuel Lopez -Figueiras 1995 « Holotypus US,
isotypus MER. Trujillo: Paramo de Cendé, Hoya de Los Carruzos,
cerca de Cerro de Los Muertos, distrito Carache, 2900 m;
acaulirrosula 33 cm alta 45 cm ancha, inflorescencia terminal
60 em, capftulos 35-38 mm diam total, disco 13-15 m diam,
involucro subhemisférico 14-16 mm diam, ifgulas amarillas o
amarillo-anaranjado claras; dominante en la localidad pero escasa
en flor, colect. 10-VI-1971 Luis Ruiz-Terdn & Manuel Lopez =
Figueiras 2101; paratypi, US, MER. Another collection: Pgramo

362 P HEY Onn0 Gerre Vol. 235 nouye

de El Jabon, 3100= -3400 m alt, vertiente oriental; rosula, hoja
argentada muy brillante, restos de inflorescencias secas,
2-xI-1969, J. Cuatrecasas, L. Ruiz-Terdn & M. Lopez-Figueiras
28217 (US, MER).

ESPELETIA RUIZII Cuatr. sp. nov.

Rosula dense foliata virescens, circa 30 cm alta 65 cm
diametro, breviter stipitata. Lamina foliorum coriacea rigida,
20-30 cm longa, 6-10(-12) mm lata, anguste lanceolato-linearis
apicem gradatim attenuata acutataque, basim versus sensim sine
sensu angustata basi 4-3 mm lata, margine revoluta; supra primo
laxe sericeo-villosa pilis longis et longissimis parallelis
ascendentibus subadpressis munita deinde (praeter basim villoso-
lanuginosam) glabrata subnitenti-viridis laevis tantum costa
impressa notata; subtus costa crasse elevata nervis secundariis
patulis prominentibus 1.5-2.5 mm inter se distantibus nervulis
minoribus prominente reticulatis, primum cum indumento superfi-
ciali laxo villoso subsericeo pilis longis et longissimis deci-
duis vestita sed costa persistente subsericeo-villosa, reliqua
cum dense crasseque indumento albo intricato-lanato persistenti
tecta. Vagina foliorum subtrapezoidea apice paulo attenuato in
adultis subrotundato, basi latiori truncata multinervata facie
adaxiali apice excepto glabra, abaxiali dense adpresse longeque
albo-villoso-sericea margine sursum apiceque longe sericeo-
barbata, 27-35 mm longae 22-33 mm latae. Folia inicialia gemmae
terminalis dense adpresseque albo-subsericeo-villosa.

Inflorescentia terminalis magna subcorymbiforme paniculata
quam rosula foliorum circa duplo longior et latior. Axis
robustus ad basim circa 5 cm diametro medullosus sursum gradatim
angustatus fistulosus angulatus striatusque ex basi ramosus ramis
alternis ascendentibus usque 30, proximalibus longioribus ad 65
em longis centrales attingentibus. Axis ramique epidermide
rubiginosa cum indumento densiusculo villoso albo pilis longis
et longissimis tenuibus subsericeis ascendentibus deinde intri-
catis aspectu lanato=barbatis tecti. Folia caulina subtendentia
rosulantia similia sed sursum gradatim breviora et indumento
externo gossypino magis copiosis vaginis latis amplectentibus
carneis vel rubris. Ramuli ultimi quam priores semper stricti-
ores dense albo-vellereo-barbati, subterminales graciles 3-10
em longi alterni vel suboppositi foliis subtendentibus sub-
aequilongis, terminales cymosi dichasiales vel monochasiales
saepe ramulo mediali brevi vel brevissimo cum capitulo termi-
nato, lateralibus longioribus etiam floriferis, bracteis sub-
tendentibus supremis pedicellos 0.5-6 em longos superantibus
vel attingentibus patulis saepe infra capitula radiatis. Lana
alba sericeo-barbata inter bracteas pedunculos et calathia
valde copiosa.

Capitula radiata erecta vel inclinata, 80-100 flores
ferentia, circulo ligularum 16-22 mm diametienti cum phyllariis
involucri exterioribus stellatis valde superato, disco convexo
10-12 mm diametienti. Involucrum herbaceum cupulatum dense

1972 Cuatrecasas, Notes on neotropical flora 363

longeque albo-lanato-barbatum, phyllariis sterilibus biseriatis
exterioribus 7-9 oblongo-lanceolatis acuminatis extremo radiatis
flores valde excedentibus, 14-9.5 mm longis 4-3.5 mm latis,
interioribus 2-4, late lanceolatis 9.5-8 mm longis 4.5-3 um
latis, facie adaxiali viridibus leviter nervatis subglabris extus
albissime copioseque barbato-lanatis; phyllariis intimis fertil-
ibus tenuioribus ovato-lanceolatis acutis acuminatis plus-minusve
arcuatis praecipue sursum longe villosis, 8-7.5 x 3.5 m.
Receptaculum convexo-subconicum 5-6 mm diametro glabrum. Paleae
exteriores femineae oblongae subacutatae curvatae subamplect-
entes circa 6 x 2 m ad apicem copiose pilosae ciliataeque pilis
obtusis. Paleae disci oblongae acutae amplectentes 5-7
paralleli-nervatae costula prominenti, dorso praecipue sursum
pilosae barbulatae pilis crassiusculis obtusis (clavatis) O.4-
0.8 mm longis.

Flores marginales feminei ligulati 32-36 in capitulo tri-
seriati. Corolla 5.2-7 mm longa; tubo 1.7-1.9 mm copiosis pilis
patulis minutis crassiusculis capitato-glandulosis 0.06-0.08 mm
longis et raris pilis clavatis 0.4-0.5 mm, apice appendice
adaxiali minuta obtuse 1-2-dentata callosa instructo; lamina
eburnea deinde lutescenti vel apicem versus aurantiaca vel
rosea, elliptico-oblonga obtusa bi-tridentata 5-7 nervata circa
2(1.8-3) um lata supra glabra subtus sparsis minutis glandulis
munita. Stylus 4-4.5 mm longis ramis sublanceolatis 1-1.2 m
longis. Achaenia nigricantia exteriora deltoidea argute tri-
angulata 2 x 1.6 m, interiora subquadrangulata 2.3 x 1.3 mm,
omnia basi acutissima.

Flores disci pseudohermaphroditi 48-64 in capitulo. Corolla
rubescenti 5.4-5.7 mm longa; tubulo crassiusculo circa 2 mm
longo sparsis glandulis pediculatis et rarissimis pilis longi-
oribus munito; limbo tubuloso-campanulato praecipue ad basim et
sursum sparse glanduloso et sparsissimis pilis clavatis 0.4-

0.5 mm praedito, dentibus triangularibus acutis crasse papillo-
seque marginatis 1-1.2 mm longis. Stylus circa 6 m longus ad
apicem dense breviterque papilloso-pilosus. Antherae 2 mm
longae basi obtusiuscule sagittatae, appendice apicali oblongo-
ovata 0.4-0.5 mm longa. Stylus circa 6 mm longus apice dense
breviterque papilloso-pilosus. Nectarium tubulosum 1 mm altum
breviter 5-denticulatum. . ‘

Typus: Venezuela, Merida: Paramo de Las Coloradas, Loma
de La Libertad, 2750-2800 m, a unos 500 m de El Portachuelo
(El Ramal) entre Santa Cruz de Mora y El Molino, municipo
Estanques, distrito Sucre, "Caulirrosuleto, roseta 30 cm alta
65 cm diam., hojas verde intensas sublucientes haz, panfeula
corimbosa terminal 105 cm diam, vainas foliares caulinares,
ejes y ramas inflorescenciales rosados o vinosos cubiertos con
indumento lanoso-algodonoso, blanco, serfceo; lfgulas cremosas
tornandose amarillentas y anaranjadas o rosadas siquiera en los
apices; flosculos rojizo vinosulos." 16 enero 1971 colect.

Luis Ruiz Teran & Manuel Lopez Figueiras 1457. Holotypus US;
isotypus MER.

364 POHVY/TrO0 LO0e sek Vol. 23, no.

ESPELETIA ULOTRICHA Cuatr. sp. nov.

Acaulirosula parva albo-lanata 15-20 cm lata 10-15 cm alta.
Caudex brevi ad 5 cm longus subterraneus crassus vel tuberosus
(3-4 cm diametro) radice robusta acuta brevi sed ramosa ramulis
crassiusculis divergentibus.

Folia coriacea linearia crasse albo-lanato vestita tacto
valde mollia, in sicco cinerascentia vel roseata vel rufescen-
tia. Lamina anguste linearis subobtusa basim versus sine sensu
attenuata et in vaginam paulo ampliatam producta 8-22 ecm longa
5-10 mm lata (sed cum indumento 10-14 mm lata et apice obtuso),
basi 5-7 mm lata (cum indumento 7-10 mm); supra epidermide
minute rugulosa tantum nervo medio impresso apparente; subtus
margine revoluta costa valde crassa notata alteris nervis in
reticulum minutum prominentem anastomosatis, alveolis cum lana
minuta candidissima repletis, sed omni architectura laminae cum
maximo indumento crispo-lanato crassissimo subdenso extus plus-
minusve arachnoideo cancellata. Vaginae oblongae apice sub-
rotundatae basi paulo angustatae nervato-striatae 20-43 mm
longa 6-10 mm latae facie adaxiali glabrae virescentes abaxiali
dense longeque sericeo-barbata. Folia inicialia gemmae termi-
nalis crasse densissimeque niveo-sericeo-villosa.

Inflorescentiae axillares scapiformes 1-5 quam rossula
2-3-plo longiores. Scapi 1-3-cephali erecti vel curvati 15-60
em longi subteneri sed ob indumento crispo lanato denso visu
crassiores, circa basim 1-4 foliis alternis linearibus 4-10 cm
longis densissime albo-lanatis instructis, saepe duobus foliis
inferioribus adjacentibus vaginis uno latere connatis; sursum
etiam parcis foliis gradatim minoribus instructis, supremis
bracteiformibus 1-2 em longis capitulum non vel vix attingen-
tibus. Indumentum scaporum crispo-lanatum superficie plus-
minusve araneosum, album sed ad extremos et in capitulis
fulvum.

Capitula radiata cernua 300-500 flores ferentia circulo
ligularum 40-45 cm diametienti, disco circa 20 mm diametro.
Involucrum cupulatum 20-25 mm latum, complanatum 40 mm dia-
metro, alborufescens, phyllariis herbaceis 19-28 sterilibus
3-4-seriatis lanceolato-subulatis acuminatisque 20-12 mm longis
2.2-1.4 mm latis densissime longe crasseque albo-lanatis,
phyllariis intimis fertilibus subulatis 12-10 mm longis 1.8-
1.2 mm latis extus dense longeque villosis barbulatisque.
Receptaculum plano-convexum 10-15 mm diametro hirsutulum pilis
tenuibus sericeis erectis 0.5-3.5 mm longis dense praeditum.
Paleae exteriores femineae oblongo-lanceolatae acutae sub-
amplectentes 6.5-7 x 1-1.4 mm costa viridula marginibus
hyalinis dorso apiceque copiose villosae. Paleae disci
hyalinae oblongae acutae amplectentes costa et sursum longe
erecto-pilosae, 6-6.8 x 1.3-1.5 mm. Pili paleae stricti acuti
erecti 1-1.5 mm longi.

Flores radii feminei ligulati 70-140 in capitulo tri-
seriati. Corolla lutea 14-18 mm longa; tubo 1.5-2.3 mm longo
apice appendice adaxiali dentiformi vel lineari 0.5-1.2 mm
longa, copiose piloso pilis crassiusculis obtusis vel

1972 Cuatrecasas, Notes on neotropical flora 365

longioribus acutis, 0.2-0.5 mm longis, patulis vel curvato-
antrorsis et sparsis glandulis globosis stipitatis; lamina
lineari obtusiucula minute obtuseque 2-3-dentata, 1.7-2 mm
lata, 7-6-nervata supra minutissime epidermo-papillosa extus
glandulis globosis subsessilibus sparsis et pilis sparsissimis
basim versus copiosioribus munita. Stylus 5.5-6 mm longus
ramis lanceolatis crasse marginatis 2-2.5 mm longis. Ovaria
obovato-oblonga basi acuta exteriora triangulata 2.5 x 1.4 mm,
interiora quadriangulata 2.7-2.8 x 1.2 m.

Flores disci pseudohermaphroditi 180-360 in capitulo.
Corolla lutea 6-7 mm longa; tubulo 2-2.5 mm longo sursum
sparsis pilis crassiusculis minutis obtusis 0.1-0.3 mm longis;
limbo tubuloso tantum basi parcis pilis, dentibus oblongo-
triangularibus acutis 1 mm longis crasse papilloseque marginatis
apice papilloso extus glandulis globosis et saepe raris (1-4)
pilis praeditis. Antherae 2.3 mm longae subobtuse sagittatae,
appendice apicali oblongo-ovata subacuta 0O.4-0.5 mm longa.
Stylus circa 7 mm longus extremo dense breviterque papilloso-
pilosus. Nectarium tubulosum crassiusculum minute denticulatum
0.5-0.6 mm longum.

Typus: Venezuela, Lara: Paramo del Jabon vertiente
oriental, 3400 mm alt, en lo alto junto al 1fmite con el estado
de Trujillo; TogaiEnt pequenhas de porte blanco, hoja blanda
densamente blanco lanuda todo alrededor; escapos monocéfalos fo)
raramente 2-3-cefalos, densamento blanco-lanudos, en el extremo
mas o menos leonados asi como el capitulo, ligulas y flosculos
amarillos; 2-XI-1969 colect. J. Cuatrecasas, L. Ruiz-Teran &
M. Lopez -Figueiras 28220; holotypus US, isotypus MER. Lara-
Trujillo: Paramo de El Jabon en Los Pocitos de El Alto (15 km
al E de Carache, Trujillo) 3100 m; Acaulirrosuleto, hojas
densamente lanosas en ambas caras, escapos axilares l-céfalos,
a@ veces Bae erat eap{tulos pendulos, ligulas y flosculos
amarillos; 2-X-1970, L. Ruiz-Teran & M. Lopez-Figueiras 929;
paratypi US, MER.

ADDITIONAL NOTES ON THE GENUS VERBENA. XIII

Harold N. Moldenke

VERBENA [Dorst.] L.

Additional & emended bibliography: Macer Floridus, De Virib.
Herb., pr. 2, xxxiii--xxxv. 1490; Brunf., Herb. Viv. Icon., ed. 1,
1: 119, 120, & [267] (1530) and ed. 2, 1: 119, 120, & [267]. 1532;
Macer, Herb. Virib. 133--135, 16h, 167, & 199--200. 1581; Gerarde,
Herb., ed. 1, 580--582, fig. 1 & 2 (1597) and ed. 2, 717--719,
fig. 1 & 2. 1633; Ray, Hist. Pl. 2: 1336 & [1967]. 1693; Sauzé &
Maillard, Fl. Dép. Deux-Sévres, ed. 1, 2 (2): 26. 1880; Thomé, Fl.
Deutschl. ): 171--173, pl. 531. 1889; Sanders, Encycl. Gard., ed.
2, 4o9--410. 1897; Druce, Fl. Berks. 390 & 61. 1898; Chadwick, N.
Y. State Mus. Bull. 12): 150. 1908; Felt, N. Y. State Mus. Bull.
12h: OS (1908) and 200: 181--182, 225, & 309. 1917; H. H. Thomas,
Round Year Gard., ed. 2, pr. 1, 122 (1920), ed. 2, pr. 2, 122
(1923), and ed. 2, pr. 3, 122, 1926; Sanders, Encycl. Gard., ed.
19, 4h7--hy9. 1930; Macself in Sanders, Encycl. Gard., ed. 21, pr.
1, 456--457 & 459. 1931; G. Klein, Handb. Pflanzenanal. 2 (1): 505
& 968 (1932), 3 (1): 57h, 583, 61h, 616, 635, 640, 6, 647, & 66k
(1932), 3 (2): 932 (1932), and (2): 867, 873, & 1860. 1933; Mac-
self in Sanders, Encycl. Gard., ed. 21, pr. 2, 56--457 & 59.
193); Nemoto, Fl. Jap. Suppl. [Nipp. Shokub. Soran-hoi] 626. 1936;
Takenouchi, Journ. Nat. Hist. Fukuoka 2: 15. 1936; Macself in San-
ders, Encycl. Gard., ed. 21, pr. 3, 456--457 & 459 (1938), ed. 21,
pre h, 456-57 & 459 (19h2), and ed. 21, pr. 5, 456-157 & 459.
1945; Schnack & Covas, Darwiniana 7: [71]--79, pl. 1--5~ 19h53
Covas & Schnack, Darwiniana 7: 85--86, fig. 1 & 2. 195; Macself
in Sanders, Encycl. Gard., ed. 21, pr. 6, 456--)57 & 459. 196;
Prased, Indian Forest. 75: 38. 1949; Hellyer in Sanders, Encycl.
Gard., ed. 22, pr. 1, 456—l57 & 459 (1950), ed. 22, pr. 2, 506—
508 (1952), and ed. 22, pr. 3, 506--508. 1956; Letty, Dyer, Ver-
doorn, & Codd, Wild Fls. Transv. 280 & 281. 1962; Anon., Cat. Sem.
Hort. Bot. Univ. Valent. 27. 1963; Anon., Delect. Sem. Hort. Bot.
Cent. Thbilis. Georg. 25. 1966; Hutchins, Sida 3: 182. 1967; Bat-
tersby, Hall, & Southgate, Journ. Chem. Soc. Lond. Sec. © 5: 721--
728. 1969; Raman, Curr. Sci. 38: 579--580. 1969; Raju, Bull. Bot.
Soc. Bengal 23: [69] & 70. 1969; S. P. & R. N. Banerjee, Bull. Bot.
Soc. Bengal 23: 168. 1969; Macer, Virib. Herb., ed. 2, pr. 2,
[86]--[88]. 1970; J. Harris Co. [Rochester, N. Y.], Care Home Gard.,
rev. ed., 21, 23, & 32. 1970; Anon., Hort. Abstr. 40: 809. 1970;
Schmelzer, Phytopath. Zeit. 67: 309. 1970; Anon., Hortic. Abstr.
hl: 1108 & 1117. 1971; Anon., Zierpflanzenbau 11: 187--189. 1971;
Khoshoo & Arora, Am, Hort. Mag. 50: 16--18, fig. 1--5. 1971; Cad-
bury, Hawkes, & Readett, Comput.-map. Fl. Warwicksh. 197. 1971; G.
E. Tucker, Castanea 37: 23. 1972; Moldenke, Phytologia 23: 257--
303. 1972; A. L. Moldenke, Phytologia 23: 317 & 318. 1972; Hinton
& Rzedowski, Journ. Arn. Arb. 53: 167. 1972.

The Druce (1898) work, cited above, although dated "1897" on

366

1972 Moldenke, Notes on Verbena 367

its title-page, was not actually published until 1898, according
to an author in Journ. Bot. 36: 10) (1898).

Beutenmueller (1907) reports the itonid "vervain leaf midge",
Itonida verbenae Beutm,, attacking members of the genus Verbena;
Felt (1911) reports the itonid, Cecidomyia sp. and the same
authority (1908) reports another itonid, Lestodiplosis verbeni-
folia Felt, from this gems. The vervain leaf midge causes mar-
ginal leaf rolls; the first-mentioned of Felt's itonids causes
irregular oval stem galls, 3--5 mm. long, and the second is
probably predaceous in rolled leaves.

Parks (1937), in speaking of V. quadrangulata Heller, notes
that "There are twenty-two other 5) species of verbena within the
state [of Texas]. These range from anmals through perennials
and through color from white to deep blue. A number of these
have been taken into cultivation and can be purchased from seed
stores. Some of the blue-flowered species produce quite large
and showy spikes of flowers but as yet have not become common in
cultivation. Those wishing to try new flowers should attempt the
cultivation of the wild verbenas found within their own terri-
tories."

VERBENA AMBROSIFOLIA Rydb.

Additional synonymy: Glandularia ambrosifolia Schnack & Covas,
Darwiniana 7: [71], in textu. 195.

Additional & emended bibliography: Schnack & Covas, Darwiniana

: [71]. 1945; Moldenke, Phytologia 23: 258. 1972.

Additional citations: NEW MEXICO: Sierra Co.: Spellenberg &
Todsen 2543 (N).

VERBENA BONARIENSIS L.

Additional bibliography: Macself in Sanders, Encycl. Gard.,
ed. ets pr. 1, 457 (1931), ed. 21, pr. 2, 457 (193k), ed. 21,
pr. 3, 457 (1938), ads,21,/pre les "57 (1942), eds 21,--pre 5; "497
(19k5 4s}, and ed. ai, pr. oe UiS7. i9h6; Hellyer in Sanders, Ency—
cl. Gard., ed. 22, pr. l, "457 (1950), ed. 22, pr. 2, 506 & 507
(1952), and ed. 22, pr. 3, 506 & 507. 1956; Moldenke, Phytologia
nee 259, 265, 289, *290, 292, & 293. 1972.

VERBENA BRACTEATA Lag. & Rodr.

Additional bibliography: Anon., Cat. Sem. Hort. Univ. Valent.
27. 1963; Moldenke, Phytologia 23: 259. 1972; A. L. Moldenke, Phy-
tologia 23: 317. 1972.

VERBENA CANADENSIS (L.) Britton

Additional bibliography: Macself in Sanders, Encycl. Gard., ed.
21, pr. 1, 457 (1931), ed. 21, pr. 2, 457 (193h), ed. 21, pr. 3,
457 (1938), ed. 21, pr. h, 457 (1942), and ed. 21, pr. 5, 57.
1945; Schnack & Covas, Darwiniana 7: [71]. 195; Macself in San-
ders, Encycl. Gard., ed. z) pr. 6, 457. 1946; Hellyer in Sanders,
Encycl. Gard., ed. 22, pr. 1, 457 (1950), ed. 22, pr. 2, 506 &
507 (1952), and ed. 23, pr. ae 506 & 507. 1956; Schmelzer, Phyto-

368 P WNT.O LeOsG ers Vol. 23, mopey

path, Zeit. 67: 309. 1971; Moldenke, Phytologia 23: 259-—-260 &
265. 1972.

VERBENA COMONDUENSIS Moldenke
Additional bibliography: Moldenke, Phytologia 23: 193. 1972.
Additional citations: MEXICO: Baja California: Moldenke & Mol-
denke 25,07 (Ws). 7. % <i

VERBENA CRITHMIFOLIA Gill. & Hook.

Additional & emended bibliography: Schnack & Covas, Darwiniana
7: [71], 72, 7h, & 75+ 1945; Moldenke, Phytologia 23: 260 & 271.
1972.

Bolkhovskikh & his associates (1969) cite the Darlington & Wy-
lie (1956) reference in the bibliography of this species as "Dar-
lington, Chrom. Numb. Flow. Pl. 1955".

VERBENA GOODDINGII Briq.

Additional bibliography: Moidenke, Phytologia 23: 23l--237.
1972.

Holmgren found this species "locally common" on rocky basalt
slopes. The corolla is described as having had its "tube pale-
yellow, lobes blue" on N. H. Holmgren 3308.

Additional citations: ARIZONA: Mohave Co.: N. H. Holmgren
3308 (N). oe

VERBENA HASTATA L.

Additional & emended bibliography: Felt, N. Y. State Mus.
Bull. 200: 182. 1917; Moldenke, Phytologia 23: 261--268. 1972;
G. E. Tucker, Castanea 37: 23. 1972.

Tucker (1972) records this species from Ashe County, North
Carolina.

VERBENA HOOKERIANA (Covas & Schnack) Moldenke
Additional & emended bibliography: Schnack & Covas, Darwiniana
7: [71] & 72. 1945; Moldenke, Phytologia 23: 270--271. 1972.

xVERBENA HYBRIDA Voss

Additional synonymy: Verbena hybrida cv. drandiflora Hort. ex
Anon., Delect. Sem. Hort. Cent. Thbil. Georg. 25, sphalm. 1966.

Additional bibliography: Sanders, Encycl. Gard., ed. 2, 09—
410 (1897) and ed. 19, hh7—lh9. 1930; Macself in Sanders, En-
cycl. Gard., ed. 21, pr. 1, 456--57 & 459 (1931), ed. 21, pr. 2,
156--457 & 459 (193k), ed. 21, pr. 3, 456-157 & 59 (1938), ed.
21, pr. bh, 456—U57 & 459 (192), ed. 21, pr. 5, LS6—LS7 & 459
(1945), and ed. 21, pr. 6, 456157 & 459. 1946; Hellyer in San-
ders, Encycl. Gard., ed. 22, pr. 1, 456—-57 & 459 (1950), ed.
22, pr. 2, 506--508 (1952), and ed. 22, pr. 3, 506--508. 1956;
Anon., Delect. Sem, Hort. Bot. Cent. Thbilis. Georg. 25. 1966;
Raman, Curr, Sci. 38: [579]--580. 1969; J. Harris Co. [Rochester,
N. rei Care Home Gard., rev. ed., 21, 23, & 32. 1970; Anon.,
Hortic. Abstr, 0: 809 (1970) and 41: 1108 & 1117. 1971; Anon.,

1972 Moldenke, Notes on Verbena 369

Zierpflanzenbau 11: 187--189. 1971; Khoshoo & Arora, Am. Hort.
Mag. 50: 16--18, fig. 1, 3, & 5. 1971; Moldenke, Phytologia 23:
271--278. 1972; A. L. Moldenke, Phytologia 23: 318. 1972.

Additional illustrations: Khoshoo & Arora, Am. Hort. Mag. 50:
16--18, fig. 1, 3, & 5. 1971.

An anonymous writer (1971) gives germination percentages and
notes on seedlings and seedling development from pelleted seeds of
this plant. Raman (1969) states that the absorption spectrum of
the acetone extract from a purple corolla showed 3 peaks in the
green-blue, green, and yellow regions, a characteristic of flora-
chrome.

An additional horticultural variety offered is

Ruffled White (Burpee) —- a "superb sweetly scented variety
producing large well-filled trusses of pure white flowers so
freely that the display has the appearance of a white carpet";
represented by Burpee seed no. 3177.

VERBENA INCISA Hook,

Additional bibliography: Sanders, Encycl. Gard., ed. 2, 09.
1897; Moldenke, Phytologia 23: 278—280 & 300. 1972; A. L. Mol-
denke, Phytologia 23: 318. 1972.

VERBENA LACINIATA (L.) Briq.

Additional bibliography: Macself in Sanders, Encycl. Gard.,
ass28s pres eh57 (93h), eds 21, prs 2, 157°(193h), ede 2h;
PEetae W571(1938) 52 eds) QA seprs: yy 57 (I9h2), ed. 21,iprs 5557
(19455, and ed. 21, pr. 6, 457. 196; Hellyer in Sanders, Encycl.
Gard., ed. 22, pr. 1, 457 (1950), ed. 22, pr. 2, 506 & 507
(1952), and ed. 22, pr. 3, 506 & 507. 1956; Raman, Curr. Sci. 38:
[579]. 1969; Moldenke, Phytologia 23: 282—28). 1972.

The "Verbena erinoides" and "Verbena ericoides" of Macself
(1931--19)6) and of Hellyer (1950--1956) are almost certainly V.
tenuisecta Briq.

VERBENA LITORALIS H.B.K.
Additional bibliography: Moldenke, Phytologia 23: 270 & 288—
295. 1972; Hinton & Rzedowski, Journ, Arn. Arb. 53: 167. 1972.

VERBENA LITORALIS var. ALBIFLORA Moldenke
Additional bibliography: Moldenke, Phytologia 23: 29h. 1972;
Hinton & Rzedowski, Journ. Arn, Arb. 53: 167. 1972,

VERBENA MACDOUGALII Heller

Additional bibliography: Moldenke, Phytologia 23: 265 & 297--
298. 19725

The Holmgrens describe this species as "locally common along
roadsides in ponderosa pine forests".

Additional citations: ARIZONA: Coconino Co.: Holmgren & Holm
gren 691 (N). eae) ae

VERBENA MENTHAEFOLIA Benth,
Emended synonymy: Verbena hintonii Moldenke, Alph. List Inval-

370 P HeAeP 6 Ty Oy Graci k Vol. 23, no.

id Names Suppl. 1: 2), in syn. 1947; Hinton & Rzedowski, Journ.
Arn. Arb. 53: 167. 1972.

Additional bibliography: Moldenke, Phytologia 23: 301--303.
1972; Hinton & Rzedowski, Journ. Arn. Arb. 53: 167. 1972.

VERBENA MICROPHYLLA H.B.K.

Additional synonymy: Glandularia microphylla Schnack & Covas,
Darwiniana 7: [71], 72, & 7h, pl. 3 D, in textu. 1945.

Additional & emended bibliography: Schnack & Covas, Darwiniana
7: [71], 72, & 7h, pl. 3 D. 1945; Moldenke, Phytologia 23: 303.
1972,

Emended illustrations: Schnack & Covas, Darwiniana 7: pl. 3 D.
195.

Material has been misidentified and distributed in some herbar-
ia as Geranium sp. On the other hand, the Mutis 1919, distributed

as us microphylla, is actually V. ciliata sie Mutis 3691 is V.

ee ieee et |

Rose 22,00 are Hiapcnotane inflata (H.B.K.) Briq.

Additional citations: ECUADOR: Chimborazo: D. H. Knight 401
(Ws). PERU: Puno: Vargas Calderén 20883 (Ac). ~ CHILE: Santiago:
Zollner 3070 (Lk). ARGENTINA: Tucum4n: Olea 236 (N).

VERBENA MINUTIFLORA Briq.

Synonymy: Verbena minutiflora "Briq. ex Moldenke" apud Angely,
FileAnal). ‘Paran,; “ad. 2-572 . 1965.

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 572.
1965; Moldenke, Phytologia 16: 98, 101, & 102. 1968; Angely, Fl.
Anal. Fitogeogr. Est. S. Paulo, ed. 1, lh: 840 & xix, map 1395.
1970; Moldenke, Fifth Sum, 1: 178 & 190 (1971) and 2: 68 & 917.
1971; Moldenke, Phytologia 23: 293. 1972.

The corollas on Hatschbach 2283 are described as having been
' "lilac" in color when fresh.

Additional citations: BRAZIL: Parand: Hatschbach 22)83 (Rf);
Reitz & Klein 17805 (N, W—25)833). Rio Grande do Sul: Rambo
15339 (Au--122317, Au--122321, Go). Santa Catarina: Smith &
Klein 1369 (N).

xVERBENA MONCHINA Moldenke

Additional synonymy: Verbena urticifolia angustifolia Bebb, in
herb,

Additional bibliography: Moldenke, Phytologia 16: 101--102.
1968; Moldenke, Fifth Sum. 1: 34—39, 4h, 45, 47, 53, & 371
(1971) and 2: 522, 651, 672, 679, 695--698, 708, & 917. 1971; Mol-
denke, Phytologia 23: 265. 1972.

Additional citations: ILLINOIS: Winnebago Co.: M. S. Bebb s.n.
[Fountaindale] (Pa). MISSOURI: Saint Louis: Drummond s.n. [St. _
Louis, N. Am.] (Au--122287); Eggert s.n. [Prairies, Sept. pt. 1891]
(Pa); hae taat sen. [St. Louis, July 182] (Au—122722).

1972 Moldenke, Notes on Verbena 371

VERBENA MONACENSIS Moldenke
Additional bibliography: Moldenke, Phytologia 10: 157-—-158.
1964; Moldenke, Fifth Summ, 1: 371 (1971) and 2: 917. 1971.

VERBENA MONTEVIDENSIS Spreng.

Additional & emended bibliography: Cabrera, Fl. Alred. Buenos
Aires 395 & 397. 1953; Darlington & Wylie, Chrom, Atl., pr. 1,
323. 1956; Schnack, Fehleisen, & Cocucci, Revist. Fac. Acron. La
Plata 35: [7], 9, ([SuJ, & 55, fig. 3. 1959; Darlington & Wylie,
Chrom., Atl., pr. 2, 323. "1961; Angely, Fl. Anal. Paran., ed. 1,
571 & 572. 1965; Troncoso in Cabrera, Fl. Prov. Buenos Aires Ss
128 & 131. 1965; Moldenke, Phytologia 16: 98, 101, & 102. 1968;
Moldenke, Résumé Suppl. 16: 7. 1968; Bolkh., Grif, Matvej., &
Zakhar., Chrom. Numb. Flow. Pl. 717. 1969; Moldenke, Fifth Summ,
1: 9, 178, 188, 190, 201, & 371 (1971) and 2: 917. "1971; Moldenke,
Phytologia 22: 489 (1972) and 23: 293 & 295. 1972.

Additional illustrations: Schnack, Fehleisen, & Cocucci, Revist.
Fac. Agron. La Plata 35: [5h], fig. cP 1959.

Recent collectors have found this plant growing in old fields
and in fertile soil of low campos. Montes reports that it is
rather abundant in Misiones, Argentina, where it is also used in
popular medicine ("de sabor muy amargo"). Additional vernacular
names (besides those previously reported by me) for the plant are
"berbena" and "verbena", and it has been encountered at 1800 m.
altitude in Salta. It has been collected in fruit (in addition
to the months previously reported) in April, July, and October.
Montes refers to it as an "annual-biennial herb". The corollas
are described as having been "blue" on Aguilar 88, Guimar&es 99,
Ibarrola 1898, J. E. Montes 765, 1719, & & 17321 tee Olea ea 198, T « Ro-

jas 11882, Ru Ruiz Huidobro 2087, | and G. J. Saiieae: 62h, violet" 0 on

Seen, anseanermees cxmetoae re

The Gallinal, Aragone, Bergalli, ee & Rosengurtt 1269,

Herb, Herter 95525, Lindman n A.3651, T. Rojas 1660, and Rosengurtt

A.12h1 & B. B.507, pr previously cited by me as Vv. Titoralis H.B.K.,
seem more . probably to represent V. montevidensis instead. The Re
Alvarez 951 and W, A. Archer 4,788, first cited by me as V. litor-
alis and now as Ve litoralis var. caracasana (H.B.K.) Briq., have
much the general a) appearance of large-spiked V. montevidensis.
Their status needs more intensive investigation.

Troncoso (1965) says of V. montevidensis: "Originaria del Uru-
guay y regién mesopot4mica vecina. Abunda en praderas higréfilas,
borde de zanjas y cercos", She cites from Buenos Aires only Cab-
rera 6298 and Pastore s.n. [Herb. San Isidro 1147], as well as Mac-
bride's type photograph 17432. Schnack and his associates (1959)
report that this plant is apomictic in its reproduction,

Material has been misidentified and distributed in some herbaria
as V. brasiliensis Vell.

Additional citations: LOUISIANA: Avoyelles Par.: J. K. Small s.

372 PHP TO MOGaa Vol. 23, Noss

n. [Burkie, May 1931] (N, N, N, N, Rf). Ouachita Par.: Thomas,
Thomas, & Thomas 708 (N). BRAZIL: Paran4: Guimar&es 99 [Herb. Fac.
rareseiek 5779] (W—2527766); Hatschbach 12075 (Ac), 14731 (N, Rf),
27659 (Ac, N); Stellfeld 1525 [Herb. Fac, Farmacia 5868] | (Ww—
2527750). Rio Grande do Sul: Palacios & Cuezzo 283a (N), 99h (N).
PARAGUAY: Lindman A.3651 (N). URUGUAY: He He '. He Bartlett 20689 (N),
21301 (N); Gallinal, Aragone, Bergalli, Campal, & Rosengurtt 1269
(N); Herb, Herter 95525 (N); Rosengurtt A.12h1 (N), B.507 (N).
ARGENTINA: Buenos Aires: R. Alvarez 311 (N), 453 (N), 609 (N),

7Th6 (N), 779 (N); A. Te Hunziker 2305 5 (W--2595170) ; Nicora 377 (W—
2595171); Tr Troncoso 357 (W—259517). Catamarca: Pierotti "h", in
part (N); Risso 263 (N (N). Chaco: Aguilar 88 (N); T. Rojas 11660
(N), 11882 ~(N). Cérdoba: T. Meyer r 12915 (N). Corrientes: “Tbar-
rola 1898 (N); Ruiz Huidobro 2087 Gian Entre Rios: F. Y. Geof-
froy 18 ( (N). Formosa: I. Morel el 2188 (N), 3072 (N), 3203 3 (N),
3385 (N), 4107 (N), 4982 (N), 5132 (N), 52h3 ( (N), 646 (N). Men-
doza: A. G. ', Schulz 613) (N). Misiones: Krapovickas, s, Cristébal,
Pire, & Tressens 15302 (Ac), 15322 (Rf); J. E. Montes 765 (N), (N),
1237 (Se--130263) , » 1479 (N, N), N), 14731 (Au—-271308) , 27659 (Au—
271283, W—2556187); Ge. G. J. Schwarz 62) (N), 821 (N), 6340 (N).
Salta: Pierotti 1025 (N). Santa Fé: R. Alvarez 973 (N). Tucum4n:
Borsini 901 (N); F. Ortiz 23 (N), sen. [30/5/45] (N).

VERBENA MORICOLOR Moldenke

Synonymy: Glandularia moricolor Schnack & Rubens, Bol. Soc. Ar-
gent. Bot. 13: 205, hyponym. 1970.

Additional bibliography: Moldenke, Phytologia 11: 73. 1965;
Schnack & Rubens, Bol. Soc, Argent. Bot. 13: 205. 1970; Solbrig,
Princ. & Meth. Pl. Biosystem. 76. 1970; Moldenke, Fifth Sum, 1:
201 (1971) and 2: 521 & 917. 1971.

VERBENA MORICOLOR Moldenke x V, PERUVIANA (L.) Britton

Bibliography: Solbrig, Princ. & Meth. Pl. Biosystem. 76. 1970;
Moldenke, Fifth Sum, 2: 917 & 970. 1971.

Solbrig (1970) describes the artificial hybridization of these
two species as well as the hybridization of V. megapotamica
Spreng. with V. peruviana: "Crosses between G[landularia] peruvia-
na and two other morphologically distinct species, G. ies, G. megapotamica
and G. moricolor, on the other hand, yielded hybrids - that were
more fertile. The cross between G. peruviana and G, megapotamica,
for example, was approximately 60 p percent pollen fertile and the
cross between G. peruviana and G. moricolor was approximately 80
percent pollen f fertile. Morphologically, G. megapotamica is simi-
lar to G. peruviana although it has flowers of a different color
and is more erect. Glandularia moricolor, on the other hand, is

quite distinct, having elliptic leaves, erect habit, smaller and

1972 Moldenke, Notes on Verbena 373

deep purple flowers, and a more compact inflorescence. Neither of
these two species grows together with G. peruviana. Glandularia
megapotamica grows in the subtropical gallery forest of northeastern
Argentina, Brazil, and Paraguay, whereas G. moricolor is a species
of the margins of the subtropical forest of northern Argentina.
Consequently, the genetic isolation between these two species and
G. peruviana is not complete. However, they are effectively iso-
lated reproductively because they do nt grow together."

As yet I have not seen any herbarium material as vouchers for
the putative hybrid, and the statement about the similarity in
habit between V, megapotamica and V. peruviana leads me to wonder
about the accuracy of the identification of one or the other of
the two plants used in the crosses. Because of this uncertainty
I have not as yet named the V. moricolor hybrid referred to above.

The hybrid between V. megapotamica and V. peruviana, on the other
hand, has been called xV. schnackii Moldenke.

VERBENA MULTICAULIS Raf.
Additional bibliography: Moldenke, Phytologia 11: 73. 1965;
Moldenke, Fifth Sum. 1: 66 (1971) and 2: 793 & 917. 1971.

VERBENA MULTIGLANDULOSA Moldenke
Additional bibliography: Moldenke, Phytologia 16: 102. 1968;
Moldenke, Fifth Sum, 1: 193 (1971) and 2: 917. 1971.

VERBENA NANA Moldenke

Synonymy: Glandularia nana Schnack & Rubens, Bol. Soc. Argent.
Bot. 13: 205, hyponym. 1o70,

Additional bibliography: Angely, Fl. Anal. Paran., ed. 1, 572.
1965; Moldenke, Phytologia 13: 211. 1966; Angely, Fl. Anal. Fito-
geogr. Est. S. Paulo, ed. 1, : 840 & xix, map 1395. 1970; Schnack
& Rubens, Bol. Soc. Argent. Bot. 13: 205. 1970; Moldenke, Fifth
Summ. 1: "178, 188, & 201 (1971) and 2: 917. 1971.

Rojas describes this species as 30--0 cm. tall and found it
growing on rocky riverbanks in Paraguay.

Additional citations: PARAGUAY: T. Rojas 13872 (N). ARGENTINA:
Formosa: I. Morel 3295 (N), 3754 (N).

VERBENA NEOMEXICANA (A. Gray) Small

Additional & emended synonymy: Verbena canescens var. neo-mexi-
cana A. Gray, Syn. Fl. N. Am., ed. 1, 2 (1): 337. 1878. Verbena
neomexicana Perry ex Moldenke, Fifth Summ, 2: 68, in syn. 1971.
Verbena neomexiczna (A. Gray) Small ex Moldenke, Fifth Sum, 2:
685, in in syn. 1971.

Additional bibliography: Howell & McClintock in Kearney &
Peebles, Ariz. Fl., ed. 2, 726 & 728. 1960; Lewis & Oliv., Am,
Journ. Bot. 8: [639] & 641, fig. 18. 1961; Hocking, Excerpt. Bot.
A.6: 91. 1963; Moldenke, Phytologia 16: 189 & 200. 1968; Whitta-
ker & Niering, Journ, Ecol. [Brit.] 56: 528. 1968; Bolkh., Grif,
Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 717. 1969; Rickett,

37h P PY TO Lover Ith Vol. 23, no.

Wild Fils. U. S. 3 (2):°365, [367], & 551, pl. 111. (1969) and \
(3): 540, [543], & 799, pl. 177. 1970; Correll & Johnston, Man.
Vasc. Pl. Tex. (Contrib. Tex. Res. Found. Bot. 6:] 1876 & 1877.
1970; Moldenke in Correll & Johnston, Man. Vasc. Pl. Tex. {[Con—
trib. Tex. Res. Found. Bot. 6:] 1315 & 1321. 1970; Moldenke, Fifth
Summ’. 12° 20° 30, 59,''62, 63, 66, °76, 371, & 396 (1971) ‘amd 2659.
672, 683--€85, 687, 705, 707, & 917. 1971; Moldenke, Phytologia
22: 485 & 500 (1972) and 23: 192, 236, 2h2, & 302. 1972.

Illustrations: Rickett, Wild Fls. U. S. 3 (2): [367], pl. 111
(in color) (1969) and (3): [543], pl. 177 (in color). 1970.

Recent collectors describe this plant as erect, becoming ) in-
ches to 1 foot tall about bushes, growing in deserts, in roadside
washes, and on silty slopes. Stewart found it "common on dry
hillsides" in Coahuila, Mexico. Kruckeberg found it to be "com-—
mon" in campground areas in oak-pinyon-juniper associations in
Brewster County, Texas. Whittaker & Niering (1968) found it grow-
ing "to some extent" on limestone in southeastern Arizona, while
in the same state Howell & McClintock (1960) describe it as "com-
mon" on foothills and in canyons at altitudes of 2000 to 6000
feet, flowering from March to October.

In addition to months previously reported by me, it has been
collected in fruit in April and June. The corollas are described
as having been "lavender" on Kruckeberg 4,796, "blue" on R. M.
Stewart 402, and "purple" on Johnston & Muller 1057 & 1067 and
Powell, Turner, & Magill 2063.0 eoly-

~~ The Ki. Mite: Mitchell sen. [h, 1/14 14/37] distributed as V. neomexicana
is actually V. bracteata Lag. & Rodr.; Atwood 2008, Ballinger s.n.
[October 2), 1959], and C. M. Rowell 5113 are Ve c canescens var.
roemeriana (Geese) Perry; “A. Davis eos Veal gooddingii Briq.; We
W. Jones s.n. [Aug. 10, 1926] is Ve gracilis Desf.; Leverich 22,
E. Marsh 29, B. Pitiaian 36, and oF M. Rowell 11635 are V. neomex-
icana var. hirtella Perry; W. W. Jones Son. Sen. [Cabullona], “Krucke—
berg 1.928 berg 928 (in part), and S. S. White 3310 are V. neomexicana var.
xylopoda - Perry; Kruckeberg 2 928 (in part) is V. menthaefolia
Benth.; C. H. Muller 821) and C. M. Rowell 1117 are V. perennis
Wooton; Pennington 7h is V. pinetorum m Moldenke; and Meebold 26696
is V. racemosa Eggert.

Additional citations: TEXAS: Brewster Co.: Kruckeberg 796
(N); C. H. Mueller 8138 (Lk). Jeff Davis Co.: Tharp & Janszen h9-
11); (Au--122363). NEW MEXICO: Grant Co.: E. L. Greene s.n. [Aug-
ust 11, 1880] (W--2606263), s.n. [Pinos Altos Mtns., August 23,
1880] (Pa). MEXICO: Chimuahaa: | Powell, Turner, & Magill 2063 (Au-
296918). Coahuila: Johnston & Muller - 193 3 (Au---301900, Mi), 1 1057
(Au-~301851), 1067 (Mi); R. Me Stewart 402 (Au--301),26) .

VERBENA NEOMEXICANA var. HIRTELLA Perry

Additional synonymy: Verbena neomexicana hirtella Perry apud
Correll & Johnston, Man. Vasc. Pl. Tex. (Contrib. Tex. Res. Found.
63] 1876. 29705

1972 Moldenke, Notes on Verbena 375

Additional & emended bibliography: Lewis & Oliv., Am. Journ.
Bot. 48: [639]—6h1, fig. 18. 1961; Hocking, Excerpt. Bot. A.6: 91
(1963) and A.9: 365. 1965; Moldenke, Phytologia 16: 189. 1968;
Moldenke in Correll & Johnston, Man. Vasc. Pl. Tex. (Contrib. Tex.
Res. Found. Bot. 6:] 1315 & 1321. 1970; Correll & Johnston, Man.
Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:] 1876. 1970; Mol-
denke, Fifth Summ. 1: 20, 59, 62, 63, & 76 (1971) and 2: 683, 685,
& 917. 1971; Molcenke, Phytologia 23: 192 & 242. 1972.

Emended illustrations: Lewis & Oliv., Am. Journ. Bot. 8: 60,
fig. 18. 1961.

Recent collectors have found this plant growing among limestone
rocks. Youngblood calls it an "herb locally frequent in rocky
sand" in Brewster County, Texas. The corollas are described as
having been "purple" on D. Youngblood 4o and "blue-purple" on
Leverich 22. Material has been misidentified and distributed in
some herbaria under the name V. neomexicana Perry. On the other
hand, the J. H. Thomas 8185, distrivuted as V. neomexicana var.
hirtella, is (in at le at least some herbaria) actually V. canescens
var. roemeriana (Scheele) Perry, as is also G. L. Webster 453,
while Tharp & Janszen 49-11), is typical V. neomexicana (A. Gray)
Small.

Additional citations: TEXAS: Brewster Co.: Leverich 22 (Lk) ; E.
Marsh 29 (Au—21))01); E. G. Marsh 68 (Au—212535) ; B. Pittman Pittman
36 (Lk); C. M. Rowell 11635 (Lk); D. Youngblood 40 (Lk). Jeff Da-
vis Co.: Tharp & & Janszen 49-114) (N). MEXICO: Durango: Johnson &
Johnson 1718 (Ws).

VERBENA NEOMEXICANA var. XYLOPODA Perry

Additional bibliography: Howell & McClintock in Kearney &
Peebles, Ariz. Fl., ed. 2, 728. 1960; Moldenke, Phytologia 16: 102
& 103. 1968; Rickett, Wild Fls. U. S. 4 (3): [543], pl. 177. 1970;
Moldenke in Correll & Johnston, Man. Vasc. Pl. Tex. (Contrib. Tex.
Res. Found, Bot. 6:] 1315 & 1321. 1970; Correll & Johnston, Man.
Vasc. Pl. Tex. [Contrib. Tex. Res. Found. Bot. 6:] 1877. 1970;
Moldenke, Fifth Summ. 1: 30, 59, 63, 66, & 76 (1971) and 2: 659,
685, 707, & 917. 1971; Moldenke, Phytologia 23:. 302. 1972.

Tlustrations: Rickett, Wild Fls. U. S. h (3): (543), pl. 177
(in color). 1970.

Recent collectors describe this plant as a perennial herb, 1 m.
tall, and found it growing on ridges, in sandy pockets of clay
flats, in juniper-grassland associations, anc in riparian habitats.
Kruckeberg found it in openings of woodlands consisting of Quercus
arizonica, Q. hypoleuca, and Pinus leiophylla. Howell & McClin-
tock (1960) describe it as having | "slightly glandular inflorescence
and a larger corolla than [the] typical form" and note that "This
is the chief Arizona representative of the species". The corollas
are said to have been "blue" on S. S. White 3099 & 3310, "lavender-

—_——

Moran describes the | plant ae "scarce".

376 P Heer LioiG Tf A Vol. 23, no. 4

Material of this variety has been misidentified and distributed
in some herbaria as V. xutha Lehm. On the other hand, the R. Run-
yon eo. distributed as V. neomexicana var. xylopoda, is Vu
cloverae Moldenke, while Re, Runyon 4869 seems to be a mixture of
V. cloverae and ve plicata | Greene.

Additional citations: ARIZONA: Cochise Co.: Pinkava, Keil, &
Lehto 14553 (N, N). Yavapai Co.: W. S. Lewis s.n. (Herb. BL Blake
956] (Ld). MEXICO: Baja California: Gentry & E Fox Fox 11735 (Mi); R.
V. Moran 14592 (Sd—-68132), 15095 (Sd - Chihuahua: Kruck-
eberg 4923 3 (N); Townsend & Barber 192 saree oan Sonora: W. 2 W. We
Jones s.n. [Cabullona] Tiona] (Sd--1133); S$ S. S. White 3099 (Sd—61803),
3310 (Sd--6180)). Tamaulipas: Stamford, |, Lauber, & “& Taylor 2381
(Se--14,7631).

XVERBENA NEQUAM Moldenke

Additional bibliography: Moldenke, Phytologia 11: 73. 1965;
Moldenke, Fifth Summ. 1: 371 (1971) and 2: 521, 666, 678, 700, &
Saly (ponte aly

xVERBENA NISA Moldenke

Additional bibliography: Moldenke, Phytologia 11: )73—7h.
1965; Moldenke, Fifth Summ, 1: 371 (1971) and 2: Bea. 689, 700, &
salto dlsfabe

VERBENA NIVEA Moldenke

Additional bibliography: Moldenke, Phytologia 13: 211--212.
1966; Moldenke, Fifth Summ. 1: 18) &’201 (1971) and 2: 685 & 917.
1971.

VERBENA NIVEA f. ROSEA Moldenke
Additional bibliography: Moldenke, Phytologia 13: 212. 1966;
Moldenke, Fifth Summ, 1: 201 (1971) and 2: 685 & 917. 1971.

XVERBENA NOACKI Moldenke
Additional bibliography: Moldenke, Phytologia 10: 192. 196;
Moldenke, Fifth Summ. 1: 371 (1971) and 2: 673, 67h, & 917. 1971.

XVERBENA NOTHA Moldenke
Additional bibliography: Moldenke, Phytologia 11: 7h. 1965;
Moldenke, Fifth Summ, 1: 371 (1971) and 2: 672, 685, & 917. 1972.

VERBENA OCCULTA Moldenke

Additional bibliography: J. F. Macbr., Field Mus. Publ. Bot. 13
(5): 61h, 615, & 627. 1960; Moldenke, Phytologia 11: 71. 1965;
Moldenke, Résumé Suppl. 16: 5 (1968) and 17: 3. 1968; Anon., Biol.
Abstr. 50 (3): B.A.S.1.C. S.201. 1969; Moldenke, Biol. Abstr. 50:
1493. 1969; Hocking, Excerpt. Bot. A.15: 422. 1970; Moldenke,
Fifth Summ, 1: 14) (1971) and 2: 917. 1971; Moldenke, Phytologia
22: 466 (1972) and 23: 28). 1972.

Recent collectors have found this plant growing in rocky-sandy

1972 Moldenke, Notes on Verbena 377

soil, flowering in March, June, and July (in addition to the
months previously reported by me), and fruiting in August. They
describe it as a shrub 3 feet tall. Fosberg calls it a "low much
branched herb, common on jalca or mountain meadow". The corollas
are described as having been "blue" on Edwin & Schunke 3692,
"violet" on Aguado 6761, "white-violet" on Lépez Guillén 3787, and
"white to purple" on F. R, Fosberg 28192.

Material has been misidentified and distributed in some herbaria
as V. berterii (Meisn.) Schau. and V. laciniata (L.) Briq. The
Ferreyra 6586, cited by Macbride (1960), is the type collection of
f. alba Moldenke.

Additional citations: PERU: Amazonas: Edwin & Schunke 3692 (N);

ukup 4960 (W--2565565). Cajamarca: F. R. R. Fosberg rg 20192 (Ac).
i aig Aguado 6761 (Tp). Lima: Lépez Guillén 3787 ("3387")
Ac

VERBENA OCCULTA f. ALBA Moldenke

Additional bibliography: J. F. Macbr., Field Mus. Publ. Bot. 13
(5): 627. 1960; Moldenke, Phytologia 10: 19). 196); Moldenke,
Fifth Summ. 1: 1h) (1971) and 2: 917. 197le

Vargas Calderén found this plant growing at 2000-2600 m. alti-
tude, in flower and fruit in October.

Additional citations: PERU: Ancash: Vargas Calderén 10276 (W—
2520241).

VERBENA OCCULTA f. AURANTIACA Moldenke

Bibliography: Moldenke, Phytologia 17: 3h. 1968; Moldenke, Ré-
sumé Suppl. 17: 3. 1968; Anon., Biol. Abstr. 50 (3): B.A.S.1.C.
S.201. 1969; Moldenke, Biol. Abstr. 50: 1493. 1969; Hocking, Ex-
cerpt. Bot. A,15: 22. 1970; Moldenke, Fifth Summ. 1: 14) (1971)
Sgee olf. LOT.

Citations: PERU: La Libertad: Angulo 1383 (Z--type).

VERBENA OFFICINALIS L.

Additional & emended synonymy: Verbena vulgo appellata Anguill.,
Sempl. 266. 1561. Verbenaca volgare del Matthioli Pona, Pl. Simp.
Bald. Mont. 3. 1595. Verbenaca recta Doi Dodon., Stirp. Hist. Pemptad.,
ed. 2, 150. 1616. Verbena lusitanica, latifolia, procerior
Tourn., Compl. Herb. 357.6 LPLOS Tans urticae-folia, canadensis,
foliis incisis, flore majore Tourn., Compl. Herb. 357. 1719. Ver-
bena communis ae flore Tourn, ex Manetti, Virid. Florent. oes
1751. Verbena vulgaris Hill, Brit. Herb. 356 & [536]. 1756. Ver-
bena foliis tripartitis, rugosis, spicis nudis gracillimis Haller,
Hist. Stirp. Ind. Helv. 1: 96--97. 1768. Verbena tetrandra spicis
filiformibus paniculatis, foliis multifido laciniatis, caule soli-
tario L. ex Haller, Hist. Stirp. Ind. Helv. 1: 96. 1763. Verbenaca
Camer. apud J. E. Sm., Engl. Fl., ed. 1, 3: 71, in syn. 1825.
Tetrandra, spicis filiformibus paniculatis, foliis multifido-

378 P ay? OD Oo A Vol. 23, no.

laciniatis caule solitario Willd, ex Lanfrossi, Giorn. Fis. Chim.
Stor. Nat. Med. & Art., ser. 2, 10: 48. 1827. Verbena communis
caeruleo flore (foliis dissectis) Micheli ex Bertol., Fl. Ital.
6: 261, in syn. 184). Verbena communis caeruleo flore Zannich.
ex Bertol., Fl. Ital. 6: 261, in syn. 184). Verbena vulgo appel-
lata Ang. ex Bertol., Fl. Ital. 6: 260, in syn. 13h. Verbenaca
volgare del Matthioli Pon. ex Bertol., als Ital. 6: 260, in syn.
18h). Verbena communis, flore caeruleo Cip. ex Bertie, Fl.
Ital. 6: 260, in syn. 13h. Verbena n. 219 Haller ex Bertol., Fl.
Ital. 6: 261, in syn. 184). Verbena spura L. ex Jacobs & Burlage,
Ind. Pl. N. C. 251, sphalm. 1958. Verbina officinalis L. ex R. I.
Patel, For. Fl. Melghat 353, sphalm. 1968. Verbena offinalis
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Abstr. 35: 180. 1965; Hocking, Excerpt. Bot. A.9: 365. 1965;
Angely, Fl. Anal. Paran., ed. 1, 570. 1965; Pitschm. & Reisigl,
Fl. Stidalp. 189. 1965; Arevalo, Fl. Espafi. 82. 1966; Hirata,

Host Range & Geogr. Distrib. Powd. Mild. 277. 1966; Bouchet &
Andy in Fryer, Eighth Brit. Weed Control Conf. Proc. 1: 31.

1966; Polunin & Huxley, Fls. Medit. 154. 1966; Airy Shaw in J.C.
Willis, Dict. Flow. Pl., ed. 7, 1176 & 1178. 1966; Hunt, Hilte-
brand, Schmid, Groger, Johne, & Mothes, Experimentia 22 (10): 656,
1966; Yotaro, Gard. Pl. World 3: 127. 1966; Hara, Fl. East. Himal.
29. 1966; Kalm, Travels N. Am., ed. Benson, pr. 2, 1: 6. 1966;
Yamazaki in Hara, Fl. East. Himal. 270. 1966; Datta & Majumdar,
Bull. Bot. Soc. Bengal 20: 103. 1966; Patzak & Rech. in Rech., Fl.
Iran. 43: 1—2 & 8. 1967; Zukowski in Pawlowskiego, Fl. Polsk. 11:
64--65. 1967; Huni, Hiltebrand, Schmid, Groger, Johne, & Mothes,
Biol. Abstr. 48: 7828. 1967; Thommen, Taschenatl. Schweiz. Fl.
177, fig. 2198. 1967; Rendle, Classif. Flow. Pl., ed. 2, 2: 501 &
505. 1967; Gilkey & Dennis, Handb. Ni. Pl. 353. 1967; Mitra, Elem.
Syst. Bot. Angiosp., ed. 2 abridg., 139. 1967; R. R. Stewart,
Pakist. Journ. Forest. 17: 515. 1967; Wherry, Bartonia 37: 13.
1967; R. K. Gupta, Season. Fls. Ind. Sum. Resorts Moos. 81, 98,
114, 132, & 21. 1967; Esfandiari, Une Prem. List. Pl. Herb, Min-
ist. Agr. Iran 252 & 253. 1967; Panigr. & Saran, Bull. Bot. Surv.
India 9: 258. 1967; Hsu, Raiwania 13: 121 & 125, fig. 27. 1967;
Abrams, Illustr. Fl. Pacif. States, pr. 2, 3: 610 & 611. 1967;
Tingle, Check List Hong Kong Pl. 38. 1967; H. C. D. de Wit, Pl.
World High. Pl. 2: 183. 1967; L. & M. Milne, Living Pl. World

38h P Havel 0) LeOnG sek Vol. 23, no.

212. 1967; Grove, Biblioth. Mycol. 2: 1h. 1967; H. Ward, Herb.
Man., ed. 2, 107--108. 1967; Ferrdndiz, Guia Med. Veg. 388—389.
1967; J. M. Watt, Lloydia 30: 1-22. 1967; Ismailov, Rest. Resur.
3: 53-57. 1967; Alzate, Pl. Medic., ed. 7, 26l--265. 1968; Mol-
denke, Résumé Suppl. 16: 5 & 10-—-12 (1968) and 17: 7. 1968; H. J.
M. Bowen, Fl. Berks. 237. 1968; Peterson & McKenny, Field Guide
Wildfls. 286, [287], & 418. 1968; Carter & Jones, Castanea 33:
203. 1968; Moldenke, Biol. Abstr. 9: 697. 1968; Moldenke, Phy-
tologia 16: 189--192 & 31. 1968; Jaitly, Guignard, & Mestre,
Compt. Rend. Seanc. Acad. Sci. Paris 267: 59. 1968; Uphof, Dict.
Econ. Pl., ed. 2, 51 & 52. 1968; Anon., Rivist. Ital. 50: [516].
1968; Sakan & Abe, Tetrahedron Lett. 1968: 2)71--2)73. 1968;
Hocking, Excerpt. Bot. A.13: 571. 1968; W. T. Pope, Wayside Pl.
Hawaii 19 & 289. 1968; Kapoor, Bull. Bot. Surv. India 10: 2.
1968; Clapham, Tutin, & Warburg, Excurs. Fl. Brit. Isles, ed. 2,
338--339. 1968; Hendrych, Act. Univ. Carol. Biol. 2: 188. 1968;
Stecher, Merck Index, ed. 8, 110). 1968; Deb, Sengupta, & Ma-
lick, Bull. Bot. Soc. Bengal 22: 199. 1968; Munz & Keck, Calif.
Fl. 686, 687, & 1679. 1968; Paton, Wild Fls. Cornwall 36. 1968;
R. I. Patel, For. Fl. Melghat 271 & 353. 1968; Munz, Suppl. Cal-
if. Fl. 101. 1968; Anon., Hort. Abstr. 39: 872. 1969; M. Martinez,
Pl. Med. Mex., ed. 5, 340-341 & 579. 1969; Battersby, Hall, &
Southgate, Journ. Chem, Soc. Lond., sec. C, 5: 721—728. 1969;
Horodysky, Waller, & Eisenbraun, Journ. Biol. Chem. 2h): 3110—
3113 & 3115. 1969; S. P. & R. N. Banerjee, Bull. Bot. Soc. Ben-
gal 232 166.91969; Fior, Nost. Fle, ede, 3,°570, fig. (2s 19en,
Bolkh., Grif, Matvej., & Zakhar., Chrom. Numb. Flow. Pl. 717.
1969; Sakan & Abe, Biol. Abstr. 50: 452. 1969; Hansen, Bol. Mus.
Munic. Funchal 2): 3). 1969; Farnsworth, Blomster, Quimby, &
Schermerh., Lynn Index 6: 267. 1969; Inouye, Ueda, Aoki, & Take-
da, Tetrahedron Lett. 1969: 2352-235). 1969; Polunin, Field
Guide Fls. Eur. 32—3)3 & 625. 1969; Hussain, Pakist. Journ.
Forest. 19: 342 & 347. 1969; Anon., Hortic. Abstr. 39: 872. 1969;
Tobe, Proven Herbal Remed. 160 & 297. 1969; Moldenke, Biol. Ab-
str. 50: 128. 1969; Anon., Biol. Abstr. 50 (1): B.A.S.1.C. S.19
1969; M. A. Rau, Bull. Bot. Surv. India 10: Suppl. 2: 63. 1969;
A. Pedersen, Bot. Tidsskr. 6): 317, [341--32], 347--349, 355--
356, & 379, fig. 50. 1969; Hocking, Excerpt. Bot. A.lh: 206.
1969; Miall, Early Naturalists, pr. 2, 25. 1969; Benchley, Cour-
ier-News [Plainfield, N. J.J], p. 11, Jan. 9. 1970; Angely, Fl.
Anal. Fitogeogr. Est. S. Paulo, ed. 1, : 825, 839, & xix. 1970;
Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tes. Res.
Found. 6:] 1877. 1970; Moldenke in Correll & Johnston, Man. Vasc.
Pl. Tex. (Contrib. Tex. Res. Found. Bot. 6:] 1318. 1970; Farns-
worth, Pharmacog. Titles 5 (2): viii & item 1429 (1970), 5 (3):
vii & item 231 (1970), 5 (7): ix & item 7248 (1970), 5 (10):
xxiv & item 12255 (1970), and 5 (11): xvii & item 15121. 1970;
Britton & Br., Illustr, Fl., ed. 2, pr. 5, 3: 94 & 95, fig. 3552.
1970; R. J. Moore, Reg. Veg. 68: 71. 1970; Anon., Biores. Index
6: 4151. 1970; "A. R.", Biol. Abstr. 51: 13600. 1970; Daoud &
Sheikh, Bull. Coll. Sci. Univ. Baghdad 11 (2): 2h—lh. 1970; E.
L. D. Seymour, New Gard. Encycl., ed. 8, 1279. 1970; Dihoru & Do-

1972 Moldenke, Notes on Verbena 385

nita, Fl. Si Veget. Podis. Babad. 68. 1970; Domville & Dunbar,
John Burroughs Nat. Hist. Soc. Bull. 8: 9). 1970; Horodysky, Wal-
ler, & Eisenbraun, Biol. Abstr. 51: lS. 1970; Willaman & Li,
Lloydia Suppl. 33 (3a): 220. 1970; Anon., Biol. Abstr. 51 (8): B.
A.S.1.C. S.222 (1970) and 51 (2h): BASIC. S.238. 1970; El-
Gazzar & Wats., New Phytol. 69: 483 & 485. 1970; Catanzaro, Fito-
terapia 1: 83. 1970; Sykes, N. Zeal. Dept. Sci. & Indust. Res.
Bull. 200: 105, 216, & 31h. 1970; Macer, Virib. Herb., ed. 2, pr.
2, (86]--[88]. 1970; N. F. Good, Biol. Abstr. 52: 2515. 1971;
Rickett, Wild Fls. U.S. 5 (2): [455], 456, & 665, pl. 152. 1971;
Priszter, Delect. Sem. Spor. Pl. Hort. Bot. Hung. 59 & 81. 1971;
Wiggins & Porter, Fl. Galdp. Isls. 997. 1971; Moldenke in Wiggins
& Porter, Fl. Gal4p. Isls. 504. 1971; Farnsworth, Pharmacog.
Titles 6 (3): xvi & item 3969 (1971), 6 (5): xiv & item 7687
(1971), and 5, Cumul. Gen. Ind. 1971; A. Love, Taxon 20: 353.
1971; Inouve in Wagner & Horhammer, Pharmacog. & Phytochem. 1971;
Brown & Wherry, Bartonia 0: 13. 1971; Anon., Biol. Abstr. 52
(5): BoA.S.I.C. S.128 & S.238. 1971; Guillarmod, Fl. Lesotho 235.
1971; Cadbury, Hawkes, & Readett, Comput.-map. Fl. Warwicksh. 197.
1971; Moldenke, Fifth Summ. 1: 16--23, 25, 27, 30, 32, 33, 39,
ha, ho, 62, 6h, 66, 76, 92, 120, 178, 18, 193, 203-211, 213—
216, 231, 23h, 238, 2hi, 2h6, 2h8, 257, 264-267, 269-271, 278,
Calpe 90, 292, 29,298, ‘303, 308, 311-313, 318, 3285°337, 3h9,
350, 352, 371, 376, 389, 468, & h71 (1971) and 2: 527, 59h, 60h,
618, 648, 619, 653, 662--664, 666--670, 672--675, 679, 680, 682,
686--688, 692—69, 696, 697, 699, 701, 702, 704, 706, 707, 709,
789, 793, & 917--918. 1971; Moldenke, Phytologia 23: 186, 192,
193, 261, 263, 26h, 269, 291, & 295. 1972; A. L. Moldenke, Phyto-
tseia) 232318. 1972.

Additional & emended illustrations: Brunf., Herb. Viv. Icon.,
ed. 1, 1: 119 (1530) and ed. 2, 1: 119. 15325 Fuchs, Hist. Plant.
Basil. 593. 1542; Durante, Herb. Nuov., ed. Rom., 69. 1585;
Matth., Disc. Valgr., ed. 1, 2: 399 & 1107. 1585; Camer. in
Matth., Pl. Epit. Util. 797. 1586; Matth., Herb. Aneb Bylinar 380
(in color). 1596; Gerarde, Herb., ed. 1, 580, fig. 1. 1597; Do-
don., Stirp. Hist. Pemptad., ed. 2, 150. 1616; Gerarde, Herb.,
ed. 2, 718, fig. 1. 1633; Lonic., Kreuterb., pr. 1, 310. 1679;
Rivin., Ord. Pl. Irreg. Monop. 81, pl. 56. 1690; Tourn., Inst.
Rei Herb., ed. 2, 2: pl. 9 (1700) and ed. 3, 2: pl. 9h. 1719;

We Curtis, Fl. Lond., ed. 1, 1 (1): pl. bk ("2") (in color).
1775; O. F. Mull. in Oeder, Icon. Pl. Fl. Dan. : 5, pl. 628 (in
color). 1775; Bulliard, Herb. France pl. 215. 1785; G. F. Hoffm.,
Deutsch. Fl. 9: pl. 2. 1791; Sowerby & J. E. Sm., Engl. Bot.,

ed. 1, 11: pl. 767 (in color). 1800; Quensel in Palmstruch,

Svensk Bot. 3: pl. 182 (in color). 180); W. Curtis, Fl. Lond., ed.
2, 1 (1): pl. 1. (in color). 1817; Alberti, Fl. Med. : pl. 228.
1317; J. E. Sm., Gramm. Bot. pl. 1, fig. 17). 1821; W. Curtis,
Fl. Lond., ed. 2, 3: pl. 90 (in color). 1835; E. Twining, Ill.
Nat. Ord. Pl. 106, fig. 1 (in color). 1855; Reichenb., Icon. Fl.
Germ. 18: pl. 91 (1292, II).1857; Boswell Syme in Sowerby, Engl.
Bot., ed. 3, 6: pl. 1018 (in color). 1866; Thomé, Fl. Deutschl. :
pl. 531 (in color). 1889; Paque, Fl. Anal. & Descr. Prov. Namur &

386 PHY DOLOGIA Vol. 23, nosis

Luxemb. 47, fig. 29). 1902; H. Wagner, Illustr. Deutsch. Fl., ed.
5, 515% 1905; Hatton, Craftman's Plant-Book & 369, fig. I & 729.
1909; Britton & Br., "T1lustr. Fl, ed. 2, ipre sh, 3295, fie pessoas
1913; L4zaro e Ibiza, Comp. Fl. Espafi., ede 33 3: 296, figi 6732
1921; Bouloumoy, Fl. * Liban & Syrie pl. 320, fig. 3. 1930; Britton
& Br., TMlustrs Fis eds $2. pr sie: 32.98; fig. "3060, 1936; Noack,
Biol. Zentralbl. 57: 386, fig. Ba "1937; F.H. & H. H. Hillman,
Seed Trade Buyers Guide 1938: pl. 12, fig. 4. 1938; Britton & prs,
Tlustre Fl (cedet2, presi 321195, figs) 3552 (19435 and ed. 2, pr.
lh, 3: 95, fig. 3552. 197; Metcalfe & Chalk, Anat. Dicot. 103k,
fig. 2h7 L. 1950; M. Martinez, Pl. Med. Vex., ed. h, 30. 19593
Emberger in Chadefaud & Emberger, Traité Bot. 2: 827 & 828, fig.
1173b & 117). 1960; Hatton, Handb. Pl. & Flor. Ornament ) & 369,
fig. I & 729. 1960; Buia, Fl. Mic. Tllustr. Rep. Pop. Rom. 01,
fig. 368. 1961; Uaeeiee & Boselt, Unsere Heilpfl. [452]. 1962;
Lonic., Kreuterb., PPG 4 6310 1962; Brownlow, Herbs & Frag.
Gard., ed. 2, pl. 10 (in’ color). 1963; Foureroy, Atlas Recon. Dir.
Ple* Plus Commun., new ed., pl. 158 [bottom]. 1963; lielchior in
Engl., Syllab. Pflanzenfam., ed. 12, 2: 436, fig. 18, A&E.
1964; Heimans, Heinsius, & Rhijsse, Geillustr. Fl. Nederl. 907.
1965; Ferr4ndiz, Guia Mod. Veg. 388. 1967; Hsu, Taiwania 13: 121,
figedene LION Thommen, Taschenatl. Schweiz. Fl. 177, ‘figs (2198
1967; P Peterson & McKenny, F Field Guide Wildfls. [287]. 1968; M.
Martinez, Pl. Med. Mex., ed. 5, 30. 1969; Fior, Nost. Fl., ed.
Bot, fig. 712. 1969; Rickett, Wild Fls. U. S. 5 (2): [h55],
ple 153 (in color). 1971.

Recent collectors have found this plant growing in sandy soil,
disturbed areas, waste ground, and clearings in evergreen forests.
They describe it as a typical ruderal plant. Koelz refers to it
as 18 inches to 5 feet tall. Manning (1965) describes its fruit
as consisting of four reddish-brown l-seeded nutlets each weigh—
ing about 0.000) gm., dispersed in mud or (in the calyx) adhering
to the coats of animals or to clothing. Bolkhovskikh and his as—-
sociates (1969) point out that the diploid number of chromosomes
for Verbena officinalis is given as 12 by Schnarf (1923) and by
Patermann (1935), but as 1) by Junell (193), Tischler (193h),
Dermen (1936), Noack (1937), and Schnack & Covas (19h). Hsa
(1967) says that at diakinesis there are seven bivalents.

The color of the corolla is given as "pale-lilac" by Paque
(1902) and Sauzé & Maillard (1872), "rose" on Koelz 15757, "rose-
lavender" on Koelz 161), "rose-purple" on feel 16839, " “purple-
blue" on Beauchamp 1178, "pale-blue" on E. H. Walker 8101, "Light—
purple" on Banerjee & Shakya 5596, maura o on E. A. Robindon
5596, "pluish-purple" on on A. | Smith 211, "purple" on nR. E. Cooper
5045 and A. Smith 50 and by Gupta 11967), "yiolet" by Deb, Sen-
gupta, & Malick (1968), "lilac" by Bouloumoy (1930), pacer
(1965), and Patzak & Rechinger (1967), and "white with bluish
tinge" by Kapoor (1968).

The time of blooming is given as July by Gattinger (189) in
Tennessee, "May to Autumn" by Tornabene (1891) in Italy, June to
October by Polunin (1969) in Europe, by Sauzé & Maillard (1880)

1972 Moldenke, Notes on Verbena 387

in Belgium, and by Heimans, Heinsius, & Thijsse (1965) in the
Netherlands, June to September by Martin (1965) in England and
Wales and by Zukowski (1967) in Poland, June to November by Ohwi
(1965) in Japan, July to September by Tatnall (1946) in Delaware,
Sauzé & Maillard (1872) in Belgium, Dietrich (182) in Germany,
Martens & Kemmler (1882) in Wirttemberg, and Thomé (1888) in Ger-
many, July to October by Druce (1897) in England, March to June
by Datta & Majumdar (1966) in India, and April to October by Ma-
heshwari (1963) in India.

Additional vernacular names applied to this plant, in addition
to the many previously reported by me, are "akicamiyu gusa",
"akimmayo", "akitsumayo", "beneran", "bé-pin-chh4u", "birbina",
"blood of Mercury", "briina", "brivina", "bukexrem", "bunj",
"colombaria", "Dankbarkraut", "devil's bane", "devil's hate",
"Eisenrich", "Eiserisch", "enchantment herb", "erba bona", "erba
colombina", "erba croce", "erba crocetta", "erba crocina", "erba
medica", "erba sacra", "erba S. Giovanni", "Zuropean verbena",
"gebrauchlicher Eisenhart", "gemeiner Eisenhart", "gogerchin otu",
"herbal vervain", "herba sacra", "herba veneris", "hierba de
todos los males", "holy plant", "horse-whip", "Tgenkraut",
"karata", "kuma-tsu-zura", "kumatuzura", myainnlin" , "ma tién
th4o", "officinis", "peristereon", "peristereum", "simplers! joy",
"spike vervain", "spurious vervain", "Taubenkraut", "tears of
Juno", "té-bfi-chh4u", "thin-bé-pin", "verbena oficinal", "verbenae
herba", "vermena", "vermenaca", "verminaca", and "verveine sauvage".

Stewart (1967) informs us that this species is a very common
weed in lower Swat, Pakistan. Carleton (1962) records the addition-
al vernacular names "countryman's treacle" [a name also applied to
Ruta graveolens], "hyssop" [also applied to Gratiola officinalis,
Hyssopus officinalis, and Teucrium pseudohyssopus], "tears-of-
Isis", and "tears-of-Juno". He tells us that the name "herb-
grace", applied to Verbena officinalis, is likewise applied to
Ruta graveolens, the name "shrubby—verbena" is applied also to
members of the genus Lantana, and "sweet-scented-verbena" to
Aloysia triphylla (L'Hér.) eee: Miall '(1969) states that
"primitive medicine gave the name of.....Verbena to Verbena of-
ficinalis and to the groundsel". Hatton (1909) reminds us that
to the ancients "Verbena was the name used for a sacrificial
herb, whatever it was". Polunin & Huxley (1966) report that "Hip-
pocrates considered this to be one of the few all-curing herbs".

Uphof (1968) avers that Verbena officinalis is an "Annual or
perennial herb. Origin uncertain, probably from the Mediterran-
ean region, escaped throughout Europe, Africa, Asia, Australia, N.
and S, America. Was in ancient times well known. Was called in
ancient Egypt 'Tears of Isis'. The entire plant, Herba Sancta,
was burned during ceremonies. At present Herba Verbenae is used
as astringents, diuretic, diaphoretic; for its action on the uter-
us and as emmenagogue. Externally employed for wounds that are
difficult to heal, ulcers and skin ailments."

Beston (1950) informs us that "There grows in the garden here
a very ancient plant which has been a part of the magic and relig-

388 Pi H Se vO (LcOuG iia Vol. 23, nos k

ion of the most diverse European cultures, in England being a sa-
cred plant of the Druids, in Scandinavia a plant of the priests
of Thor, in Greece and Rome a plant so holy that no other might
be used to brush the altars of Olympian Jove. Yet it is nota
stately plant but a quite simple one, almost a thing of the road-
sides and the fields. 'Hierobotane' the Greeks called it, the
'Holy Plant'; and *Herba Sacra! the Romans. To us it is Spike or
Herbal Vervain, the Verbena Officinalis of Linnaeus. Whatever
list of annuals I make for the garden, this ancient herb is sure
to be written down. It has enough garden presence of a rustic
kind to justify its inclusion, being in no way boorish or uncivil,
and it is easy to start from seed and easy to grow....Seed....is
still used in domestic medicine as a help in the early stages of
colds and fevers.,...to be had....only from European firms. The
herb is not particular about soil but likes a sunny place, and I
have put my plants well back in the border, planting them in a
close-growing line, and facing them with some shorter and more
compact perennial. To those interested in magic and religion,
there is no herb in the garden more worthy of attention, for
this simple plant without fragrance, without an outer look of
power, without a flower of significance, was singled out from
among all other plants and herbs as the most sacred of the grow—
ing things of earth between the Pillars of Hercules and the
roots of the Caucasus."

Clair (1961) gives us another fascinating account of this

plant:
"Bring your garlands, and with reverence place
The vervain on the altar. BEN JONSON

Strange that a herb of such insignificant appearance should have
held so important a place in mankind's estimation through long
centuries; but it was a sacred and magic herb in Persia, in an-
cient Greece and Rome, and in ancient Britain, where the Druids
had a special reverence for it, ordering it to be gathered about
the rising of the great dog star, but only when neither the sun
nor moon were above ground at the time to see it. Moreover, those
who uprooted it were expected to place upon the spot where it had
grown honey in the comb to make amends for having deprived the
earth of so holy a herb.

"The Greeks called it hiera botane, or Holy Herb, and in France
to this day Vervain is popularly called Herbe sacrée. From time
immemorial it was the symbol of enchantment, for white and against
black magic. In philtres it engendered love, and was one of the
plants dedicated to Venus, for did not Venus Victrix wear a crown
of Myrtle interwoven with Vervain? The Romans purified their
houses with it to ward off evil spirits, and with it they swept
the altars of their gods.

"Like Dill, it possessed the power of overcoming the spells of
the sorcerer, being ' 'gainst witchcraft much avayling'. Edith
Wheelwright tells us that the Welsh in the Middle Ages called it
"Devil's Bane', and after cutting it in the dark, brought it into
the churches to use as a sprinkler of holy water.

"The Romans were great observers of the custom of New Year's

1972 Moldenke, Notes on Verbena 389

gifts, and those royal colleagues Romulus and Tatius ordained
that every year Vervain should be offered to them with other gifts
as an augury of good fortune for the coming year.

"Vervain has always taken an important place in herbal medicine
since the days of Dioscorides. It was considered efficacious in
cases of scrofula, and Lupton, in his Book of Notable Things
(1660), says 'the root of Vervain hanged at the neck of such as
have the king's evil, it brings a marvellous and unhoped help’.

It was also held to be a cure for the ague and in that curious
book of Dr John Schroder, the Chymical Dispensatory (1669), we
read that 'some cure Tertians and Quartans peculiarly therewith:
in the Tertian they take the third joynt from the Earth and gath-
er it by pulling it upward and give it to be drunk; and in a Quar-
tan the fourth joynt,' This I fancy the good doctor cribbed from
Dioscorides. But, as Gerarde says: "many odde old wives fables
are written of Vervaine.'"

The common names, "Pigeon's grass", "columbine", etc., were
applied to this plant because doves were said to be fond of
hovering around it. "As an herbe of good omen Vervain was one of
those associated with the Eve of St John, and in an old book
called Ye Popish Kingdome appears the following couplet:

"And young men round about with maides coe dance in every streete,
With garlands wrought of Mother-wort, or else with Vervaine
sweete.'

eooeeelt is slightly aromatic when bruised....In tase it is bit-
ter and contains a glucoside, verbenalin, which is employed in
herbal medicine as a febrifuge and in nervous complaints. Some
say that as an anti-thermic it compares favorably with quinine...

'Here holy vervayne and here dill,

'Gainst witchcraft much availing.'

The Muses! Elysium.....

'Trefoil, Vervain, John's Wort, Dill,

Hinder witches of their will.'......

‘Whosoever weareth Vervin or Dill

May be bold to sleep on every hill.'"

Coon (1963) explains that "The constituent which brings Verbena
into the medical field is a bitter glucoside and tannin, a simple
infusion (2 teaspoons to 1 pint) being employed as a diaphoretic,
tonic, and expectorant. There are, in herbal literature, no
strong claims made for its efficacy......An exploration of the
story of vervain leads us down some ancient avenues and provides
an explanation for belief in the efficacy of the plant in herbal
medicine. This plant was first used by the Romans. They gave us
the name 'verbena', which to them meant any one of a number of
plants used in sacrifices, purgation and supplications. Finally
the name was attached to one particular plant, and the virtues as-
cribed to Verbena by the Romans were passed along through the cen-
turies, until, in the Middle Ages, it was said to have been a plant
which, growing on the Mount of Calvary, staunched the wounds of the
Saviour. The transferral of virtues from pagan to Christian (it
has happened in our Christmas celebrations) was not unusual, and ver-
bena early became one of the holy herbs associated with St. John."

[to be continued]

CHEILOTHELA VAGINATA AND SYRRHOPODON STEYERMARKIT

TWO NEW MOSS SPECIES FROM VENEZUELA

Harold Robinson
Smithsonian Institution, Washington, D.C. 20560.

Examination of recent Venezuelan collections has revealed
the following two very distinctive species of mosses.

Cheilothela vaginata H.Robinson, sp. nov. (Figs. 1-3).

Planta dioica? dense caespitosa pallide viridis inferne
sordida. Caules usque ad 4 cm longi simplices vel parce ramosi.
Folia caulina e basi late oblonga vaginata superne abrupte
anguste linearia erecto-patentia ca. 2.5 mm longa 0.8 mm lata,
margine minute crenulata erecta; nervis prope basin 65y latis,
ad apicem subulatis indistincte; cellulis basilaribus angustis
8-12, latis ad 40-95 longis laevibus pellucidis, mediis et
superioribus bistratosis quadratis 7 x 7-l4uy vel brevioribus
oblongis ad extrema amba papillosis.

Venezuela. Estado Mérida: a 2 kms de la Ciudad de Mérida,
sobre rocas, sitio hfmedo. elev. 2,500 m, 22 Feb. 1971, Nora
Faria 1 (US, holotype; D.Griffin, isotype).

The bistratose lamina and the protruding cell ends of the
leaf relate the new species to the common Cheilothela chilensis
(Mont.) Broth. of the Dominican Republic, the Andes south to
Chile and of New Zealand and Kerguelen. The leaf base of the
new species is entirely different in being broadly vaginate and
the plant has a superficial resemblance to Dicranella vaginata
(Hook.) Card.

Syrrhopodon steyermarkii H.Robinson, sp. nov. (Figs. 4-6).

Planta dioica? laxe caespitosa sordide viridis inferne
fulvescens. Caules usque ad 8 cm longi simplices vel parce
ramosi. Folia caulina e basi late ovata vaginata superne sensim
lineari-lanceolata erecto-patentia 6-7 mm longa ca. 1.3 mm lata,
margine superne multistratosa argute duploserrata erecta; nervis
prope basin ca. 100u latis, superne percurrentibus laevibus;
cellulis cancellinarum paucis ca. 15, latis 35-4Oy longis;
cellulis basilaribus plerumque rhomboideis vel oblongis 7-12u,
latis 12-50, longis flavorubescentibus, parietibus nodulosis;
cellulis superioribus quadratis vel breviter oblongis ca. 6y
latis 5-14 longis laevibus; cellulis marginalibus exterioribus
quadratis, interioribus elongatis in basi intramarginaliter
decurrentibus.

390

1972 Robinson, Two new moss species 391

Venezuela. Territoria Federal Amazonas: Cerro Yapacana,
alrededores del campamento a lo largo del rio en las faldas en
la parte suroeste, Lat. 3° 45' N, Long. 66° 45' W. altura: 825
metros, 4 May 1970, J.A.Steyermark 103154 (US, holotype; VEN,
isotype).

The new species of Syrrhopodon is of that group with very
reduced cancellinae and mostly brownish to reddish leaf bases.
The species is most obviously distinctive in the thickened
margins which are continued for a distance intramarginally at
the lower end.

In the same series of collections from the Guayana Highland
region there is a specimen of perhaps the most closely related
species, Syrrhopodon rupestris Mitt. (Territorio Amazonas,

a lo largo de rio Yatua, Cerro Arauicaua, on bluffs at base of
igneous outcrop, 11-12 April 1970, J. A. Steyermark 102544).
The specimen has been compared with a portion of the type
material (Brazil, Spruce 14) and agrees in all respects
including habitat. The new specimen is apparently only the
second known collection of the species and the first from
within the boundaries of Venezuela. Both the foregoing species
appear quite distinct from the more widely distributed

S. rigidus Hook. & Grev. which has much shorter leaves and
shorter stems with less distinct leaf bases.

392 Pipipepeo sO Ger: A Vol. 23, no.

DSO

CBO

RADO™DAOQ2
GF

OS

(?
coer

Figs. 1-6. Venezuelan mosses. 1-3. Cheilothela vaginata.
1. Leaves. 2. Leaf cross-sections. 3. Upper marginal cells of
leaf. 4-6. Syrrhopodon steyermarkii. 4. Leaves. 5. Upper
leaf cells. 6. Leaf margin at top of sheathing base.

STUDIES IN THE EUPATORIEAE (ASTERACEAE). LXXI.

A NEW GENUS, HATSCHBACHIELLA,

R. M. King and H. Robinson
Smithsonian Institution, Washington, D.C. 20560.

Attempts to complete the reorganization of the
Eupatorieae of Southern Brazil have revealed a natural
group of two species with unenlarged hirsute style
bases. For this group we establish a new genus,
Hatschbachiella.

he genus is notable for the small narrow leaves
with nearly entire margins and for the corymbose
inflorescences with subimbricate obtuse phyllaries.
The appearance is similar to the related genus Austro-
eupatorium and one of the species has been frequently
confused with a species of the latter genus. Through
the kindness of Mr. Charles Jeffrey of the Royal
Botanic Gardens, Kew, the type specimen of Eupatorium
tweedianum has been seen and it proves to have the
setiferous achene and short but distinct small-celled
carpopodium which we recognize as characters of
Hatschbachiella. We have previously placed the species
in Austroeupatorium on the basis of misidentified
specimens oF the superficially similar Eupatorium
laetevirens Hook. & Arn. which has only ETE on the
achene and has an elongate carpopodium of very Large
celles,

A second genus of close relationship to Hatsch-
bachiella is Stomatanthes which also has setiferous
achenes and shorter small-celled carpopodia. Stomat-
anthes differs from Hatschbachiella in general aspect
by the usually pyramidally to thyrsoid-paniculate
inflorescences and differs in detail by the unusually
short stout and straight filaments of the anthers.

The new genus is named for Dr. Gerdt Hatschbach
of the Museu Botanico Municipal, Curitiba, Brazil.
Dr. Hatschbach has been an active collector of the
Southern Brazil flora.

Hatschbachiella R.M.King & H.Robinson, genus
novum Asteracearum (Eupatorieae). Plantae herbaceae
erectae pauce ramosae. Caules teretes. Folia
opposita vel alterna subsessilia vel brevipetiolata,
laminis anguste ellipticis integris vel remote
serrulatis. Inflorescentiae corymbosae paniculatae.

ye

394, Peles Ab CO) 1 (OY (e) ae I Vol. 23, nO.

Involucri squamae subimbricatae 12-15 bi-triseriate
hirsutae et glanduliferae obtusae; receptacula plana
glabra. Flores 10-12 in capitulo; corollae anguste
infundibulares 5-lobatae extus glanduliferae intus
glabrae, cellulis angustis parietibus sinuosis, lLobis
aequilateraliter triangularibus sublaevibus; filamenta
antherarum in parte inferiore longa, in parte super-
lore angusta, cellulis quadratis vel elongatis,
parietibus annulate ornatis, cellulis exothecialibus
subquadratis, appendicibus antherarum late ovatis;
Styli inferne non incrassati hirsuti, appendicibus
linearibus vel anguste clavatis distincte papillosis;
achaenia prismatica 5-costata setifera et gland-
ulifera; carpopodia valde distincta brevia, cellulis
subquadratis non inflatis, parietibus subnodulosis;
Pappus setiformis uniseriatus, setis 30-40 scabris
ad apicem tenuibus, cellulis apicalibus acutis.
Species typica: Eupatorium tweedieanum Hook. &
Arn.

Our studies of the genus indicate that it
contains the following two species.

Hatschbachiella polyclada (Dusen ex Malme) R.M.King
& H.Robinson, comb. nov. Eupatorium polycladum
Dusen ex Malme in Svensk. Vet.-Akad. Hand1l. Ser..
vie xi. Nos 2. 39. L93Se Brazile.

Hatschbachiella tweedieana (Hook. & Arn.) R.M.King &
H.Robinson, comb. nov. Eupatorium tweedieanum
Hook. & Arn. in Hook. Comp. Bot. Mag. 1: 242,
1835 (1836). Argentina, Brazil, Paraguay, and
Uruguay.

Acknowledgement
This study was supported in part by the National

Science Foundation Grant GB- 20502 to the senior
author.

STUDIES IN THE EUPATORIEAE (ASTERACEAE). LXXII.

NOTES ON THE GENUS KOANOPHYLLON.

R. M. King and H. Robinson
Smithsonian Institution, Washington, D.C. 20560.

In the months since the circumscription of the
genus Koanophyllon (King & Robinson,1971) three
points have arisen that warrant publication. These
include a neotypification of the type species, a new
species and further notes on one of the common central
american species.

Richard Howard of Harvard University has rightly
called our attention to the problem of neotypification
of the type species of the genus Koanophyllon. From
all indications there is no existing specimen of
K, tinctoria of Arruda de Camara and the nature of
the original work suggests that no herbarium specimen
was ever prepared. In view of this fact we take the
opportunity to select a neotype that will properly
affirm the identity of the species with the Later
described and widely used name Eupatorium laeve DC.
Among the specimens cited by de Candolle (1836) under
the original description of E. Laeve and recorded on
the microfiche series of the de Candolle herbarium
in Geneva is a collection by Lhostky from Rio de Jan-
eiro, Brazil. We propose that this specimen be
recognized as the neotype of Koanophyllon tinctoria
and as the lectotype of Eupatorium laeve DC.

Though the genus Koanophyllon ranges from the
Southwestern United States south to Argentina, the
center of distribution seems to be unquestionably
Guatemala and the adjacent areas. It is from a
closely adjacent region in the Maya Mountains of the
interior of British Honduras that the following new
Species has been collected.

Koanophyllon sorensenii R.M.King & H.Robinson, sp. nov.
Sart rues 0.5 m altus. Caules et petioli

minute dense fulvo-tomentosi. Folia opposita petio-

lata, petiolis ad 1 ecm longis; Lamina usque ad 10.5

em longa 3.5 cm Lata ovato-elliptica, basi acuta,

margine remote crenulata, apice vix acuminata,

paginis utrinque puncto-glanduliferis, inferioribus

minute albo-tomentosis; nervatura pinnata; inflores-

centia laxe paniculata; capitula ca 8 mm alta;

395

396 PHY T0415 O1G 2.2 Vol. 23, no.

involucri squamae bi-triseriatae ca. 15 acutae
exteriores extus minute dense fulvo-tomentose;
flores ca. 10; corolla tubularis, Lobis longe tri-
angularibus extus glanduliferis; appendices anther-
arum breves profunde fisses; styli ad apicem vix
clavati; achaenia pauce setifera et glandulifera;
Pappi setae tenues ca. 30, Grana pollinis ca. 20,
diam. breviter spinosa.

British Honduras: along "Pine Ridge" of
Chiquibul Rd. near junction of Little Vaqueros Creek,
zone of pine and melastomaceous scrub, scattered
oaks present also. 19 August 1971. Low shrub,

0.5 m tall, rays absent, style branches pale greenish
yellow. Paul D. Sorenson 7129 (Holotype US!).

The new species has the general habit of

Koanophyllon solidaginoides, K. celtidifolia and

K. hondurensis but Lacks the ternervate condition

at the base of the leaf. The corolla Lobes and style
branches also differ by being narrower than usual for

the genus

A special effort has been made to further
analyse the rather distinctive Central American
species Koanophyllon albicaulis(Schultz-Bip. ex Klatt)
R.M.King & H.Robinson. A number of interesting
comments have been obtained by reviewing comments on
various labels. The plant ranging from western
Mexico to Honduras is common in moist thickets and
is known by at least two common names in British
Honduras "Water wood" and "Sholte schnook (Mayan)
(Old woman's walking stick)". Of more significance
is a comment on a Honduras collection (Standley
33173)" "Used to? color .green ‘cord, ‘cloth ete.” ie
would seem that K. albicaulis Like K. tinctoris is
a dye plant. There remains a possibility the dye
chemical is different though related since the
comment indicates a green dye while the Brazilian
dye is referred to as indigo. Perhaps chemical
taxonomists will be induced to study the problem in
more detail.

References

Decandolle, A.P. 1836. Ordo CII. Compositae. Prodr.
5: 4-695,

King, R.M. & H.Robinson. 1971. Studies in the Eupator-
ieae (Asteraceae). LXIV. Phytologia 22: 147-152.

STUDIES IN THE EUPATORIEAE (ASTERACEAE). LXXIII.

THE GENUS, OPHRYOSPORUS.

R. M. King and H. Robinson
Smithsonian Institution, Washington, D.C. 20560.

The genus Ophryosporus is a large and natural
group of Eupatorieae occuring mostly in the Central
and Southern Andes of South America. The group is
consistent in the reduced form of the anther appendage
and has been long recognized as a distinct genus.

The clavate form of the style branches and uniform
structure of most flower parts has also helped others
to recognize the genus. It is necessary here to make
only slight revisions.

The genus is extended here to include one
additional species previously placed in a separate
monotypic genus, Trychinolepis B.L.Robinson from
southern Peru. The scale-like pappus caused B.L.
Robinson to establish the separate genus though the
relationship to Ophryosporus was recognized. The
knowledge of unreliability of the pappus and the fact
that it is the only difference leads to the present
reduction of the genus.

The type species of Ophrysporus along with other
Chilean species present a ora distinctive appear-
ance with small shorter petioled leaves and more
spikelike inflorescences. The type species even has
hairs on the inside of the corolla in some specimens,
but there is not enough basis in our estimation for
a distinct genus or subgenus.

Three Mexican species of Decachaeta were placed

at one time in Ophryosporus by B.L.Robinson because
of their short anther appendages. These species can
be distinguished from Ophryosporus easily by the
alternate rather than opposite Leaves, the larger
number of flowers per head, the distinct tuft of
hairs on the receptacle, the anther appendage being
reflexed and undivided instead of inflexed and
divided, and by the cells of the anther collar lack-

ing annular thickenings. No species of Ophryosporus
1s presently recognized north of Colombia.

Ophryosporus Meyen, Reise 1: 1834-35,
397

398 PHY TCEOG FA Vol. 237 Mowe

Trychinolepis B. L. Robinson, Contr. Gray Herb.
Ness 80S 621 L923;

Woody herbs to subshrubs, sparingly branched
often with prominent axillary fasicles. Leaves
opposite, petioles ca. 1 mm.long or up to l cm. long,
blades minute or Large and broadly lanceolate to
elliptical, usually cuneate at base, margins coarsely
to scarcely serrate. Inflorescence corymbose, a
thryse or paniculate with corymbose branches.
Involucre of 4-8 imbricate, subequal phyllaries, in
1-2 series, receptacle slightly convex, glabrous,
3-12 flowers per head; corollas constricted below,
tubular to funnelform above, 5-lobed, lobes with short
slightly to distinctly projecting cells; stomates
absent; anther collar with short cells below and
distinct annular thickenings on the walls. Anther
appendage very small, double inturned remnant,
filaments short or rather long, inserted well above
base; style base without enlarged node, glabrous.
Style branch expanded at tip, smooth and thickened,
narrower and mamillose below. Achenes prismatic
5-ribbed, with small headed glands or setae or both;
base of achene usually twisted with wedge shaped
groups of cells prominent. Carpopodium distinct,
slightly to strongly tapered, with small quadrate
rather thin-walled cells. Pappus of bristles or
broad laciniate scales.

Type species: Ophryosporus triangularis Meyen

Our studies indicate that the genus contains
the following 29 species.

Ophryosporus angustifolius B.L.Robinson, Contr. Gray
Hemboin.saro0ss, 1950. Bolivia.

Ophryosporus apricus B.L.Robinson, Contr. Gray Herb.
He So oe ok, LLoZe Perwe

Ophryosporus ets (Griseb.) Hieron., Engl.
Bot. Jahrb. 1897. Argentina, Bolivia.

Ophryosporus bipinnatifidus B.L.Robinson, Proc. Amer..
AGads Doro. EOL. treru,

Ophryosporus charua (Griseb.) Hieron., Engl. Bot.
Jahrb. 12007055 1897. Argentina’.

age King & Robinson, The genus Ophryosporus 399

Ophryosporus chilca (H.B.K.) Hieron., Engl. Bot.
Jahrb. 22: 706. 1897. Peru.

Ophryosporss clavulatus Griseb., Goett. ABH. 24:

° 79. Argentina.

Ophryosporus cumingii (Sch.-Bip.) Benth., Benth. &
poxcet, Getts 2: 188." 1873. Bolivia;

Ophryosporus eleutherantherus (Rusby) B.L.Robinson,

ontr. Gray Herb. 61: 27. 1920. Bolivia, Peru.

Ophryosporus freyreysia (Dallm.) Baker, Mart. Fl.
Brags 6C€2): Leés. 1876. Brazil.

Ophryosporus hoppii (B.L.Robinson) R.M.King & H.
Robinson, comb. nov. Trychinolepis hoppii B.L.
Robinson, Contr. Gray Herb. n.s. 80: = 1928.

Peru.

Ophryosporus johnstonii B.L.Robinson, Contr. Gray
Herb. /7: 4. £926. Chile.

Cphryosporus kuntzei Hieron., Engl. Bot. Jahrb. 22:
: 974 Bolivia.

eee porns laxiflorus Baker, Mart. Fl. Bras. 6(2):

76. Brazil.

Ophryosporus Lorentzii Hieron., Engl. Bot. Jahrb.
22: 706. 1897. Argentina.

Ophryosporus macbridei B.L.Robinson, Contr. Gray
Herb. neSs 75: 4. LO2K. Peru.

Ophryosporus macrodon Griseb., Goett. ABH. 24: 173.
1879. Argentina, Bolivia.

Ophryosporus organensis Cabr., Arq. Jard. Bot. Rio
Janeiro, : TEAR. Gauloos. BRAZLL.

Ophryosporus origanoides (Meyen & Walp.) Hieron.,
Engl. Bot. Jahrb. 22: 707. 1897. Bolivia,
Ecuador, Peru.

Ophryosporus ovatus B.L.Robinson, Proc. Amer. Acad.
55: 5. LOLS. Peru.

Ophryosporus pachychaeta Baker, Mart. Fl. Bras. 6(2):
Sis L375 Brazil.

1,00 Pi WeEy0 LOxGeIok Vol. 23, tome

Ophryosporus paradoxus (Hook. & Arn.) Benth., Benth.
& Hook, ft. Gen. 2:4239. L873. Chile:

Ophryosporus piquerioides (A.P.Decandolle) Benth.,
Benth. & Hook. £. Gen. 22239, 1873.° Argentine,
Bolivia, Chile, Peru.

Ophryosporus regnellii Baker, Mart. Fl. Bras. 6(2):
188. 1876. Brazil.

Ophryosporus saltensis Hieron., Engl. Bot. Jahrb.
te. wee >

22: 705. 1897. Argentina.

Ophryosporus serratifolius (H.B.K.) B.L.Robinson,
Contr. Gray Herb. n.s. 90: 3. 1930. Golonbres

Ophryosporus sodiroi Hieron., Engl. Bot. Jahrb. 29:
3% L930. Ecuador.

Ophryosporus steinbachii B.L.Robinson, Contr. Gray
Herbs, ness //2 De L926%, Bolivias

Ophryosporus triangularis Meyen, Reise 1: 402. 1834.
Chile.

Ophryosporus venosissimus (Rusby) B.L.Robinson, Proc.
Amer. Acad. GL: 271. 1905. Bolivia.
Acknowledgement
This study was supported in part by the National

Science Foundation Grant GB- 20502 to the senior
author.

‘ky

13 “2 eR
: a ANICAT

GALS
Z PHYTOLOGIA

Designed to expedite botanical publication

Vol. 23 July, 1972 No. 5

CONTENTS

RUDD, V. E., A new species of Myrocarpus (Leguminosae) and a brief
ION PRE MAIN, oc ops Joe by'! Ds iia Wd tata ecw eateries Fete 401
KING, R. M., & Robinson, H., Studies in the Eupatorieae (Asteraceae).
LXXIV. New species of Critonia, Fleischmannia and

NTIS ge 59S Ke Bae nach owt cay Pe had a cas whine ele 405
KING, R.M., & ROBINSON, H., Studies in the Eupatorieae (Asteraceae).
LXXV. A new genus, Cronquistianthus .............45. 409

MOLDENKE, H.N., A Fifth Summary of the Verbenaceae, Avicenniaceae,
Stilbaceae, Dicrastylidaceae, Symphoremaceae, Nyctanthaceae,
and Eriocaulaceae of the World as to Valid Taxa, Geographic

Distribution, and Synonymy. Supplement 1. ............. 413
MILDNER, R., & ROGERS, C. M., New names and a new combination in

South American Linum (Linaceae) ............0024455 439
SEYMOUR, F. C., Second Expedition to Nicaragua, 1969-1970 ........ 440
GRIFFIN, D., Ill, Pohlia annotina (Hedw.) Lindb. var. loeskei Crum,

Steere and Anderson in Guatemala. .............2 25005. 452
MOLDENKE, H. N., Seven undescribed subspecific taxa in the

Eriocaulaceae and Verbenacede..........022 cece eeeee 453
ne WW, Cr. V AFICRUNE-ONCIGIUINS 30% oon eo Ro os Sw ols ee re 455
MOLDENKE, H. N., Additional notes on the genus Verbena. XIV....... 460
WURDACK, J. J., Certamen Melastomataceis XIX ...........-0045. 474
ESA” SE. NIOUAE SOWIOWNE. CFs! hoa sol Ps + dish wn mw Seneca miele ow 502
Index to authors in Volume Twenty-three... 2.2... 0 ee eee eens 503
Index to supra-specific scientific names in Volume Twenty-three........ 503
Publication dates for Volume Twenty-three. .... 2.0.00 022 0222 eees 512

Published by Harold N. Moldenke and Alma L. Moldenke

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

Price of this number, $2; per volume $8 in advance,
or $8.50 at close of volume; foreign orders 50 cents extra

A NEW SPECIES OF MYROCARPUS (LEGUMINOSAE)
AND A BRIEF RESUME OF THE GENUS

Velva E. Rudd, Smithsonian Institution

The small genus Myrocarpus Freire Allemao has hitherto been
known only from southern Brazil, Paraguay, and northeastern Ar-
gentina. The new species from Venezuela, described below, is an
interesting disjunct.

Like the related genera, Myroxylon L. f. and Myrospermum
Jacq., it is balsamiferous, and when sterile can be recognized
by leaflets with pellucid dots and dashes. Myrocarpus has smaller
flowers, the pods are elongate, compressed, usually 3-5-seeded,
with marginal wings, in contrast to those of Myroxylon and
Myrospermum which are commonly 1l-seeded, with a basal wing.

Some of the species of Myrocarpus are poorly known, and I am
reserving judgment as to how "good" they are. The following key
and descriptions are based on what information I have at hand.

Myrocarpus, with Myroxylon and Myrospermum, has been placed
in the faboid tribe Sophoreae by most authors. Hutchinson (Gen.
Fl. Pl. 1: 314. 1964) included Myrocarpus in his expanded inter-
pretation of the tribe Cadiae, which he transferred from the
Caesalpiniaceae to the Fabaceae, but retained Myroxylon and Myro-
spermum in the Sophoreae. Much more study is needed of the genera
transitional between the caesalpinioid and faboid Leguminosae be-
fore a satisfactory understanding and classification can be
reached. For the time being, I prefer to retain Myrocarpus in the
Sophoreae sensu latior.

MYROCARPUS Freire Allem&o, Diss. Leg. 26 Oct. 1847.

Trees, unarmed, balsamiferous; leaves alternate, imparipin-
nate, (1-) 3-9 (-10)-foliolate; leaflets alternate, ovate to lan-
ceolate-ovate, glabrous, with pellucid dots and dashes; stipules
minute, caducous; stipels lacking; inflorescences axillary or
pseudoterminal, many-flowered spicate racemes; bracts minute,
deltoid; bracteoles lacking; flowers small, fragrant, about 6-8
mm. long; calyx turbinate-campanulate, subsericeousm 3-4 om. lom
with 5 short, subequal lobes, the vexillar lobes sometimes con-
nate; corolla with (3-4) 5 free, whitish, linear, clawed, sub-
equal, glabrous petals about 1 m. wide; stamens 6, 8, or 10, the
filaments free, subequal, mostly exceeding the petals, the an-
thers uniform, subcordiform, basifixed, dehiscing by lateral
slits; ovary brevi-stipitate, usually 3-5-ovulate, the style
short, glabrous, the stigma minute, terminal; fruit elongate,
laterally compressed, samaroid with marginal wings, commonly 3->>-
seeded; seeds compressed fusiform or oblong, the hilum apical.

Type species: M. fastigiata Freire Allemao.
ol

02 Pon vet Os sONG ek Vol. 23, no. 5
Key to species

Leaflets 1.5-2.7 cm. long, 1-1.5 cm. wide, acute to obtuse;
fruit 7-8 mm. wide, the wings about 2 m. wide, the semini-
ferous body 3.5-4 mm. wide; ovary lightly pubescent, glab-
rescent; flowers Dec.-Jan. Brazil: Rio de Janeiro.

1. M. fastigiata.

Leaflets (2-) 2.5-9 cm. long, 1.5-6 cm. wide, acute to acuminate,
occasionally obtuse or emarginate; fruit 10-20 mm. wide, the
wings 3-6 mm. wide, the seminiferous body 3-9 mm. wide;
ovary glabrous.

Fruit with body 8-9 mm. wide; leaves 3-4-foliolate, the axis
1.5-4 em. long; flowers Sept. Brazil: eastern Sao Paulo.
3. M. leprosus.
Fruit with body 3-6 mm. wide; leaves (1-) 4-9-foliolate, the
axis (3-) 5-12 cm. long.
Length of fruit 4.5-9 cm. long, the width 1-2 cm. the body
about 3 m. wide, the wings 3-6 mm. wide; flowers May-
Dec. Brazil: southern Minas Gerais and Rio de Janeiro
south to Santa Catarina; Rraguay; nrtheastern Argentina.
2. M. frondosus.
Length of fruit 9-19 cm. long, the width 2 cm., the body 5-6
mm. wide, the wings 5-8 mm. wide flowers Jan. ? (fruit in
April). Venezuela: western Apure. hy M. venezuelensis,

1. MYROCARPUS FASTIGIATUS Freire Allem&o, Diss. Leg. 26 Oct.184¥7.
Type: F. Freire Allemao s.n., Brazil, Rio de Janeiro, "sil-
vis primaevis" (holotype R 7; isotypes BM, G, F neg. 28193
ex G)i

large tree; leaves 5-9 (-10)-foliolate, the axis about 3.5-
6.5 cm. long; leaflets ovate to elliptic, 1.5-2.7 cm. long, 1-
1.5 cm. wide, acute to obtuse, occasionally emarginate, the base
rounded to acute; flowers (in January and February) about 6 m.
long, the calyx 2-3 mm. long; stamens 6, 8, or 10 (fide Freire
Allemao); ovary pubescent, glabrescent; fruit about 2.5-6 cm.
long, 7-8 mm. wide, the seminiferous body 3.5-4 mm. wide, the
wings 2 mum. wide.

Distribution: Known only from the area of the type collec-
tion.

Local names: cabureiba; oleo pardo.

1972 Rudd, New species of Myrocarpus 03

2. MYROCARPUS FRONDOSUS Freire Allemao, Diss. Leg. 22 Sept. 1848,
Type: F. Freire Allem&o s.n., Brazil, Rio de Janeiro, "sil-
vis ieommbaet (holotype R 7; isotypes BM, G, F neg. 28194
ex G).

Leptolobium ? punctatum Bentham, Linnaea 22: 526. Sept. 1849.
Type: A. F. Regnell II.653, Brazil, Minas Gerais, Caldas,
26 Aug. 1868 (in flower) Theloteve K; isotypes NY, US).

Myrocarpus r riensis Hallier f. Meded. Rijks. Herb. Leiden
NO. 27: io. 1915. Type: K. Fiebrig 165 (as 105), Paraguay,
oo de Altos, 29 Sept. 1902 (holotype L; isotypes P,
US).

Myrocarpus frondosus forma paraguariensis (Hallier f.) Hassler,
Repert. Nov. Spec. Fedde 16: 233. 1919.

Tree, to about 25 m. tall; leaves (1-3) 5-10-foliolate, the
axis (3-)5-12 cm. long; leaflets ovate to lanceolate-ovate, (2-)
2.5-7 cm. long, 1.5-3.5 cm. wide, acute to acuminate, sometimes
obtuse or emarginate, the base rounded to acute; flowers (in May-
December) about 6-8 mm. long, the calyx 3-4 mm. long; stamens 10;
ovary glabrous; fruit 4.5-9 cm. long, 1-2 cm. wide, the body
about 3 mm. wide, the wings 3-6 mm. wide.

Distribution: In forest, southern Brazil, Paraguay, and
northeastern Argentina, at elevations of 100-1200 m.

Local names: cabreuna; cabreuva; ibird-payo; ivird-paye 2
incienso; oleo pardo.

Specimens examined:
BRAZIL: Minas Gerais: Caldas, Regnell II-653, 16 Sept. 1865 US).
Rio de Janeiro: Novo Friburgo, Glaziou 11908 (NY, P, US). Petro-
polis, Glaziou 8446 (Ny; US), 11906 (P, US), 11907 (P).
Guanabara:—Corcovado, Glaziou 6516 (P, US). Tijuca, Glaziou 6837
Tingua, Glaziou 8649 (P). Rio de Janeiro, Horto Florestal, Herb.
no. 111914 (NY). Sdo Paulo: Areias, Kuhlmann 4422 (ny, US). Cab-
reuva, Hoehne 31001 (NY). Souzas, Moreira de Souza s. n. (NY).
Parana: Paranagua, Hatschbach 2480 (US). Campina Grande do Sul,
Hatschbach 5212 (US), 17812 (US). Morrétes, Hatschbach 22125 (NY).
Santa Catarina: Bom Retiro, Lomba Alta, Smith, Reitz, & Klein
1957 (NY, US). Herval Velho, above Rio Leao, Smith & Reitz 12417
(US). Xanxeré, Smith & Reitz 12496 (NY, US). Itapirange, Rio Uru-
guai, Smith & Reitz 12660 (NY, US). Sado Miguel d'Oeste, Smith &
Reitz 12809 (US). Uruguaf, Vila Rica, by Rio do Peixe, Smith &
Reitz 120924 (US). Rafael, Ibirama, Klein 678 (US). Ibirama, Reitz
& Klein 3835 (NY, US). Serra do Matador, Rio do Sul, Reitz &
Klein 7246 (NY, US), 8291 (US). Alto Matador, Rio do Sul, Reitz &
Klein 7588 (NY, us). Rio do Meio, Lauo Mueller, Reitz & Klein
8228 (NY, US). Sanga da Areia, Jacinto Machado, Reitz & Klein
9290 (wx}, 9369 (US).

Oh PHYTO LO G Tk Vol. 23, toae

PARAGUAY: Montes del Parque, Rojas 8649a (US). Villarrica,
Jorgensen 3605 in part (US). Limpio, Hassler 3226 (NY, P). Rio
Paraguay near Concepeidn Hassler 7335 (wy, Ee US). Yparacay,
Hassler 12228 (L, NY, us), 12682 (L, NY, US). Yaguaron, Balansa
3112 @, P, US).

3. MYROCARPUS LEPROSUS Pickel, Arq. Bot. Estad. S. Paulo, n. sen
3: 161, tab. 41. 1955. Type: B. J. Pickel s.n., Brazil, Sado
Paulo, Santos, "in fragosis montis S. Theresiae," 29 Sept.
1949 (flowers), 27 Nov. 1949 (immature fruit) (hdotype SPSF
no. 3462; isotype SP no 53474).

Small trees; leaves 3-4-foliolate, the axis 1.5-3.5 (-4) cm
long; leaflets ovate to obovate, 5-9 cm. long, 3-4.5 cme wide,
acuminate, obtuse, or retuse, the base acute or rounded; flowers
(in September) about 8 mm. long, the calyx 4 mm. long; stamens
8-10; ovary glabrous; fruit (immature ) 5 cm. long, 1. cm. wide,
the seminiferous body 8-9 mm. wide, the wings 3-4 m. wide.

Distribution: Known only from the type collections.

I have not seen material of M. leprosus. The above descrip-
tion is based on that of Pickel. The species seems to be very
close to M. frondosus, differing chiefly in the size of the fruit

4, MYROCARPUS VENEZUELENSIS Rudd, sp. nov.
Type: J. A. Steyermark, G. Bunting, & C. Blanco 101748,
Venezuela, Apure, Reserva Forestal San Camilo, "selva siem-
preverde a lo largo de la Quebrada de la Cristalina hasta la
vega del Cerro La Nulita," 250-280 m. alt., 2 April 1968
(holotype US no. 2622434; isotypes NY, VEN).

Arbor, M. frondosus affinis, sed imprimis fructibus majus-
culis differt.

Tree about 25 m. tall; leaves 5-7-foliolate, the axis 6-12
cm. long; leaflets ovate, 4-8 cm. long, (2-) 2.5-4 cm. wide, acu-
minate, the base rounded to subcordate; complete flowers not seen,
calyx 3-4 mm. long; fruit 9-19 cm. long, 2 cm. wide, the semini-
ferous body 5-6 mm. wide, the wings 5-6 mm. wide.

Distribution: Known only from the type collection.

Local name: barbasco.

STUDIES IN THE EUPATORIEAE (ASTERACEAE). LXXIV.

NEW SPECIES OF CRITONIA, FLEISCHMANNIA AND HEBECLINIUM.

R. M. King and H. Robinson
Smithsonian Institution, Washington, D.C. 20560.

The revised classification of the Eupatorieae
has greatly simplified the identification of specimens
and has facilitated the recognition of previously
undescribed species. Four new species, belonging
to the following three genera, are described here.

Critonia dominicensis R.M.King & H.Robinson,
Sp. nov. Frutices ca. 2 m alti laxe ramosi. Caules
teretes vel obscure angulati glabri. Folia opposita
longe petiolata, petiolis 1.5-2.0 mm lLongis; Lamina
anguste Lanceolata 12-20 cm longa 3-5 cm Lata glabra
distincte punctata longe attenuata remote serrulata
base anguste cuneata, nervis pinnatis secundariis
utrinque 6-7. Inflorescentiae dense corymbosae;
pedicelli glabri. Capitula 3-6 fasciculata ca. 6 mm
alta; flores 5; involucri squamae ca. 20, imbricatae
4-5-seriatae valde inaequilongae interiores anguste
oblongae facile deciduae glabrae; corollae anguste
tubulares ca. 3.5 mm lLongaeglabrae, lobis Longe tri-
angularibus; appendices antherarum vix Longiores
quam latae; appendices stylarum Lineares; achaenia
temeriter setifera, late costata, basi angustata;
Pappi setae ca. 35 ad apicem leniter dilatatae.
Grana pollinis ca. 20, diam.

Type: DOMINICA: British West Indies: Fond Baron
Estate. Abundant shrubs ca. 2 m. tall, shade, flowers
all past anthesis. 18 December 1967. Robert Merrill
King 6301. (Holotype US !).

The species is in the group with Critonia dalea
but the long attenuate leaf apex is very distinctive.
The following species should also be added to the
genus Critonia though the anther appendage is some-
what shorter.

Critonia platychaeta (Urban) R.M.King & H.Robinson,
comb. nov. Eupatorium platychaetum Urban, ‘
Notizbl. Bot. Gart. Berlin 8: 23. 1921. Jamaica.

hos

06 PHY TOLOG IA Vol. 23, no. 5

Fleischmannia cuatrecasasii R.M.King & H.Robinson,
sp. nov. Plantae herbaceae vel suffrutescentes
erectae usque ad 3 dm altae, pauce vel multo ramosae
in sicco olivaceae vel atratiores. Caules teretes
puberuli. Folia opposita vix petiolata, petiolis 1-2
mm longis; Lamina linearis elliptica usque ad 2.7 cm
longa, 0.3 cm lata, remote pauce serrulata ad apicem
anguste obtusa base anguste cuneata trinervata, supra
subglabra, subtus in nervis et marginem minute
puberula. Inflorescentiae lLaxe cymosae; pedicelli
puberuli 2-5 mm Longi. Capitula 3.5 mm alta; flores
ca. 20; involucri squamae 20-25 subimbricatae tri-
seriatae inaequilongae interiores ca. 2.5 mm lLongae,
anguste oblongae, breviter acutae vel oblongae, base
distincte callosae; corollae anguste infundibulares
1.5-1.7 mm lLongae, extus inferne glabrae, Lobis
equilateraliter triangularibus extus saepe pauce
breviter setiferis; thecae antherarum 0.5-0.7 mm
longae; appendices parvae vix longiores quam Latae
superne crenulatae; achaenia glabra vel perpauce
setifera; carpopodia valde tumescentia Latiora quam
longa; pappi setae ca. 25. Grana pollinis 18-20,
diam.

Type: COLOMBIA: Valle del Cauca: Quebrada La Brea,

near Cordoba. Flowers pink. November 7, 1944, Earl

L. Core 1549 (Holotype US!). Paratypes El Valle:

Rio Calima: Quebrada de La Brea. Alt. 30-40 m.May 1946
Richard Evans Schultes & Mardoqueo Villarreal 7359
(UST). Valle: Rio Calima (region del Choco): entre
Pailon y El Coco. 50 met. alt. 23 Mayo 1946. J. Cuatre-
Casas 21245 (Facultad de Agronomia del Valle f).

The species is related to F. misera of the Choco
but is distinct by the more erect habit, narrower
More remotely serrulate margins, darker color and
firmer texture of the leaves. According to Dr. Cuatre-
Casas the species occurs in a distinctive ecological
niche in the region largely Lacking composites in the
Pacific Lowlands of Colombia, a zone inland from the
outer coastal zone where the Eupatorian genus Tubero-
styles is found.

Hebeclinium reedii R.M.King & H.Robinson, sp.
SSS
nov. Suffrutices erecti. Caules teretes dense Longe
hirsuti. Folia opposita longe petiolata, petiolis
3-4 cm longis; Lamina Late ovata usque ad 12 ecm Longa,
8 cm lata, distincte breviter acuminata duplo-serrata
basi Late cuneata supra in nervis breviter hirsutis,

1972 King & Robinson, New species 07

subtus in nervis et nervulis hirsutis, nervis pinna-
tis secundariis utrinque ca. 4-5, ca. 45° ascendent-
ibus. Inflorescentiae lLaxe cymosae; pedicelli
breviter hirsuti. Capitula plerumque 3-fasciculata,
ca. 6 mm alta; flores 20-25 albi; involucri squamae
ca. 40 imbricatae quadriseriatae valde inaequilongae
anguste oblongae obtusae superne puberulae; receptacula
convexa ubique sclerotica pilosa; corolla anguste
infundibulares ca. 3 mm Longa intus glabra, lobis
breviter triangularibus extus dense setiferis, setis
brevibus; appendices antherarum vix lLongiores quam
latae; achaenia glabra; pappi setae ca. 25 ad apicem
distincte breviter dilatatae. Grana pollinis 18-20u
diam.

Type: PANAMA: Darien: Cerro Pirre. Matita, flores
blancas, sin olor. 11 de Abril 1967. Narciso Bristan
464, (Holotype US!).

The species is apparently closest to Hebeclinium
peeeeeehtcum of adjacent Colombia but leaves of the
atter are more nearly entire or slightly crenulate
with undersurface more densely pubescent. The petioles
of the latter species are generally shorter and
thicker and the phyllaries are narrower at the tips
and more sparcely pubescent.

We take great pleasure in naming this new species
for Dr. Clyde F. Reed of Baltimore, Maryland who very
generously donated the type specimen to the United
States National Herbarium.

Hebeclinium costaricense R.M.King & H.Robinson,
sp. nov. Hebeclinio reedii simile sed caulibus minus
pubescentibus, fFoliis minus acuminatis, laminis 9-16
em longis 6-9 cm lLatis margine leniter crenulatis,
subtus in nervulis hirtellis, receptaculis magis
convexis, floribus ca. 40 in capitulo, achaeniis
pauce glanduliferis.

Type: Costa Rica:Alajuela. Near La Laguna, 6 to
8 km. south of Villa Quesada. Alt. 1200 m. Feb. 19,
1966. Flowers white, herb 1.5 meters tall. Cut over
forest area, mountains. Antonio Molina R., Louis O.
Seem ans 5 William C. Burger & Bruce Wallenta 1/545.
E Paratype Costa Rica. Cataratas de San Ramon..
February 28, 1931. A.M. Brenes 13564 (F !).

The new species is clearly distinct from either
of the two species of Hebeclinium previously known

08 PH YD O"L OGD & Vol. 23, no. 5

from Costa Rica. From H. macrophyllum, the new species
differs most obviously by the broadly cuneate rather
than cordate leaf-base and by the broader tipped less
glaucous phyllaries. Hebeclinium hygrohyaeum (B.L.
Robinson) R.M.King & H.Robinson has Longer elliptical
leaves with nearly entire margins. The new species

is actually most closely related to H. reedii described
above from eastern Panama. The panamanian species is
most easily distinguished by the closely doubly

serrate leaf margin but differs more significantly in
the less convex receptacle bearing fewer flowers and

by the glabrous achene.

Acknowledgement
This study was supported in part by the National

Science Foundation Grant GB-20502 A #1 to the senior
author.

STUDIES IN THE EUPATORIEAE (ASTERACEAE). LXXV.

A NEW GENUS, CRONQUISTIANTHUS .

; _R. M. King and H. Robinson
Smithsonian Institution, Washington, D.C. 20560.

One of the most important elements of the
Eupatorieae in the Andean Region is a complex of
rather woody species with nearly smooth corolla lobes,
plain styles, and well-developed anther appendages.
Many of the species have distinctive pale pubescence
on the underside of their leaves. One genus related
to the complex, Lourteigia, has been described
previously (King & H.Robinson, 1971) having achenes
greatly constricted under the pappus, having 20 or
ca. 40 flowers per head and having dense clusters of
hairs on the outer surface of the corolla Lobes. A
second genus of the complex, Cronquistianthus, is
described here on the basis of 13 species that lack
the constriction under the pappus, have 8-18 flowers
per head and have only glands or scattered hairs on
the outer surface of the corolla lobes. A marked
characteristic of the new genus is the usually
stopper-shaped asymmetrical carpopodium set off by
numerous thin-walled easily collapsing wedge-shaped
cells. None of the other related genera have such
distinctly delimited carpopodia.

There are some other features of the new genus
that are notable. The glands on the corolla are
more like those of Ophryosporus and Helogyne in Lack-
ing much enlarged apical cells. The enlarged tips
of the style branches of some species are also
reminiscent of Ophryosporus. Ophryosporus is totally
distinct by the SaaEE anther Pree Ce: and Helogyne
differs by its densely glanduliferous achenes ea
very broad style branches. The involucres of some
species of Cronquistianthus, especially C. chamaedri-
folius, have strongly imbricated phyllaries and
resemble those of Chromolaena. The latter genus has
a totally different type of carpopodium with straight
sides and an outer sheath of cells while the typical
element has papillose corolla lobes and a distinctive
form of anther collar. One species of Cronquist-
ianthus, C. organoides, has scattered hairs inside of
the corolla, a feature not found in any other species
of the genus, even the closely related C. niveus. A
feature that does seem consistent throughout

409

10 PEYTOLOGIA Vol. 23, no. 5

Cronquistianthus is the form of the pappus setae
GEE taper to a narrow rather smooth tip.

In naming the new genus we honor again Dr.
Arthur Cronquist of the New York Botanical Garden

whose philosophical guidance and objectivity have
been particularly welcome to the authors.

Cronquistianthus R.M.King & H.Robinson, genus
novum Asteracearum (Eupatorieae). Plantae frutes-
centes erectae pauci- vel multi-ramosae. Caules
teretes, glabrescentes; ramulis puberulis. Folia
opposita breviter vel longe petiolata, laminis
linearibus, Lanceolatis vel ovatis acutis utrinque
pubescentibus subtus interdum dense albotomentosis,
margine integris vel serratis. Inflorescentiae dense
corymbosae, capitulis subsessilibus saepe in glomer-
ulis congestis. Involucri squamae imbricatae valde
inaequilongae tri-quinqueseriatae, 12-25, obtusae;
receptacula plana glabra. Flores 8-18 in capitulo;
corollae tubulares vel anguste infundib-
ulares 5-lobatae, lobis aequilateraliter triangular-
ibus vel longioribus lLaevibus extus glanduliferis
vel breviter setiferis, glandibus vix capitatis;
filamenta antherarum alte inserta, in parte superiore
angusta, cellulis quadratis vel elongatis, parietibus
transverse annulatis, cellulis exothecialibus sub-
quadratis, appendicibus longioribus quam latis
interdum retusis; styli inferne non nodulosi glabri,
appendicibus breviter papillosis ad apicem Linear-
ibus vel abrupte dilatatis; achaenia prismatica
plerumque 5-costata pauce vel multo setifera; carpo-
podia distincta subobturaculiformia valde inaequalia,
cellulis parvis subquadratis vel rotundatis multi-
seriatis, parietibus aliquantum incrassatis; pappus
setiformis uniseriatus, setis ca. 30-35 contiguis ad
apicem sensim angustioribus, cellulis apicalibus
acutis,.

Species typica: Eupatorium niveum H. B. K,

Our studies of the genus indicate that it
contains the following 13 species.

Cronquistianthus chamaedrifolius (H.B.K.) R.M.King &
-Robinson, comb. nov. Eupatorium chamaedri-
folium H.B.K., Nov. Gen. et Sp. 4: 88-89, ed.
fol. 1818. Ecuador, Peru.

Cronquistianthus chotensis (Hieron. ) R.M.King & H.
Robinson, comb. nov. Eupatorium chotense Hieron.
Engl. Bot. Jabxrb .,36)- Lee. L905... -Pexar,

1972 King & Robinson, Gems Cronquistianthus Te
Cronquistianthus desmophyllus (B.L.Robinson) R.M.King

& H.Robinson, comb.nov. Eupatorium desmophyllum
B.L.Robinson, Contr. Gray fess Nee Yar os Leed.

Peru.

Cronquistianthus glomeratus (A.P.Decandolle) R.M.King
& H.Robinson, comb. nov. Eupatorium glomeratum

A.P.Decandolle, Prodr. 5: 154, 1836. Peru.

Cronquistianthus kalenbornianus (B.L.Robinson) R.M.
King & H.Robinson, comb. nov. Eupatorium
kalenbornianum B.L.Robinson, Contr. Gray Herb.
nes. OL: 8. 1920. Peru.

Cronquistianthus lLavandulaefolius (A.P.Decandolle)
R.M.RKing & H.Robinson, comb. nov. Eupatorium
Llavandulaefolium A.P.Decandolle, Prodr. 5: L154.

836. Peru.

Cronquistianthus leucophyllus (H.B.K.) R.M.King & H.
obinson, comb. nov. Eupatorium leucophyllum
H.B.K., Nov. Gen. et Sp. G: 90. ed. fol. 8.
rer.

Cronquistianthus niveus (H.B.K.) R.M.King & H. Robins.
comb. nov. Eupatorium niveum H.B.K., Nov. Gen.
et Sp. 4: 90. ed. fol. L818. Colombia, Ecuador.

Cronquistianthus origanoides (H.B.K.) R.M.King & H.
Robinson, comb. nov. Eupatorium origanoides
H.B.K., Nov. Gen. et Sp. 4: 89. ed. fol. 18.
Colombia, Ecuador.

Cronquistianthus pseudoriganoides (Hieron.) R.M.King
& H.Robinson, comb. nov. Eupatorium ps

eud-

origanoides Hieron., Engl. Bot. Jahrb. 29: 10.
Ll - Ecuador.

Cronguistianthus rugosus (H.B.K.) R.M.King & H.Robin-
son, comb. nov. Eupatorium rugosum H,B.K.., Node.
Gen. et Sp. 4: 89. ed. fol. -Colombia,
Ecuador.

Cronquistianthus urubambensis (B.L.Robinson) R.M.King
& H.Robinson, comb. nov. Eupatorium urubambense
B.L.Robinson, Contr. Gray Herb. n.s. 60: 58.

1919. Peru.

Cronquistianthus volkensii (Hieron.) R.M.King & H.
Robinson, comb, nov. Eupatorium volkensii Hieron.,

412 P. He ¥t) Osly OG wk Vol. 23, no. 5
Engl. Bot. Jahrb. 40: 370. 1908. Peru.

Reference

King, R.M. & H.Robinson. 1971. Studies in the
Eupatorieae (Compositae). XXXV. A new genus,
Lourteigia. Phytologia 21:28-30.

Acknowledgement

This study was supported in part by the National
Science Foundation Grant GB-20502 A #1 to the senior
author.

A FIFTH SUMMARY OF THE VERBENACEAR, AVICENNIACEAE, STILBACEAE,
DICRASTYLIDACEAE, SYMPHOREMACEAE, NYCTANTHACEAE, AND
ERIOCAULACEAE OF THE WORLD AS TO VALID TAXA, GEOGRAPHIC

DISTRIBUTION, AND SYNONYMY

Supplement 1

Harold N. Moldenke

Addenda & errata to Part I:

The name, "Phyla nodiflora var. reptans (H.B.K.) Moldenke", is
to be corrected to read "Phyla nodiflora var. reptans (Spreng.)
Moldenke" in the 58 places where the name occurs.

UNITED STATES OF AMERICA:

Maine:

Eriocaulon pellucidum Michx. [Isle au Haut]
Verbena hastata L. [Isle au Haut]

Massachusetts:

Eriocaulon parkeri B. L. Robinson [Barnstable County]

Connecticut:

Verbena simplex Lehm. [Middlesex County]

New Jersey:

Eriocaulon pellucidum Michx. [Cape May County]
Verbena simplex Lehm. [Cape May County]
Verbena stricta Vent. [Cape May County]
North Carolina:
Clerodendrum trichotomum var. fe ineum Nakai [Surry Ceunty]
Verbena hastata L. [Ashe County]
Verbena urticifolia L. [Ashe County]
South Carolina:
Lantana horrida H.B.K. [Orangeburg County]
Phyla lanceolata (Michx.) Greene [Georgetown County]
Verbena brasiliensis Vell. [McCormick County]
Verbena urticifolia L. [Georgetown County]
Florida:
Clerodendrum bungei Steud. [Dade County]
Duranta repens var. alba (Masters) L. H. Bailey [Monroe Coun-
ty]
Lippia alba (Mill.) N. E. Br. [Monroe County]
Vitex agnus-castus L. [Dade County]
Vitex glabrata R. Br. [Dade County]
Mississippi:
Clerodendrum bungei Steud. [Claiborne County]

Ohio:

Verbena bracteata Lag. & Rodr. [Gallia County]

Indiana:

413

yi) PHY T 0 L0G ik Vol. 23, no. 5

Verbena bipinnatifida Nutt. [Lake County]
Verbena canadensis (L.) Britton [Franklin County]
Verbena simplex Lehm. [Franklin County]
Verbena stricta Vent. [Franklin County]
Minnesota:
Verbena bracteata Lag. & Rodr. [Houston County]
Verbena hastata L. [Houston County]
Verbena hastata f. rosea Cheney
South Dakota:
Verbena hastata L. [Jones Island]
Missouri:
Verbena brasiliensis Vell. [Saint Louis]
Louisiana:
Phyla incisa Small [Saint Bernard Parish]
Oklahoma:
Verbena bipinnatifida Nutt. [Roger Mills County]
Texas:
Verbena plicata Greene [Armstrong County]
Vitex negundo var. intermedia (P'ei) Moldenke [Donley County]
Oregon:
Terbena officinalis var. albiflora Strobl [Multnomah County]
MEXICO:
Bouchea nelsonii Grenz. [Veracruz]
Bouchea spathulata var. longiflora Moldenke [Coahuila]*
Callicarpa acuminata var. argutedentata Moldenke [Chiapas &
Tamaulipas ]
Callicarpa acuminata var. pringlei (Briq.) Moldenke [Hidal-
go, Quintana Roo, San Luis Potos{, Tamaulipas, Veracruz,
& Yucatdn]
Callicarpa pringlei Briq. — to be deleted
Citharexylum affine D. Don [Veracruz]
Citharexylum oleinum (Benth.) Moldenke [Oaxaca]
Cornutia pyramidata var. isthmica Moldenke [Yucatdn]
Eriocaulon molinae L. 0. Williams [Jalisco]
Lantana velutina f. macrophylla Moldenke [Guerrero & Oaxaca]*
Lantana velutina f. violacea Moldenke [Oaxaca]
Lippia oleia var. ovatifolia Moldenke -- delete the as-
terisk
Lippia mcvaughii Moldenke [Jalisco]
Lippia pringlei f. intecta Moldenke [Jalisco]
Phyla incisa Small [Hidalgo]
Verbena elegans var. asperata Perry [Veracruz]
Vitex agnus-castus L. [Nuevo Leén]
YUCATAN ISLANDS:
Callicarpa acuminata var. pringlei (Briq.) Moldenke [Cozumel]
Callicarpa p pringlei Briq. -=- to be deleted
GUATEMALA =
Bouchea prismatica var. brevirostra Grenz. [El Petén]

1972 Moldenke, Fifth Summary Supplement 1s

Callicarpa acuminata H.B.K. [Guatemala]

Callicarpa acuminata var. argutedentata Moldenke [El Petén]

Citharexylum cooperi Standl. [Izabal }

Citharexylum crassifolium Greenm. [(Chimaltenango & Santa
Rosa]

Citharexylum donnell-smithii Greenm. (San Marcos & Zacapa]

Citharexylum hirtellum Standl. [Izabal]

Citharexylum mocinni D. Don [El Quiché, Guatemala, Quezal-
tenango, & Sacatepéquez]

Citharexylum pterocladum Donn. Sm. [Chimaltenango & El Petén]

Citharexylum steyermarkii Moldenke [Zacapa]

Cormtia grandifolia (Schlecht. & Cham.) Schau. [Guatemala &
Sacatepéquez]

Cormutia pyramidata var. isthmica f. albida Moldenke [El
Petén]+#

Duranta guatemalensis Moldenke [Huehuetenango]

Lantana hirta Grah. [El Quiché]

Lantana hispida H.B.K. [Chiquimula]

Lantana microcephala A. Rich, [El Petén]

Lantana trifolia L. [El Quiché & Guatemala]

Lippia alba (Will.) N. E. Br. [Alta Verapaz & Chiquimula]

Lippia cardiostegia Benth. [Alta Verapaz, Chimaltenango, El
Quiché, & Sacatepéquez ]

Lippia chiapasensis Loes. [Baja Verapaz, Solol4, & Totonic-
apan]

Lippia curtisiana Moldenke -- delete Baja Verapaz, Sololé,
& Totonicapdn

Lippia hypoleia var. ovatifolia Moldenke [El Progreso]

Lippia substrigosa Turcz. [Alta Verapaz & Chiquimula]

Petrea volubilis L. ([Chimaltenango, El Progreso, El Quiché,
Escuintla, Jutiapa, & San Marcos]

Phyla nodiflora (L.) Greene [Chiquimula]

Phyla scaberrima (A. L. Juss.) Moldenke [Alta Verapaz, Ret-
alhuleu, Sacatepéquez, & Sololé]

Phyla stoechadifolia (L.) Small [Alta Verapaz & Jutiapa]

Priva aspera H.B.K. [El Quiché, Escuintla, & Jalapa]

Priva mexicana (L.) Pers. [Escuintla, Jalapa, & Progreso]

Stachytarpheta jamaicensis (L.) Vahl [Jutiapa]

Verbena litoralis H.B.K. [Baja Verapaz, Chimaltenango, El
Quiché, Jalapa, Retalhuleu, & Sacatepéquez]

Verbena teucriifolia Mart. & Gal. [Guatemala, Huehuetenango,
San Marcos, Solold, & Totonicap4n]

Vitex kuylenii Standl. [Alta Verapaz]

BRITISH HONDURAS:
Citharexylum crassifolium Greem,
Citharexylum hirtellum Standl.
HONDURAS :
Aegiphila laxicupulis Moldenke

416 PH Y-T'0 LOG TA Vol. 23, no. 5

Aegiphila panamensis Moldenke
Bouchea nelsonii Grenz.
Callicarpa acuminata var. argutedentata Moldenke
Eriocaulon molinae L. 0. Williams -- delete the asterisk
Lantana achyranthifolia Desf.
Lantana glandulosissima Hayek [Atl4ntida]
Lippia cardiostegia Benth. [Intibuc4]
Phyla scaberrima (A. L. Juss.) Moldenke [Morazdn]
Priva mexicana (L.) Pers.
Stachytarpheta angustifolia (Mill.) Vahl
Vitex gaumeri Greerm. [Atl4ntida]
GULF OF HONDURAS ISLANDS:
Avicennia germinans (L.) L. [Soldier Cay]
Lantana involucrata var. odorata (L.) Moldenke [Caulker &
Long Cays]
NICARAGUA :
Aegiphila deppeana Steud.
Ghinia spicata (Aubl.) Moldenke
Stachytarpheta jamaicensis (L.) Vahl [Matagalpa]
Vitex kuylenii Standl.
PANAMA :
Aegiphila costaricensis Moldenke
BAHAMA ISLANDS:
Avicennia germinans (L.) L. [Cat]

CUBA:
Callicarpa resinosa Wright & Moldenke -- delete "Pinar del
Rio "
JAMAICA :

Lantana angustifolia x L. camara Adams*

Lantana angustifolia x L. trifolia Adams*

Lantana camara x L. insularis Adams*

Lantana involucrata x L. reticulata Adams+
VIRGIN ISLANDS:

Lantana involucrata f. rubella Moldenke [Anegada]
LEEWARD ISLANDS:

Duranta repens L. [Montserrat]

Holmskioldia sanguinea Retz. [Dominica]

Petrea kohautiana Presl [Marie Galante]

Vitex divaricata Sw. [Marie Galante]
NORTHERN SOUTH AMERICAN ISLANDS:

Lantana armata Schau. [Margarita]

Lantana involucrata L. [Orchila & Punta Brava]

Stachytarpheta coccinea Schau. -- to be deleted
COLOMBIAN CARIBBEAN ISLANDS:

Duranta coriacea Hayek [San Andres]

Lantana armata Schau. [San Andres]

Lantana maxima Hayek [San Andres]

Lippia alba (Mill.) N. E. Br. [San Andres]

1972 Moldenke, Fifth Summary Supplement 417

COLOMBIA :
Aegiphila elata Sw. [Arauca]
Eriocaulon spruceanum Te viviparum Moldenke — delete the as-

terisk
Paepalanthus andicola var. villosus Moldenke — delete the as-
terisk
Stachytarpheta cayennensis f, albiflora Moldenke [Antioquia]
Stachytarpheta elatior Schrad. [Vichada]

VENEZUELA :
Aegiphila glandulifera Moldenke [T4chira]
Aegiphila odontophylla Donn. Sm. [Mérida]
Clerodendrum speciosissimum Van Geert [Mérida]
Duranta coriacea Hayek -—— to be deleted
Duranta mutisii f. serrulata Moldenke [Aragua]
Duranta steyermarkii Moldenke [Mérida]
Gmelina arborea Roxb. [Barinas]
Holmskioldia sanguinea Retz. [Mérida]
Lantana armata Schau. [Gudérico, Mérida, & Trujillo]
Lantana armata var. velutina Moldenke (Mérida, T4chira, & Tru-
ee as{iiio),. PERS no ih
Lantana boyacana Moldenke [Aragua]
Lantana camara L. [Barinas & Mérida]
Lantana cujabensis Schau. [Barinas & Mérida]
Lantana fiebrigii Hayek [Mérida]
Lantana glutinosa Poepp. [Barinas]
Lantana maxima Hayek [Barinas]
Lantana trifolia f. hirsuta Moldenke [Mérida]
Lippia alba (Mill.) N. E. Br. [Mérida]
Lippia hirsuta L. f. —- to be deleted
Lippia micromera Schau, [Mérida]
Paepalanthus andicola var. villosus Moldenke [T&chira]
Stachytarpheta elatior var. jenmani Moldenke [Cojedes}
Syngonanthus huberi Ruhl. [Bolivar]
Tectona grandis L. f. [Trujillo]
xVerbena hybrida Voss [Mérida]
Verbena parvula Hayek [Mérida]
Verbena tenera Spreng. [Mérida]
FRENCH GUIANA:
Stachytarpheta urticaefolia (Salisb.) Sims
ECUADOR:
Duranta mutisii f. serrulata Moldenke [Loja]
PERU:
Aegiphila cordata Poepp. [Hudnuco]
Aegiphila haughtii Moldenke {San Martin]
Aegiphila spicata (Rusby) Moldenke [Hudmco & Loreto]
Avicennia germinans (L.) L. [Pancho Island]
Citharexylum flexuosum var. glaberrimum Moldenke [Ancash]#*

418 Pen yb OV LOG A Vol. 23, no. 5

Lantana zahlbruckneri Hayek [Ancash]
Verbena bonariensis L. [Arequipa & Lima]
Verbena cuneifolia Ruiz & Pav. [San Martin]
Verbena parvula parvula Hayek [Loreto]
Vitex cymosa Bert. [Tumbes]
Vitex gigantea H.B.K. [Tumbes]
Vitex schunkei Moldenke [Loreto]#

BRAZIL:
Aegiphila chrysantha Hayek [Pard4]
Aegiphila gloriosa var. paraénsis Moldenke [Pard]*
Aegiphila microcalycina Moldenke [Roraima]*
Amasonia lasiocaulos Mart. & Schau. [Acre]
Citharexylum reitzii Moldenke -—~ to be deleted
Eriocaulon dictyophyllum f. viviparum Moldenke [Mato Grosso]*
Eriocaulon kunthii Korn. — to be deleted
Eriocaulon spruceanum f. viviparum Moldenke [Mato Grosso]*
Lantana camara var. angustifolia Moldenke [Mato Grosso]
Lantana cujabensis Schau. [Acre]
Lantana micrantha Briq. [Mato Grosso]
Lantana trifolia L. [Acre]
Leiothrix hirsut: hirsuta f. vivipara Seed [Guanabara]*
Lippia sidoides Lippia sidoides Cham. [Maranhi
Paepalanthus di dichotomus var. ins ea Moldenke [Minas Ger-

ais ]*
Paepalanthus fulgidus var. zuloagensis Moldenke [Amaz6énas]*
Paepalanthus planifolius (Bong.) Korn. [Rondénia]
Paepalanthus rhizocephalus Alv. Silv. [Mato Grosso]
Petrea rivularis Moldenke [Amazénas]
Stachytarpheta angustifolia (Mill.) Vahl [Paré]
Stachytarpheta gesnerioides var. cuneata Schau. [Distrito Fed-
eral }

Stachytarpheta goyazensis Turcz. [Distrito Federal]
Stachytarpheta simplex Hayek [S&o Paulo]
Syngonanthus densifolius Alv. Silv. [Goids]
Syngonanthus helminthorrhizus (Mart.) Ruhl. [Distrito Federal]
Syngonanthus humboldtii var. glandulosus Gleason [Amazénas]
Verbenoxylum reitzii (Moldenke) Troncoso [Santa Catarina]*
Vitex panshiniana iana Moldenke [Mato Grosso]
Vitex triflora var. hirsuta Moldenke [Acre]*

BOLIVIA:
Aegiphila spicata (Rusby) Moldenke — delete the asterisk
Lantana chamissonis (D. Dietr.) Benth. [Tarija]
Verbena cabrerae Moldenke [Tarija]

PARAGUAY :
Verbena guaranitica (Troncoso) Moldenke
CHILE:

Junellia tridactylites (Lag.) Moldenke [Curicé]

1972 Moldenke, Fifth Summary Supplement 419

ARGENTINA:
Aloysia meyeri Moldenke { Jujuy]
Lantana grisebachii Stuck. [Jujuy]
Lantana hassleri Briq. [Corrientes]
Lantana minasensis var. hispida Moldenke [Corrientes]*
Verbena aristigera S. Moore [Corrientes]
Verbena glutinosa Kuntze [La Pampa]
Verbena guaranitica (Troncoso) Moldenke [Corrientes & Misiones]
Verbena litoralis var. caracasana (H.B.K.) Briq. [Tucumdn]
Verbena megapotamica Spreng. [Corrientes & Entre Rfos]
Verbena ovata Cham. [Corrientes]
Verbena parodii (Covas & Schnack) Moldenke [La Pampa]
Verbena perakii (Covas & Schnack) Moldenke [Catamarca & Cér-
doba
Verbena pulchra Moldenke [Corrientes]
Verbena radicata var. glabra (Hicken) Moldenke [Mendoza]* —
this is the corrected entry
Verbena tristachya Troncoso & Burkart [Lechiguanas Island]
MACARONESTA:
Lantana camara var. aculeata (L.) Moldenke [Gomera & Palma]
Phyla nodiflora (L.) Greene [Gomera & Gran Canaria]
Verbena bonariensis L. [Gomera]
Verbena supina L. [Fuertaventura, Lanzarote, & Tenerife]
ITALY:
Verbena bracteata Lag. & Rodr.
Vitex agnus-castus var. laciniosa Ces.*
SICILY:
Verbena supina f. erecta Moldenke
Vitex agmus-castus f. f. alba (West.) Rehd.
PANTELLERIA :
Verbena officinalis L.
MALTA:
Verbena officinalis L.
COMINO ISLAND:
Vitex agnmus-castus L.
GOZO ISLAND:
Verbena officinalis L.

Vitex agnus-castus L.
ALGERIA:
la nodiflora var. canescens (H.B.K.) Moldenke
TUNISIA:
Phyla nodiflora var. canescens (H.B.K.) Moldenke
Vitex agnus-castus L.
CYRENAICA:
Verbena officinalis L.
Vitex agnus-castus L.
MALI:
Avicennia africana P. Beauv. [Senegambia]

20 Pin wer Oo wore Ts Vol, 23, no. 5

Lippia abyssinica (Otto & Dietr.) Cuf. [Soudan]
SUDAN:

Clerodendrum cordifolium (Hochst.) R. Br. [Dafur]
ERITREA :

Clerodendrum discolor (Klotzsch) Vatke
ETHIOPIA :

Eriocaulon richardi Korn. —— to be deleted
REPUBLIC OF SOMALI:

Clerodendrum hildebrandtii Vatke

Lantana viburnoides var. velutina Moldenke

Lippia somalensis Lippia somalensis Vatke -- delete the asterisk

Vitex tangensis Gi Gurke
SENEGAL:

Lippia multiflora Moldenke
REPUBLIC OF GUINEA:

Lippia abyssinica (Otto & Dietr.) Cuf.
SIERRA LEONE:

Premna quadrifolia Schum. & Thonn.

Vitex | chrysocarpa hrysocarpa Planch.

Vitex ferruginea Schum. & Thonn,
IVORY COAST:

Vitex simplicifolia Oliv.
GHANA :

Lantana rugosa Thunb.

Lippia abyssinica (Otto & Dietr.) Cuf.

TOGOLAND :
Lippia abyssinica (Otto & Dietr.) Cuf.
NIGERIA :

Clerodendrum capitatum var. conglobatum (J. G. Baker) Thomas
Lantana rugosa Thunb. [Northern & Southern]

ZAIRE:

Premna quadrifolia Schum. & Thonn.
RWANDA :

Clerodendrum discolor (Klotzsch) Vatke
UGANDA :

Clerodendrum discolor var. oppositifolium Thomas
Clerodendrum myricoides var. camporum Gtirke
Clerodendrum schweinfurthii var. bakeri (Giirke) Thomas
CHOLE ISLAND:
Clerodendrum glabrum E. Mey.
KENYA:
Clerodendrum buchholzii Gurke
Clerodendrum capitatum (Willd.) Schum. & Thonn.
Clerodendrum cordifolium (Hochst.) R. Br.
Clerodendrum discolor var. oppositifolium Thomas
Clerodendrum fischeri Gurke
Clerodendrum makanjanum Winkler

1972 Moldenke, Fifth Summary Supplement 421

Clerodendrum scheffleri Girke
Clerodendrum triplinerve. Rolfe
Clerodendrum umbellatum Poir.
Lantana rhodesiensis Moldenke
Lippia grandifolia Hochst.
Lippia somalensis somalensis Vatke
Premna angolensis Giirke
Vitex negundo L.
ANGOLA:
Clerodendrum polycephalum J. G. Baker [Kongo]
ZAMBIA :
Clerodendrum capitatum var. conglobatum (J. G. Baker) Thomas
RHODESIA :
Clerodendrum capitatum (Willd.) Schum. & Thonn.
Kalaharia uncinata f. rubra Moldenke
MALAWI :
Clerodendrum discolor var. diimmeri Thomas
Kalaharia uncinata f. rubra Moldenke
NAMIBIA:
Clerodendrum discolor var. crenatum Thomas
Kalaharia uncinata f. rubra Moldenke
SWAZILAND:
Clerodendrum triphyllum (Harv.) H. H. W. Pearson
LESOTHO:
Clerodendrum triphyllum (Harv.) H. H. W. Pearson
Eriocaulon abyssinicum Hochst.
Eriocaulon bauri N. E. Br.
Eriocaulon dregei Hochst.
Verbena officinalis L.
SOUTH AFRICA:
Clerodendrum discolor var. crenatum Thomas [Transvaal]
Clerodendrum ternatum var. lanceolatum (Gtrke) Moldenke [Bech-
vanaland]}
Eriocaulon abyssinicum Hochst. [Cape Province, Natal, & Trans-
vaal |
Eriocaulon sonderiamum Korn. [Natal]
Lantana rugosa Thunb. [Cape Province]
Verbena rigida Spreng. [Orange Free State]
COMORO ISLANDS:
Avicennia marina (Forsk.) Vierh. [Euphrates]
Clerodendrum leucophloeum Balf. f.
Premna obtusifolia R. Br. [South]
Stachytarpheta jamaicensis (L.) Vahl [West]
IRAN:
Eriocaulon cinereum R. Br.
Vitex negundo L.
AFGHANISTAN:

Vitex negundo L.

22 P.HY-T0.L..0,G:1 4 Vol. 23, no. 5

PAKISTAN:
Caryopteris odorata var. integrifolia Moldenke [Northwestern

Provinces ]
Caryopteris paniculata C. B. Clarke [Poonch]
Gmelina arborea Roxb. [Northwestern Provinces]
Verbena officinalis L. [Karachi]

BANGLADESH:
Clerodendrum urticifolium (Roxb.) Wall.
NEPAL:

Caryopteris paniculata C. B. Clarke
Eriocaulon exsertum Satake*
Eriocaulon kathmanduense Satake*
Eriocaulon obclavatum Satake*
Eriocaulon trisectoides Satake
BHUTAN:
Caryopteris odorata (Hamilt.) B. L. Robinson
Clerodendrum kaempferi (Jacq.) Sieb.
INDIA:
Aloysia triphylla (L'Hér.) Britton [Madras]
Avicennia marina (Forsk.) Vierh. [New & West Islands]
Avicennia marina var. acutissima Stapf & Moldenke [Saurashtra]
Avicennia officinalis L. [Sagar Island]
Caryopteris grata Benth. [Kashmir]
Caryopteris odorata (Hamilt.) B. L. Robinson [Madhya Pradesh]
Caryopteris odorata var. integrifolia Moldenke [East Punjab &
Uttar Pradesh]
Caryopteris paniculata C. B. Clarke [Uttar Pradesh]
Clerodendrum inerme (L.) Gaertn. [Rajasthan; Manauli & New Is-
lands]
Clerodendrum kaempferi (Jacq.) Sieb. [Khasi States]
Clerodendrum macrostachyum Turcz. [Khasi States]
Clerodendrum paniculatum L. [Kerala]
Clerodendrum philippinum var. simplex Moldenke [Mysore]
Clerodendrum phlomidis L. f. [Saurashtra]
Clerodendrum urticifolium (Roxb.) Wall. [Assan]
Clerodendrum venosum Wall. [Khasi States]
Eriocaulon collinum Hook. [Kerala]
Eriocaulon leucomelas Steud. [Andhra Pradesh]
Eriocaulon luzulaefolium Mart. [Andhra Pradesh]
Eriocaulon nilagirense Steud. [Kerala]
Eriocaulon polycephalum Hook. f. — delete the asterisk
Eriocaulon robustum Steud. — delete the asterisk
Eriocaulon setaceum L. [Bihar]
Eriocaulon thwaitesii Korn. [Mysore]
Phyla nodiflora (L.) Greene [Sagar Island]
Premna barbata Wall. [Kashmir]
Premna latifolia var. viburnoides (Wall.) C. B. Clarke [Manauli

1972 Moldenke, Fifth Summary Supplement 423

Island]
Premna obtusifolia R. Br. [Hare Island]
Sphenodesme ferruginea (W. Griff.) Briq. [Assam]
Sphenodesme pentandra var. wallichiana (Schau.) Munir [Khasi
States
Stachytarpheta urticaefolia (Salisb.) Sims [Kerala]
Tectona grandis L. f. [Kerala]
Verbena officinalis L. [Bihar]
Vitex altissima f. alata (Willd.) Moldenke [Madras & Mysore]
Vitex pinnata L. [Gujarat]
CHAGOS ARCHIPELAGO:
Phyla nodiflora (L.) Greene [Diego Garcia]
Premna obtusifolia R. Br. [Diego Garcia]
Stachytarpheta jamaicensis (L.) Vahl [Diego Garcia]
CEYLON:
Lantana camara L.

BURMA:
Clerodendrum indicum f. semiserratum (Wall.) Moldenke [Upper

Burma }*
Clerodendrum macrostachyum Turcz. [Tenasserim]
Clerodendrum nutans Jack [Upper Burma]
Clerodendrum semiserratum Wall. -- to be deleted
Clerodendrum serratum (L.) Moon [Tenasserim]
Congea griffithiana Munir [Upper Burma]
Glossocarya mollis Wall. [Tenasserim]
Gmelina arborea var. glaucescens C. B. Clarke [Tenasserim & Up-
per Burma]

Holmskioldia sanguinea Retz. [Upper Burma]
Vitex axillaris Wall. —- to be deleted
Vitex limonifolia Wall. [Tenasserim]
Vitex peduncularis Wall. [Tenasserim]
Vitex scabra Wall. [Upper Burma]

NICOBAR ISLANDS:
Gmelina elliptica J. E. Sm.
Vitex negundo L.

MERGUI ARCHIPELAGO:
Clerodendrum paniculatum L. [Mergui]
Congea griffithiana Munir [Mergui]
Congea velutina Wight [Mergui & Tavoy]
Congea vestita W. Griff. [Mergui]
Sphenodesme ferruginea (W. Griff -) Briq. (Mergui & Tavoy]

Sphenodesme griffithiana Wight (Mergui & Tavoy]
Vitex axillaris Wall. [Tavoy]*

CHINA:
Caryopteris incana (Thunb.) Miq. [Kwangsi]
Caryopteris incana var. nana (Dreer) Moldenke {[Chekiang, Kiang-

si, & Kwangtung]

42h P Hit) T4OuL, On GoTok Vol. 23, no. 5

Caryopteris incana var. szechuanensis Moldenke [Szechuan]*
Caryooteris mongholica Bunge [Kansu]
Caryopteris terniflora Maxim. [Szechuan]
CHINESE COASTAL ISLANDS:
Caryopteris incana var. nana (Dreer) Moldenke [Honam]
THAILAND:
Callicarpa brevipetiolata Merr.
Clerodendrum godefroyi Kuntze
Clerodendrum schmidtii var. macrophyllum Moldenke*
Congea connata Fletcher -- delete the asterisk
Eriocaulon polycephalum Hook. f.
Eriocaulon robustum Steud.
KOH CHANG ISLAND:
Congea connata Fletcher
INDOCHINA :
Clerodendrum godefroyi Kuntze -- delete the asterisk
Eriocaulon cinereum R. Br. [North Vietnam]
Eriocaulon sexangulare L. [North Vietnam]
Sphenodesme involucrata var. paniculata (C. B. Clarke) Munir
{[Cochinchina

MALAYA:
Clerodendrum deflexum Wall. [Singapore]

Clerodendrum paniculatum L. [Malacca]
Gmelina elliptica J. E. Sm. [Penang & Singapore]
Peronema canescens Jack [Penang]
JAPAN:
Caryopteris incana f. macrophylla Moldenke*
Eriocaulon decemflorum f. aberans Satake*
RYUKYU ISLAND ARCHIPELAGO:
Vitex trifolia var. subtrisecta (Kuntze) Moldenke [Ishigaki]
FORMOSA :
Caryopteris incana var. nana (Dreer) Moldenke
PHILIPPINE ISLANDS:
Clerodendrum bethunianum Lowe [Samar]
Clerodendrum cumingianum Schau. [Luzon]
Clerodendrum curranii Elm. [Culién & Paragua]
Clerodendrum inerme (L.) Gaertn. [Palmas]
Clerodendrum intermedium Cham. [Basilan & Masbate]
Clerodendrum kaempferi (Jacq.) Sieb. [Luzon; delete "Negros"]
Clerodendrum klemmei var. puberulum Moldenke [Luzon]*
Clerodendrum minahassae var. brevitubulosum H. J. Lam [Biliran,
Mindoro, & Samar]
Clerodendrum minahassae var. grandicalyx Moldenke [Leyte]+*
Clerodendrum philippinum var. simplex Moldenke [Samar]
Clerodendrum trichotomum Thunb. [Sabtan]
Clerodendrum umbellatum var. speciosum (Dombrain) Moldenke [Lu-
zon

1972 Moldenke, Fifth Summary Supplement 2s

Holmskioldia sanguinea Retz. [Luzon]
BONIN ISLANDS:
Callicarpa glabra Koidz. (Chichijima]
GREATER SUNDA ISLANDS:
Avicennia marina f. angustata Moldenke (Sarawak]*
Avicennia officinalis L. [Sarawak]
Premna canescens Hansford [Java]*
WALLIS & FUTUNA ISLANDS:
Clerodendrum inerme (L.) Gaertn. [Futuna]
Stachytarpheta urticaefolia (Salisb.) Sims [Futuna & Wallis]
Vitex trifolia var. bicolor (Willd.) Moldenke [Futuna]
LOYALTY ISLANDS:
Vitex trifolia var. bicolor (Willd.) Moldenke [Urea]
NEW CALEDONIAN ISLANDS:
Gmelina lignum-vitreum Guillaum. [New Caledonia]*
Holmskioldia sanguinea Retz. [New Caledonia]

FIJI ISLANDS:
Faradaya neo-ebudica var. puberulenta (Moldenke) Moldenke [Tai-
levu]
AUSTRALIA:

Eriocaulon australe R. Br. [South Australia]

Eriocaulon cinereum R. Br. [South Australia]

Eriocaulon nanum R. Br. [South Australia]

Eriocaulon nutans F. Muell. [Northern Territory]*

Eriocaulon pallidum R. Br. [South Australia]

Eriocaulon stillulatum Hook. [Northern Territory]*
CULTIVATED:

Aloysia triphylla (L'Hér.) Britton [South Africa]

Bouchea fluminensis (Vell.) Moldenke [England]

Callicarpa dichotoma (Lour.) K. Koch [Russia]

Callicarpa japonica Thunb. [Russia]

Callicarpa japonica f. albibacca Hara [New Jersey]

xCaryopteris clandonensis Simmonds [Canada]

Caryopteris grata Benth. [Burma]

Caryopteris incana var. nana (Dreer) Moldenke — delete the

asterisk me

Clerodendrum aculeatum (L.) Schlecht. [Florida]

Clerodendrum glabrum E. Mey. [England]

Clerodendrum japonicum (Thunb.) Sweet [England]

Clerodendrum petasites (Lour.) S. Moore [Scotland]

Clerodendrum philippinum Schau. [Spain]

Clerodendrum philippinum var. simplex Moldenke [England]

Clerodendrum speciosissimum Van Geert [Philippines & Scotland]

Clerodendrum thomsonae Balf. f. [Scotland]

Clerodendrum thomsonae var. delectum L. H. Bailey [England]

Clerodendrum thomsonae var. variegatum L. H. Bailey [England]

Clerodendrum tomentosum (Vent.) R. Br. [England]

1,26 PY TO 10.0 7 & Vol. 23, no. 5

Clerodendrum trichotomum Thunb. [Canada & Russia]

Clerodendrum ugandense Prain [Jamaica]

Clerodendrum umbellatum var. speciosum (Dombrain) Moldenke
[England]

Congea tomentosa Roxb. [Martinique]

Duranta repens L. [Ghana]

Gmelina arborea Roxb. [Ghana & Venezuela]

Holmskioldia sanguinea Retz. [Burma, China, Colombia, Guatema-
la, Haiti, & Nepal]

Lantana camara L. [Russia]

Lantana camara var. alba Moldenke [Venezuela]

Lantana camara var. flava (Medic.) Moldenke [Venezuela]

Lantana camara var. mista (L.) L. H. Bailey [New Jersey & Ven-
ezuela]

Lantana camara var. mutabilis (Hook.) L. H. Bailey [Venezuela]

Lantana involucrata L. [Venezuela]

Lantana montevidensis (Spreng.) Briq. [Venezuela]

Lippia alba (Mill.) N. E. Br. [Jamaica]

Petrea arborea H.B.K. [England]

Petrea arborea var. broadwayi Moldenke [England]

Petrea kohautiana Pres] [England]

Petrea kohautiana var. alba (Freeman & Williams) Moldenke [Eng-
land

Petrea ee var. albiflora (Standl.) Moldenke [England]

Phyla nodiflora var. canescens (H.B.K.) Moldenke [Algeria &
Tunisia]

Premna esculenta Roxb. [England]

Premna latifolia Roxb. [England]

Premna obtusifolia R. Br. [England]

Tectona grandis L. f. [England & Ghana]

Teucridium parvifolium Hook. f. [New Zealand]

Verbena ambrosifolia Rydb. [Texas]

Verbena bipinnatifida Nutt. [Texas]

Verbena bracteata Lag. & Rodr. [Spain]

Verbena crithmifolia Gill. & Hook. [Argentina]

Verbena flava Gill. & Hook. [Argentina]

Verbena glutinosa Kuntze [Argentina]

Verbena hookeriana (Covas & Schnack) Moldenke [Argentina]

xVerbena hybrida Voss [Armenia, Australia, Belgium, Egypt,
Lithuania, Luxemburg, Malaysia, Maryland, Portugal, Rho-
desia, Romania, South Africa, & Tasmania|

Verbena microphylla H.B.K. [Argentina]

Verbena origenes R. A. Phil. [Argentina]

Verbena parodii (Covas & Schnack) Moldenke [Argentina]

Verbena perakii (Covas & Schnack) Moldenke [Argentina]

Verbena peruviana (L.) Britton [Hungary, Peru, Spain, & Tortola]

Verbena peruviana (L.) Britton x V. elegans H.B.K. [Massachusetts]

1972 Moldenke, Fifth Summary Supplement 27

Verbena peruviana (L.) Britton x ¥. moricolor Moldenke (Massa-

chusetts ]+
Verbena peruviana (L.) Britton x V. pulchella Sweet [Massachu-
setts ]+

Verbena phlogiflora Cham. [Formosa]

Verbena platensis Spreng. [Guatemala]

Verbena radicata Moldenke [Argentina]

Verbena rigida Spreng. [Jamaica]

Verbena santiaguensis (Covas & Schnack) Moldenke x V. peruvi-
ana ct Britton [Argentina]* "apa

Verbena santiaguensis (Covas & Schnack) Moldenke x V. phlogi-
flora Cham. [Argentina]* =

Verbena santiaguensis (Covas & Schnack) Moldenke x V. pul-

chella Sweet [Massachusetts ]#* “=

xVerbena teasii Moldenke [India]

Verbena tenera Spreng. [Spain]

Verbena tenuisecta Briq. [Guatemala & Honduras]

xVerbena wingei Moldenke [India]

xVeronicena haartmani Moldenke — to be deleted

Vitex agnus-castus L. [Canada, England, Jamaica, Luxemburg,
Mexico, Russia, & South Africa]

Vitex agnus-castus f. alba (West.) Rehd. [England]

Vitex negundo L. [Kenya & Russia]

Vitex negundo var. heterophylla (Franch.) Rehd. [Canada]

Vitex trifolia L. [England]

Addenda & errata to Part II:

Aegiphila chotzkiana Cham. = A. lhotzkiana Cham.

Aegiphila hoffmannioides Standl. & Steyerm. = ?Hoffmannia lenti-
cellata Hemsl., Rubiaceae

Aegiphila integrifolia (Jacq.) Jacq. = A. integrifolia (Jacq.)
Jacks.

Aegiphila knotzkiana Cham. = A. lhotzkiana Chan.

Aegiphila umbellata Vell. = Feramea corymbosa Aubl., Rubiaceae —
this is the corrected entry

Aegiphyla pendula Moldenke = Aegiphila pendula Moldenke
Agmus castus Cast. = Vitex agnus-castus L.

Aloysia gracillissima Mears = A. gratissima (Gill. & Hook.) Tron-

coso
Aloysia gratissima (Gill.) Troncoso = A. gratissima (Gill. & Hook.)
Troncoso

Aloysia hatschabachii Moldenke = A. hatschbachii Moldenke
Aloysia reichei Moldenke = A. reichii Moldenke

Avicennia officinalis Kurz = A. alba Blume

Avicennia officinalis sec. auct. afr. = A. marina (Forsk.) Vierh.
Callicarpa americana Sessé & Moc. = C. acuminata var. pringlei

(Briq.) Moldenke -- this is the corrected entry

428 PHYTOLOGIA Vol. 23, no. 5

Callicarpa bodinieri var. lyu (Léveillé) Rehd. = C. bodinieri var.
lyi (Léveillé) Rehd.
Sates ereoclona Hansford = C. erioclona Schau.
Sos formosensis Hansford = = Cs formosana Rolfe
Callicarpa japonica 'Leucocarpa' McGourty = C. japonica f. albi-
bacca Hara
Callicarpa oreoclona Hansford = C. erioclona Schau.
Callicarpa priglei B: Briq. = C. acuminata var. pringlei (Briq.)
Moldenke -- this is the corrected entry
Callicarpa pringlei Briq. = C. acuminata var. pringlei (Briq.)
Moldenke a
Callicarpa pringleii Briq. = C. acuminata var. pringlei (Briq.)
Moldenke —- this is the corrected entry
Callicarpa vestita Wall. ex C. B. Clarke = C. vestita Wall.
Callicarpa n. 9 Hook. f. & Thoms. = Caryopteris paniculata C. B.
Clarke
Caryopteris incams Miq. = C. incana (Thunb.) Miq.
Caryopteris mastacanthus var. clandonensi Barbey = x0. clandonen-
sis Simmonds
Caryopteris mongolica Maxim. = C. mongholica Bunge
Caryopteris nepataefolia ae = C. nepetaefolia (Benth.) Maxim.
Caryopteris odorata (Roxb.) B. Robinson = C. odorata (Hamilt.)
B. L. Robinson
Caryopteris serratum (L.) Moon = Clerodendrum serratum (L.) Moon
Cellicarpa Hansford = Callicarpa L.
Cellicarpa neene Hansford = Callicarpa arborea var. psilocalyx

(H. J. Lam) Moldenke
eee ae Mold. = Citharexylum B. Juss.

Citahrexylum reitzii Mold. = Verbenoxylum reitzii (Moldenke) Tron-
coso

Citharexylum cordatum Stevens = C. caudatum L.

Citharexylum fructiculosum L. = C. fruticosum L.

Citharexylum hintonii Moldenke = C. hintoni Moldenke

Citharexylum mocinnii var. longibracteolatum Moldenke = C. mocinni
var. longibracteolatum Moldenke a

Citharexylum reitzii Moldenke = Verbenoxylum reitzii (Moldenke)
Troncoso

Clandularia J. F. Gmel. = Verbena [Dorst.] L.

Clandularia caroliniensis J. F. Gmel. = Verbena canadensis (L.)
Britton

Clerodendrin Westcott = Clerodendrum Burm.

Clerodendrin thomsonae Westcott = Clerodendrum thomsonae Balf. f.

Clerodendron bethuneanum Law. = Clerodendrum bethunianum Lowe

Clerodendron bethunianum Lour. = Clerodendrum bethuniamum Lowe

Clerodendron blancoi Merr. = Clerodendrum minahassae var. brevitub-—
ulosum H. J. Lam

Clerodendron buchanani Roxb. = Clerodendrvm buchanani (Roxb.) Walp.

1972 Moldenke, Fifth Summary Supplement 429

Clerodendron bungeana Ledin = Clerodendrum bungei Steud.
Clerodendron castaneaefolium Klotzsch = Clerodendrum viscosum

Vent.
Clerodendcron commersonii (Lam.) Spreng. = Clerodendrum inerme (L.)
Gaertn. eae ea tee
Clerodendron discolor Klotzsch = Clerodendrum discolor (Klotzsch)
Vatke

Clerodendron formicarium Hansford = Clerodendrum triplinerve Rolfe

Clerodendron glabrum angustifolia Sims = Clerodendrum glabrum E.
Mey.

Bete tantion glandulosum Wall. = Clerodendrum colebrokiamum Walp.

Clerodendron gratum Kurz = Caryopteris paniculata C. B. Clarke

Clerodendron gratum Wall. = aryopteris grata Benth.

Clerodendron hereophyllum Miq. = Peronema canescens Jack

Clerodendron inermis Gaertn. = Clerodendrum inerme (L.) Gaertn.

Clerodendron macrostachyum Turcz. = Clerodendrum m macrostachyum
Turez.

Clerodendron paniculata L. = Clerodendrum paniculatum L.

Clerodendron petasites S. Moore = Clerodendrum petasites (Lour.) S.
Moore

Clerodendron robecchii Chiov. = Clerodendrum robecchii Chiov.

Clerodendron robecchii var. macrophyllum Chiov. = Clerodendrum
robecchii var. macrophyllum Chiov.

Clerodendron sansibarense Gurke = Clerodendrum sansibarense Giirke

Clerodendron semiserrata Wall. = Clerodendrum indicum f. semiser-
ratum (Wall ~) Moldenke

Clerodendron semiserratum Wall. = Clerodendrum indicum f. semi-
serratum (Wall. y Moldenke

Clerodendron similis Merr. = Clerodendrum mindorense Merr.

Clerodendron siphonanthus var. semiserrata C. B. Clarke = Clero-
dendrum indicum f. semiserratum (Wall.) Moldenke -- this is
the corrected entry

Clerodendron siphonanthus var. semiserrata (Wall.) C. B. Clarke =
Clerodendrum indicum f. semiserratum (Wall.) Moldenke

Clerodendron s Squamatum var. urticifolia C. B. Clarke = Cleroden-
drum urticifolium (Roxb.) Wall.

Clerodendron thompsoni: thompsonii Hook. = Clerodendrum thomsonae Balf. f.

Clerodendron thomsoniae x C. splendens Adams = Clerodendrum umbel-
latum var. r. speciosum (Dombrain) Moldenke

Clerodendron trichostomum Hansford = Clerodendrum trichotomum
Thunb,

Clerodendron ?trichotomum Wall. = Clerodendrum villosum Blume

Clerodendron verticillata Don = Clerodendrum indicum (L.) Kuntze

Clerodendron yambesiacum Baker = Clerodendrum guerkei J. G. Baker

Clerodendron sp. L. = Clerodendrum infortunatum L.

Clerodendron Sp. n. 563 W. Griff. = Clerodendrum bracteatum Wall.

Clerodendrum acerbianum (Vis.) Benth. & Hook. = C. acerbiamum (Vis-

430 PHYTOL OG 1 & Vol. 23, no. 5

ian.) Benth.
Clerodendrum buchananii Roxb. = C. buchanani (Roxb.) Walp.
Clerodendrum inermi Farnsworth = C. inermis (L.) Gaertn.
Clerodendrum macrosiphon Hook. = Ce inciswm var, macrosiphon

(Hook. f. y J. G. Baker

Clerodendrum minnahassee Buswell = C. minahassae Teijsm. & Binn.

Clerodendrum phlomides L. f. = C. phlomidis L. f.

Clerodendrum phlomoides L. = C. phlomidis L. f.

Clerodendrum speciosissimum Van Beert. = C. speciosissimum Van
Geert

Clerodendrum triplinerva Rolfe = C. triplinerve Rolfe

Clerodendrum ugandensis Prain = C. ugandense Prain

Cmelina Caaudhuri = Gmelina L.

Congea tomentosa var. coerulea (Griff.) Clarke = C. tomentosa Roxb.

Congea velutinus Wight = C. velutina Wight

Congeae sp. W. Griff. = Sphenodesme involucrata (Presl) B. L. Rob-
inson

Cornutia grandifolia var. quadrangularis Moldenke = C. grandifolia
var. quadrangularis %rst. & Moldenke

Cornutia pyramidata var. ismithia Moldenke = C. pyramidata var.
isthmica Moldenke

Diototheca repens Raf. = Phyla nodiflora var. reptans (Spreng.)
Moldenke -- this is the corrected entry

Duranata Dennis = Duranta L.

Duranata mitisii Dennis = Duranta mutisii L. f.

Duranta plunieri "A. B." = D. repens L.

Elaeagnon theophrasti Lob. = Vitex agnus-castus L.

Erimus laciniatum L. = Verbena laciniata (L.) Briq.

Erimus peruviams L. = Verbena peruviana (L.) Britton

Eriocaulon callocephalum Alv. Silv. = E. liguiatun (Vell.) L. B.
Sm. — this is the corrected entry

Eriocaulon elichrysoides Kunth = E. ligulatum (Vell.) L. B. Sm. —
this is the corrected entry

Eriocaulon heudeloti N. E. Br. = E. heudelotii N. E. Br.

Eriocaulon kunthii Korn. = E. ligulatum (Vell.) L. B. Sm. — this
is the corrected entry

Eriocaulon kunthii var. j Korn. = E. ligulatum (Vell.) L. B. Sm. —
this is the corrected entry

Eriocaulon kunthii var. @ Korn. = E, ligulatum (Vell.) L. B. Sm.
=- this is the corrected entry _

Eriocaulon kunthii var. ? Korn. = E. ligulatum (Vell.) L. B. Sm. —
this is the corrected entry

Eriocaulon kunthii var. Y Korn. = E. ligulatum (Vell.) L. B. Sm. —
this is the corrected entry

Eriocaulon richardi Korn. = E. abyssinicum Hochst.

Eriocaulon richardii Korn. = Es abyssinicum Hochst. -- this is the
corrected entry

1972 Moldenke, Fifth Summary Supplement 431

Eriocaulon sexangulare Auct. [ex Cuf.] = E. abyssinicum Hochst.

Eriocaulon sexangulare Auct. [ex Ruhl.] = ie cinereum R, Br. —
this is the corrected entry

Eriocaulon willdenowianum Moldenke = E. willdenovianum Moldenke

Gesnera pilosa Glaz. = Amasonia hirta B Benth.

Gesneria pilosa Glaz. = = Amasonia hirt: hirta Benth.

Ghinia spicata Moldenke = G. = G. spicata ata (Aubl.) Moldenke

Glandularia J. G. Agardh -- a section of Sargassum C, Agardh, Fu-
caceae

Glandularia ambrosifolia Schnack & Covas = Verbena ambrosifolia
Rydb.

Glandularia cabrerae (Mold.) Tronc. = Verbena cabrerae Moldenke

Glandularia canadensis L. Small = Verbena canadensis (L.) Britton

Glandularia dissecta (Willd. ex Spreng. eng.) Schnack & Covas = Verbena
dissecta Willd.

Glandularia elegans Solbrig = Verbena elegans H.B.K.

Glandularia elegans x peruviana Solbrig -—- see under Verbena ele-
gans H.8.K. x V. peruviana (L. ) Britton

Glandularia e elegans x x p pulchella Solbrig -- see under Verbena ele-
gans H.B.K. x V. pulchella Sweet

Glandularia elegans x stellaroides Solbrig — see under Verbena
elegans H.B.K. x V. stellarioides Cham.

Glandularia megapotdmica Schnack & Covas = Verbena megapotamica
Spreng.

Glandularia microphylla Schnack & Covas = Verbena microphylla H.B.K.

Glandularia peruviana x megapotémica Schnack & Covas = xVerbena
schnackii Moldenke

Glandularia pinnaliloba Schnack & Rubens = Verbena pinnatiloba
(Kuntze) Moldenke

Glandularia pulchra Mold. = Verbena pulchra Moldenke

Glandularia pulchella x peruviana viana Solbrig — see under Verbena

ruviana (L.) Britton x V. pulchella Sweet

Glandularia santiaguenensis Covas & Schnack = Verbena santiaguen-

sis (Covas & Schnack) Moldenke

Glandularia santiaguensis x G. peruviana Schnack & Covas -- see un-
der Verbena santiaguensis (Covas & Schnack) Moldenke x v.

peruviana ana (L.) Britton

eects oe rn (Cham.) Tronc. = Verbena sessilis (Cham.)

Kuntze
Glandularia sessilis Schnack & Rubens = Verbena sessilis (Cham.)
Kuntze

Glandularia stellatioides (Cham.) Schnack & Covas = Verbena stel-
larioides Cham.
Glandularia sulfurea (D. Don) Schnack & Covas = Verbena sulphurea

D. Don
Glandularia temiisecta Small = Verbena temuisecta Briq.

432 PHYTOLOGIA Vol. 23, no. 5

Glossocarya linnaei Benth. = G. scandens (L. f.) Moon

Gmelina asiatica Wall. [in part] = G. elliptica J. E. Sm.

Gmelina filipensis Cham. = G. philippensis Cham,

Gmelina sp. n. 2 W. Griff. = G. elliptica J. E. Sn.

Hastingia coccinea Konig = Holmskioldia sanguinea Retz.

Hastingia scandens Roxb. = Holmskioldia sanguinea Retz.

Hippia aegyptiaca Delchev. = Phyla nodiflora var. reptans (Spreng.)
Moldenke -- this is the corrected entry

Holmskioldia sangiunea Retz. = H. sanguinea Retz.

Holmskoéldia Retz. = Holmskioldia Retz.

Holmskoldia re Retz. = Holmskioldia sanguinea Retz.

Hymenofyram s Wall. = Hymenopyramis Wall.

Hymenofyranus siamensis Wall. = Hymenopyramis siamensis Craib

Hymenospyranis Wall. = Hymenopyramis Wall.

Hymenospyranis brachiata Wall. = Hymenopyramis brachiata Wall.

Junellia glauca var. cisandina Niederl. = J. glauca var. cisan-
dina (Niederlein) Moldenke

Lantana cujabensis var. punctuata Moldenke = L. cujabensis var.
punctata Moldenke

Lantana horrida var. parviflora Schau. = L. horrida H.B.K.

Lantana jamaicensis x reticulata Adams = L. fucata var. antillana
Moldenke

Lantana lasiocarycina Goodland = Lippia lasiocalycina Cham.

Lantana megapotamica megapotamica (Spreng.) Tronc. = L. montevidensis (Spreng.)

Taare ae
Lantana mondevidensis (Spreng.) Briq. = L. montevidensis (Spreng.)
Briq.

Lantana nana Price = L. camara var. hybrida (Neubert) Moldenke
Lantana ovatifolia var. ovatifolia Britton = L. ovatifolia Britton
Lantana ovatifolia var. reclinata R. W. Long = = L. depressa depressa Small
Lantana repens Sessé & Moc. = Phyla nodiflora var. reptans
Spreng.) Moldenke -- this is the corrected entry
Lantana reticulata x jamaicensis Stearn = L. fucata var. antilla-
ees Moldenke
Lantana semantha Pettingill = L. camara L.
Lantana urticifolia Mill. = L. “urticaefolia Mill.
Lantana zahlbruchneri Hayek = = Le zahlbruckneri Hayek
Lippia Endl. = Phyla Lour.
Lippia aegyptiaca Carr. = Phyla nodiflora var. reptans (Spreng.)
Moldenke -—- this is the corrected entry
Lippia aegyptiaca Delchev. = Phyla nodiflora var. reptans
(Spreng.) Moldenke —- this is the corrected entry
Lippia alba (Mill.) N. E. Briq. = L. alba (Mill.) N. E. Br.
Lippia asperifolia asperifolia A. Rich. ex Marthe = L. javanica
(Burm. f.) Spreng.
Lippia bolandieri Schau. = L. graveolens H.B.K.

1972 Moldenke, Fifth Summary Supplement 433

Lippia canescens Rich. = Phyla nodiflora var. reptans (Spreng.)
Moldenke -- this is the corrected entry
Lippia canescens Robinson = Phyla nodiflora var, reptans (Spreng.)
Moldenke —— this is the corrected entry
Lippia citridora H.B.K. = Aloysia triphylla (L'Hér.) Britton
Lippia dulcis mexicana Kraemer = Phyla scaberrima (A. L. Juss.)
Moldenke
Lippia gerhtii Moldenke = L. gehrtii Moldenke
Lippia gerthii Angely = L. gehrtii Moldenke
Lippia grandifolia Hochst. ex A. Rich. = L. grandifolia Hochst.
Lippia lanceolata Rose = Phyla nodiflora var. reptans (Spreng.)
Moldenke -- this is the corrected entry
Lippia ligustrina Kearney & Peebles = Aloysia gratissima (Gill. &
Hook.) Troncoso
Lippia nodiflora Eggers = Phyla nodiflora var. reptans (Spreng .)
Moldenke -- this is the corrected entry
Lippia nodiflora Millsp. = Phyla nodiflora var. reptans (Spreng.)
Moldenke -- this is the corrected entry
Lippia nodiflora Robinson & Greenm. = Phyla nodiflora var. rep-
tans (Spreng.) Moldenke — this is the corrected entry
Lippia a nodiflora var. repanda (H.B.K.) Kuntze = Phyla nodiflora
var. reptans (Spreng.) Moldenke -- this is the corrected en-

try

Lippia nodiflora nodiflora var. reptans (H.B.K.) Kuntze = Phyla nodiflora
var. r. reptans (Sore "Headaches -- this is the corrected en-
try

Lippia reptans Humb. = Phyla nodiflora var. reptans (Spreng.)
Moldenke -- this is the corrected entry

Lippia reptans Humb. & Bonpl. = Phyla nodiflora var. reptans
fereste-) Moldenke -- this is the corrected entry

Lippia reptans H.B.K. = Phyla nodiflora var. reptans (Spreng.)

Moldenke -- this is the corrected entry

Lippia reptans Humb. & Kunth = Phyla nodiflora var. reptans
Spreng.) Moldenke — this is the corrected entry

Lippia reptans Kunth = Phyla nodiflora var. reptans (Spreng.)
Moldenke — this is the corrected entry

Lippia reptans L. = Phyla nodiflora var. reptans (Spreng.) Molden-
ke -- this is the corrected entry

Lippia reptans (Spreng.) H.B.K. = Phyla nodiflora var. reptans
(Spreng.) Moldenke

Lippia saturejaefolia Mart. & Schau. = L. satureiaefolia Mart. &
Schau.

Lippia seriphioides (Mold.) A. Gray = Acantholippia seriphioides
(A. Gray) Moldenke

Lippia trifida var. gracilis Phil. = cantholippia deserticola
(R. A. Phil.) Moldenke

Litsea luzonica Blanco = Symphorema luzonicum (Blanco) Fern.-Vill.

Luehea speciosa Willd. -- in the Tiliaceae

43h PHYTOL OG? & Vol. 23, no. 5

Mesanthemum rosenii Pax = M, roseni Pax.

Nyctanthaceae Kundu = Nyctanthaceae Kundu & De

Nyctanth|acjeae J. G. Agardh = Nyctanthaceae Kundu & De — this is
the corrected entry

Nyctantheae J. G. Agardh = Nyctanthaceae Kundu & De — this is the
corrected entry

Obletia verbenolacoea Rozier = Verbena canadensis (L.) Britton

Paepalanthus calaussiamus Korn. = P. claussenianus Korn.

Paepalanthus lamarckii Knuth = P. lamarckii Kunth

Petraea violacea Angely = Petunia violacea Lindl., Solanaceae

Petraevitex trifolia Merr. = Petraeovitex trifoliata Merr.

Petrea arborea var. Broadway Chittenden = P. arborea var. broad-
wayi Moldenke

Petrea aspera L. = P. aspera Turcz.

Petrea kohautiana var. anomalo Chittenden = P. kohautiana var.
alba (Freeman & Williams) Moldenke

Phyla Lourt. = Phyla Lour.

Phyla betulaefolia Greene = P. betulaefolia (H.B.K.) Greene

Phyla nodiflora var. reptans (H.B.K.) Moldenke = P. nodiflora

var. reptans (Spreng.) Moldenke

Phyla nodiflora reptans (H.B.K.) Moldenke = P. nodiflora var.
reptans (Spreng.) Moldenke -- this is the corrected entry

Phyla nodifloros var. reptans (H.B.K.) Moldenke = P. nodiflora
var. reptans (Spreng.) Moldenke -- this is the corrected
ent

Phyla mre) Greene = P. nodiflora var. reptans (Spreng.) Mol-
~ denke -— this is the corrected entry

Phyla reptans H.B.K. = P. nodiflora var. reptans (Spreng.) Mol-

denke
Phyla reptans (H.B.K.) Greene = P. nodiflora var. reptans (Spreng.)

Moldenke -- this is the corrected entry
Phyla stochadaefolia (L.) Small = P. stoechadifolia (L.) Small
Phyla strigosa (Mart. & Gal.) Moldenke = P. strigulosa (Mart. &
~ Gale) Moldenke
Phyla strigulosa var. parviflora Moldenke = P. strigulosa var.
sericea (Kuntze) EEPa es
Phyla yucata yucatana var. parviflora Moldenke = P, strigulosa var. ser-
icea cea (Kuntze) Moldenke
Phylla incisa Williams = Phyla incisa Small
Phylla scaberrima (A. L. Juss.) Moldenke = Phyla scaberrima (A.

L. Juss.) Moldenke

Premna integrefolia L. = P, obtusifolia R. Br.

Salix amerina Diosc. = Vitex agnus-castus L.

Sezegleewia luconiensis Turcz. = Symphorema luzonicum (Blanco)
Fern.-Vill.

Sherardia nodiflora, fruticosa, foliis subrotundis serratis Houst.

[in part] = Ph Phyla nodiflo: nodiflora var. _ var. reptans (Spreng.) Moldenke —

1972 Moldenke, Fifth Summary Supplement 435

this is the corrected entry
Spartothamnella juncea Briq. = S. guncea (A. Cunn.) Briq.
Sphenodesma ? Wall. = Sphenodesme eryciboides Kurz
Sphenodesma acuminata acuminata Wight = Sphenodesme pentandra Jack
Sphenodesma eryciboides Kurz = Sphenodesme eryciboides Kurz

Sphenodesma ferrugineum Wight = Sphenodesme racemosa (Presl) Mol-
denke

Sphenodesma grossum Kurz = Sphenodesme eryciboides Kurz
Sphenodesma jackianum Wight = Sphenodesme griffithiana eso ie
Stachytarphaeta cayen oa ae (L. C. Rich.) Vahl = Stachytarphe

L

cayennensis C. Rich.) Vahl
eae tete ti hintonii Moldenke = S. hintoni Moldenke
Stilbe albiflora > pilosa A. DC. = as albiflora E. Mey.

Symphar ema Vill. = Symphorema Roxb,

Sympharema luzonicum (Blanco) Vill. = Symphorema luzonicum (Blan-
co) Fern.-Vill.

Symphorema involucratum Kew = S. polyandrum Wight

Symphorena F.-Vill. = Symphorema Roxb.

Symphorena luzonicum F.-Vill. = Symphorema luzonicum (Blanco)

Fern.-Vill.
Syngonanthus ceranthemoides (Bong.) Ruhl. = S. xeranthemoides
(Bong.) Ruhl.

eee a densiflorus var. gabresc Anon. = S. densiflorus
+ glabrescens brescens Moldenke

ee Beha pinrisaeie Imam = T. philippinensis Benth.
Tetrandra, spicis filiformibus paniculatis, foliis multifido-

a PEP caule solitario Willd. = Verbena officinalis L.
Teucridium paucifolium A. Cunn. = T. parvifolium Hook. f.
Timotecia Moldenke = Casselia Nees & Mart.
Timotecia confertiflora Moldenke = Casselia confertiflora (Mol-

denke) Moldenke
Urbania eganioides Phil. = U. egaflioides R. A. Phil.
] Lis

Veraena Lam. = Verbena [Dorst.

Veraena erinoides Lam. = Verbena laciniata (L.) Briq.

Verbena americana, spici multiplici, foliis urticae angustissimis,
floribus purpureis Tourn. = V. hastata L.

Verbena aukletia Jacq. = V. canadensis (L.) Britton

Verbena beipinnalifida Nutt. = V. bipinnatifida Nutt.

Verbena bininnafillia Abrigo = V. bipinnatifida Nutt.

Verbena bipinnatifolia Nutt. = V. bipinnatifida Nutt.

Verbena bracteosa x hastata Rydb. = xV. crookshanksi Moldenke —
this is the corrected entry

Verbena braliliense Vell. = V. brasiliensis Vell.

Verbena clemensorum Moldenke = xV. clemensorum Moldenke

Verbena communis caeruleo flore Zannich. = V. officinalis L.

4,36 PHYT@LOG IA Vol. 23, no. 5

officinalis L.

Verbena communis, flore caeruleo Cup. = V. officinalis L.

Verbena amend eee Grey = V. canadensis (L.) Britton

Verbena elata Jan = V. alata Sweet

Verbena ericoides Macself =~ V. tenuisecta Brig.

Verbena foemina Brunf. = Senecio vulgaris L., Carduaceae

Verbena foemina Trag. = Sisymbrium officinale L., Brassicaceae

Verbena foliis laciniatis s Ss superioribus, tripartitis, spicis an-
~ gustis longissimis Haller = V. officinalis L. is

Verbena foliis tripartitis, rugosis, spicis nudis gracillimis Hal-
ler = V. officinalis L.

Verbena fruticosa Mill. [in part] = Phyla nodiflora var. reptans
(Spreng.) Moldenke -- this is the corrected entry

Verbena fruticosa Millsp. [in part] = Phyla nodiflora var. reptans
(Spreng.) Moldenke -- this is the corrected entry

Verbena frutieosa Mill. [in part] = Phyla nodiflora var. reptans
(Spreng.) Moldenke -- this is the corrected entry

Verbena frutieosa Mill. & Houst. [in part] = Phyla nodiflora var.
reptans (Spreng.) Moldenke -- this is the corrected entry

Verbena glauca Gill. = Junellia selaginoides (Kunth) Moldenke

Verbena globifera Nocca = Phyla nodiflora var, reptans (Spreng .)
Moldenke -- this is the corrected entry

Verbena gooddingi nepetifoloia Tidestr. = V. gooddingii var.
nepetifolia Tidestr. a

Verbena goodmanii Moldenke = xV. goodmani Moldenke

Verbena hastata f. hastata Seymour = V. hastata L.

Verbena hastato-urticifolia Bebb = xV. , engelmannii Moldenke

Verbena hybrida cv. drandiflora Hort. = xV. hybrida Voss

Verbena lusitanica, latifolia, procerior Tourn. = V. officinalis
Ibe

Verbena macdougalli Shinn = V. macdougalii Heller

Verbena megapotamica var. phlogiflora Cham. = V. phlogiflora Cham.

Verbena melindre Schau. = V. peruviana (L.) Britton

Verbena officinalis L. x Veronica maritima L. = Veronica spuria L.,
Scrophulariaceae -- this is the corrected entry

Verbena offinalis Cham. = V. officinalis L.

Verbena offinalis var. glabrescens Cham. = V. glabrescens (Cham.)
Herter

Verbena peruviana Moldenke = V. peruviana (L.) Britton

Verbena peruviana 'Chiquita' Graf = V. tenera var. maonetti Regel

Verbena phlogiflora Cham. emend. Moldenke = V. phlogiflora giflora Cham.

Verbena plicata var. plicata Devor = V. plicata Greene

Verbena recta sive mas Fuchs = Sisymbrium officinale L., Brassica-
ceae

Verbena 3: sacra Gerarde = V. supina L. — this is the corrected entry

Verbena Santiaguensis Darlington . & Wylie = V. santiaguensis (Covas &

1972 Moldenke, Fifth Summary Supplement 437

Schnack) Moldenke

Verbena sphedroides Angely = V. ephedroides Chan.

Verbena sulphurea f. fuscorubra Skottsberg = V. sulphurea var.
fuscorubra Skottsberg

Verbena supina Dodon. = V. supina L.

Verbena tenera mahonetti Hay & Synge = V. tenera var. maonetti
Regel ee ea = Tee noe Ge

Verbena tenuifolia Tourn. = V. supina L.

Verbena tetrandra spicis filiformibus, paniculatis, foliis multi-
fido laciniatis, caule solitario L. = V. officinalis L.

Verbena townsendi Svenson = V. townsendii Svenson

Verbena triphylla Lam. = Aloysia triphylla (L'Hér.) Britton

Verbena urticae-folia, canadensis, foliis incisis, flore majore
Tourn, = V. officinalis L.

Verbena urticifolia angustifolia Bebb = xV. moechina Moldenke

Verbena urticifolio-hastata Bebb = V. hastata L.

Verbena vulgo appellata Ang. = V. officinalis L.

Verbena n. 219 Haller = V., officinalis L.

Verbenaca recta Dodon. = Verbena officinalis L.

Verbenaca supina Dodon. = Verbena supina L.

Verbenaca supina Gesn. = Ajuga sp., Lamiaceae -—- this is the
corrected entry

Verbenaca supina sive foemina Fuchs = Verbena supina L.

Verbenaca volgare del Matthioli Pon. = Verbena officinalis L.

densis (L.) Britton
Verbenna frummondii Baxt. = Verbena canadensis (L.) Britton
Verbina officinalis L. = Verbena officinalis L.
Vermicularia Tode -- in the Melanconiaceae
Vermicularia Tode ex Fries = Vermicularia Tode, Melanconiaceae

Vermicularia pseudosphaeria Tode -- in the Melanconiaceae
Vermicularia pubescens Tode -- in the Melanconiaceae

Veronica spuria Haartm. = V. spuria L., Scrophulariaceae --
this is the corrected entry

Vervena Kundu & De = Verbena [Dorst.] L.

Viter Aubreville = Vitex Tourn.

Viter fosteri Aubreville = Vitex ferruginea Schum. & Thonn.

Vitex Trag. = Vitex Tourn.

Vitex agnus castus var. trifolia Kurz = V. trifolia L.

Vitex agnus-castus var. latifolia (Mi11.) Loud. = V. agnus-castus
f. latifolia (Mill.) Rehd.

Vitex agnus-castus var. rosea Rehd. = V. agnus~castus f. rosea
Rehd.

Vitex agnus-castus # laciniosa Ces. = V. agnus-castus var.
laciniosa Ces. ey

Vitex agnus-castus 'Latifolia' McGourty = V. agnus-castus f. lat-

438 PLB-Y 20) GONG. 5 ok Vol. 23, no. 5

ifolia (Mill.) Rehd.

Vitex altissima (f. subglabra) Thwaites = V. altissima f. subgla-
bra Thwaites

Vitex cunninghamii Schau. = V. glabrata R. Br.

Vitex gigantia H.B.K. = V. gigantea H.B.K.

Vitex heterophylla Kurz = =v. quinata (Lour.) F. N. Will.

Vitex laciniosa Arcang. = ‘aA agmus-castus var. laciniosa Ces.

Vitex negundo "Heterophylla" Sherk & Buckley = V. negundo var.

heterophylla (Franch.) Rehd.

Vitex sex-dentata Wall. = Caryopteris grata Benth.

Vitex n. 12 Hook. f. & Thoms. = V. urceolata C. B. Clarke

Vitex n. 18 Hook. f. & Thoms. = Ve leucoxylon L. f.

Vitex nn. 10 and 18 Hook. f. & Thoms. = V. glabrata var. bombaci-
~~ folia (Wall.) Moldenke

Vitex nn. snnerL5, 16 Hook. f. & Thoms. = V. gamosepala W. Griff.

Volkameria urticifolia Wall. = Clerodendrum urticifolium (Roxb.)
Wall.

Volkameriae sp. W. Griff. = Clerodendrum venosum Wall. & Caryop-
teris odorata (Hamilt.) B. L. Robinson

Zapani ia nodiflora | Spreng. = Phyla nodiflora var. reptans (Spreng .)
meiteres — this is the corrected entry

Zapania reptans Spreng. = Phyla nodiflora var. reptans (Spreng .)
Moldenke -- this is the corrected entry

Addenda and errata to Parts IV, V, VI, VII, VIII, and X will not
be published at this time, with the single exception that an
inaccurate statement on page 786 should be corrected. Dr.
Otto Degener informs me that it is not his policy to use a
double author citation when a taxon is shifted from a binom-
ial with an incorrectly spelled generic portion to one with
the "corrected" generic name. He informs me that if he did
this in any instance in the past it was merely in an effort
to comply with the International Code as it existed at that
time. This, then, conforms with my policy also in such cases
where there are "orthographic variants" of generic names,
e.g., Clerodendron vs. Clerodendrum and Sphenodesma vs.
Sphenodesme.

It is perhaps worthwhile to mention here that a few complete copies
of the original "Fifth Summary", 97) pages, in 2 volumes, are
still available and may be ordered direct from either Editor
of PHYTOLOGIA for $25 plus 80 cents postage & wrapping (U.S.A.
addresses) or $1.62 postage & wrapping (foreign addresses) .

NEW NAMES AND A NEW COMBINATION IN SOUTH AMERICAN LINUM (LINACEAE)

R. Mildner & C. M. Rogers, Wayne State University, Detroit, Mich.

During the course of revising the native South American
species of Linum, the following new names and new combination have
been found necessary.

L. SMITHII Mildner, sp. nov. Species affinis L. organensi a qua
stipulis glandulosis, sepalis denticulatis, foliis in inflores-
centia redactis differt. Type: Reitz & Klein 7999, Brazil: Bom
Jardin, Santa Catarina (holotype: UC!; isotypes: G!, NY!, US!).
The specific epithet is given in honor of Lyman B. Smith, one of
the collectors, and, among his many interests, a student of the
flora of Santa Catarina.

L. smithii is found in eastern portions of the states of
Santa Catarina and Rio Grande do Sul in southern Brazil.

L. BURKARTII Mildner, sp. nov. Herba 18-50 cm. alta; sepala
ovata, acuminata, 3.7-4.4 mm. longa, glandulosa-denticulata;
stipulae glandulosae nullae; styli liberi, 3.5-4.5 mm. longi;
fructus ovatus, acutus, 2.2-2.5 mm. longus. Type: Rosengurtt
5755, Uruguay: Florida (holotype: SP!; isotype: MVM!). The
specific epithet is given in honor of Arturo Burkart, longtime
student of the flora of Argentina.

L. burkartii is found in southern Uruguay and in Entre Rios
and Buenos Aires provinces of eastern Argentina.

L. PROSTRATUM Domb. ex Lam. var. PARVUM (Johnst.) Mildner, comb.
nov. Type: Johnston 3549, Peru: Mollendo (holotype: GH!)
L. parvum Johnst., Cont. Gray Herb. 85: 172. 1929.

L. MACRAEI Benth. var. MARTICORENAE Mildner, var. nov. A var.
macraei caulibus pubescentibus, habitu humili et compacto
differt. Type: Mildner 15a, Chile: open grassy fields and road-
cuts, about 23 km. from Concepcion on road to Chillan (holotype:
US!; isotypes: CONC!, K!, UC!, WUD!). The varietal epithet is
given in honor of Prof. C. Marticorena, student of the flora of
Chile, who was most helpful to the senior author while he was
collecting in the Concepcion area.

Var. marticorenae is found near the coast in Chile, from
Valparaiso southward to a few km. south of Concepcion.

Field work in South America was supported in part by a
Sigma Xi Grant-In-Aid of Research to the junior author.

439

SECOND EXPEDITION TO NICARAGUA, 1969-1970

Frank C. Seymour

That the flora of Nicaragua is rich in variety is one of the features
‘which attracts botanists to return again and again. A large factor in
providing such variety may be the extremes of rainfall. At one extreme
the rainfall is reported by B. W. Taylor as zero during the month of
February in Managua. At the other extreme is San Juan del Norte, on
the east coast at the mouth of Rio San Juan, where the joking remark
is that it rains all the time. There 823 mm of rain arereported du-
ring July.

Differences of soil must account for some of the variety in flora.
Clay appears to be the most widely distributed soil, but volcanic ash
and even volcanic clinkers are conspicuous types, while large tracts
of land such as those at Waspan and between Siuna and Limbaikan are
characterized by sand. B. W. Taylor has estimated that the number
of species mounts up to 10,000 although the area of the whole country is
smaller than New England, which, with a notably varied flora, supports
only about 2900 species, about .043 per square mile. Nicaragua, with
its 57,915 square miles and 10,000 species, accordingly can boast of
.172 species per square mile. This is in spite of the fact that 3, 475
square miles of its area are covered by water. Thus if the estimate is
not too high, Nicaragua luxuriates in about four times as many species
per square mile than New England.

The personnel of this second expedition consists of only three men,
all of whom participated in the first expedition. The three are: (1)
Frank C. Seymour, the writer of this article, then associate curator
of the Pringle Herbarium of the University of Vermont; (2) John T.
Atwood, Jr., then a graduate student in the University of Vermont,
majoring in botany; (3) Eduardo Narvaez S., thena student in the
Escuela Nacional de Agricultura y Ganaderia in Managua. Members
of the first expedition who were unable to return were deterred mostly
for financial reasons. We have been unable tofind a foundation which

could make us any grant,with the result that each member of our party
had to pay his own expenses.

Acknowledgements. For identifications, we are indebted to Dr.
Henry K. Svenson for Cyperaceae; to Mr. John T. Atwood for Orchi-
daceae; to Dr. Lyman B. Smith and Mr. Atwood for Bromeliaceae;

0

1972 Seymour, Second Expedition to Nicaragua rel

NICARAGUA eee

Botanical Expedition
1969 -1970

Localities of collections
marked by black circles.

Departments
by number

. Boaco

. Carazo

. Chinandega

. Chontales

. Comarco del Cabo

. Esteli 13. Matagalpa

. Granada 14. Nueva Segovia

. Jinotega 15. Rio San Juan COSTA

. Leon 16. Rivas RICA
. Madriz 17. Zelaya

. Managua 18. Lake Managua

2. Masaya 19. Lake Nicaragua Map drawn by Ruth F Benoit

OonNoaourwhs

hy2 P.HY,T O.L:06.1-A Vol. 23, no. 5

to Dr. Harold N. Moldenke for Verbenaceae; to Dr. R. M. Tryon for
_some Pteridophyta; to Dr. Bernice G. Schubert for Desmodium.

The dates of the second expedition, Dec.15, 1969 to Jan. 29, 197Q
were only two weeks later in the season than the first, chosen largely
because it was the time of college vacations when students could come.

On December 15, 1969, Seymour left Burlington, Vermont, by air-
plane, arriving in Managua the next day. For five days he collected a-
lone, until, on December 20, Atwood joined him in a trip to Teustepe.
By December 23, Narvaez joined us. On the trip to Sebaco, Placido Me-
na B., a student in the same Escuela Nacional, collected with the other
three.

We wish to express our appreciation again to Dr. Gustavo Jarqu in
B., then director of the Escuela National de Agricultura y Ganaderia
and his staff for their generous hospitality in welcoming us to their dor-
mitory and making available to us the facilities of their botanical labora-
tory.

A day -by-day record of our collections follows. At the endof a
day's record are given the collection numbers of each person.

Dec. 15. Managua, Route 1, in the region of the Escuala National de

Agricultura y Ganaderia. This is a remarkable area for its rich black
soil and abundant water supply under ground. On this, the first day of
arrival, only a few roadside weeds were taken. Seymour 2333-2340.

Dec. 16. Tipitapa, Route 1, just a few kilometers east of the Escu-
ela and with the same type of habitat, including a shallow muddy pond,
a cotton field and a swampy meadow. Eichornia azurea (Sw.) Kunth
and Tithonia rotundifolia (Miller) Blake were flourishing. Seymour
2341-2347

Dec. 16. Potrerillos, Dept. Boaco, Route 7. A fault line extending
approximately from the Gulf of Fonseca in the northwest to San Juan
del Norte in the southeast, runs about 20 km east of Tipitapa. East of
this fall line, one enters a region of very dry infertile soil where lies
the tiny village of Potrerillos. Altho it is in the Dept. of Boaco, the
soil and vegetation are extraordinarily different from the rain-forests
and lush growth of the eastern part of Dept. Boaco. Collections in
Potrerillos were made along the roadside and in a stomy field. Char-
acteristic species were Amaranthus scariosus Bth. and Jacquinia pun-
gens Gray. Seymour 2348-2371.

Dec. 17. No collecting.

1972 Seymour, Second Expedition to Nicaragua U3

Dec. 18. Managua, near the above-named Escuela, which will be re-
ferred to in the following pages simply as the Escuela. Seymour 2372-
2376.

Dec. 18. Nagarote, Dept. Leon, on the unnumbered route. Altho
this locality was chosen because it was near to Lake Managua ( not
more than 3 km from the western shore), the soil was parched and dry,
consisting of fine volcanic ash. Unlike the region of the Escuela and
Tipitapa, here the streams were low and dried up. Altho in the "Area
Code", it is designated as "Seasonal Tropical Forest", little forest of
any kind was observed. Antigonon leptopus Hk. & Arn. and Tecoma
stans (L.) HBK. were conspicuous. Seymour 2377-2392.

Dec. 19. No collecting.

Dec. 20. Teustepe, only 10 km east of Potrerillos on Route 7, al-
so in the Dept. Boaco. Atwood having arrived, Seymour and Atwood,
collected at first on the south side of the road, where in spite of scrub-
by growth, nevertheless the soil was dry and brooks dried up. On cros-
sing to the north side of the highway, we forded the Malacatoya River.
Here were far more interesting species including Hymenachne amplexi-
caulis (Rudge) Nees, Salix chilensis Molina and Muntingia Calabura L.
The river here is a wide shallow stream with an abundance of water bub-
bling over the many loose stones. Atwood 2392-2422. Seymour 2423 -
2457. Unintentionally numbers 2413-2422 were used twice.

Dec. 21. Managua, casual collections near the Escuela. Seymour
2458-2462.

Dec. 22. Near the Escuela, Atwood 2463, Mangifera indica L.

Dec. 23. Sebaco, Dept. Matagalpa, Route 1. At this point, Rio
Grande is a small stream but with an abundance of water gliding over
a sandy bed. High above its present level, it had left debris in trees
and shrubs to show what it could do in the rainy season. Dividing into
pairs, Narvaez and Seymour explored downstream, Atwood and Mena up-
stream. Most of the vegetation was familiar, but Cyperus canus Presl
in good fruit was unusual. Atwood 2464-2496. Mena 2497-2521. Narvaez
2522-2529. Seymour 2530-2548.

Dec. 24. Casual collections near the Escuela. Atwood 2549 -
2553. Seymour 2554-2555.

Dec. 25. Calabazas, Dept. Matagalpa, Route 1, 80 km north, a
short distance south of Cuidad Dario. Having no apparent means of
celebrating Christmas, keeping busy collecting was the best antidote
for homesickness. For the first time we met a crowd of men, naturally

AAA PY (Ph OuOvG Vol. 23, no. 5

not at work on this holiday. Habitats were,asusual, parched. A tiny
pond, mostly dried up, still retained some soft black muck. Exposed
clay mud had been baked hard.

Seymour having dropped his knapsack onto a low clump of bushes
thought nothing of it until a group of men began showing an extraordinary
interest init. They flourished their machettes and pulled out a snake !
It was a harmless species. This was one of the few times when we saw
any snake, dead or alive. Atwood 2556-2573. Seymour 2574-2603.

Dec. 26. Managua, Route 1. The Escuela being not far from the
shore of Lake Managua at its southern end, Seymour, Atwood, and
Narvaez attempted to hike across country to the shore. After an in-
terminable walk in the broiling sun, still no lake was in sight, so we
gave up the lake and collected a few numbers along lanes and in scrubby
growth. Climbing a tree we brought down a species of Phoradendron.
Nothing appeared more notable than Cyperus panamensis (Clarke) Brit-
ton and Fuirena umbellata Rottb. Atwood 2604-2614. Seymour 2622-
2629, 2594. Narvaez 2615-2621.

Dec. 27. El Viejo, Dept.Chinandega. For our first sampling of the
flora of this department, we collected along a brook near where it emp-
ties into the Atoya River. The vegetation proved to be very similar to
what we had found all along the dry strip of land a few kilometers east
of the Pacific Ocean. Legumes were abundant, among them Desmodium
triflorum (L.) DC. and D. procumbens (Miller) Hitchc. Weare indebt-
ed to Dr. Bernice G. Schubert of the Harvard Herbaria for identifying
the specimens of this genus. Atwood 2630-2646. Narvaez 2647-2656.
Seymour 2657-2678.

Dec. 28. No collecting.

Dec. 29. The breakdown of buses is not always opportune, but it
was this time. In all our travels this is the only time when our bus
broke down. As we entered Dept. Chontales along Route 7, where it
crosses Cano Gordo, at 150-151 km, our bus came to an unexpected
stop. To improve each shining hour, we nearly filled two field presses
here while we waited. Hyptis pectinata Poit. and Borreria suaveolens
Mey. were growing near the river. Narvaez, better than collecting,
helped to persuade the bus to roll again. Atwood 2679-2685. Seymour
2686-2703.

Nicaragua has a very efficient system of locating places along the
highway. At intervals of 1 kilometer, posts give the number of kilomet-
ers distant from the capital, Managua. What could be more efficient to
aid botanists in locating their collections? When near a highway, we
used this means of identifying our locality.

1972 Seymour, Second Expedition to Nicaragua . Ls

A further advantage of the breakdown was that when we arrived in
Santo Tomas, Dept. Chontales, in rain-forest, not only had we already
made collections, but also we were rested again and ready to start anew.
The score for the day in terms of the number of specimens collected
became: Atwood159; Narvaez 124(a small score because of working on
the bus); Seymour 216. Grand total 489.

In Santo Tomas, we sampled a variety of habitats from a low wet
spot to an open hillside anda luxuriant thicket. Panicum stoloniferum
Poiret was clambering, vine-like, over other vegetation, such as Eleo-
charis retroflexa (Poiret) Urban. A small scrawny. tree yielded Catopsis
Berteroana (Schultes f.) Mez and several orchids including Scaphyglottis
cuneata Schl. We are indebted to Dr. Henry K. Svenson for identifying
our Cyperaceae and to John T. Atwood for identifying our Orchidaceae
and many Bromeliaceae; to Dr. LymanB. Smith for identifying many of
our Bromeliaceae. Atwood 2704-2738. Narvaez 2739-2750. Seymour
2751-2804.

Dec. 30. No collecting.

Dec. 31. The regionof the Escuela, although much cultivated, yield-
ed a surprising variety. When time was lacking to go farther afield, we
took a closer look at some promising spot nearby. So, on this date Sey-
mour and Atwood: hiked to a Heliconia swamp over the line in Tipitapa.
Sagittaria lancifolia L. was growing luxuriantly as well as Heliconia
latispatha Bth. Atwood 2805-2825. Seymour 2826-2847.

Jan. 1, 1970. Near the Escuela again, Seymour 2848, Panicum
maximum Jacq.

Jan. 2. Byairplane, we descended upon Siuna, a mining community
in the northern partofDept. Zelaya. With not even a breathing time, we
boarded a truck and reached Limbaikan(Limbaika) on the Prinzapolka
River toward the end of the afternoon. While waiting for a boat to take us
down the river, Atwood grabbed some epiphytes and hid them in the crotch
ofa tree where he found them in fine condition a few days later, onour re-
turn trip. Towhile away the time on the boat, from 6 p.m. until nearly
3 a.m., we studied the stars. On arrivingin PuertoIsabel on the coast
at the mouth of the river, we fellinbed, tooexhausted evento undress.
At Limbaikan, Atwood 2849-2853.

Jan. 3. Puerto Isabel, on the Caribbean Coast at the mouth of the
Prinzapolka River. Collecting within a few feet of the sea proved very
profitable. The land was all verylow, andsofar as we could see, level.
There appeared to be no forest, only scrubby growth and grassland,
obviously pastured. Atwood picked up Rhynchospora nitida(Torr.) Wood
var. hispaniolica Kukenthal, new toCentral America. He and Narvaez

hh6 Pane THO io Gtk Vol. 23, noe

both brought Kyllinga peruviana Rottb., a strikingly characteristic
sedge. Seymour added a second sedge new to Nicaragua, Eleocharis
Sellowiana Kunth. The large number of Cyperaceae in this locality was
very remarkable. Atwood 2854-2881. Narvaez 2882-2893. Seymour
2894-2935.

Jan. 4. Puerto Isabel, again. The second day at this same locality
yielded Cephaelis tomentosa (Aubl.) Vahl and Lipocarpha maculata (Mx.)
Torr. , the latter not found elsewhere. Atwood 2936-2959. Narvaez
2960-2964 . Seymour 2965-2991.

Jan. 4. Leaving Puerto Isabel at six o'clock in the evening, we
traveled all night by boat up the Prinzapolka River, and via Limbaikan
returned to Siuna.

Jan. 5. Siuna. In spite of an almost sleepless night before, we
picked up here some of the plants at which we could only gaze longingly
two days before as we hurried on our way. This is the place where
oranges and grapefruit were lying on the ground in abundance, with no
one to pick them up, delicious as they were. We enjoyed the delightful
hospitality of the La Luz Mining Co. Emilia sonchifolia (L.) DC. was
especially common on a hill in this mountainous region. Eragrostis
pusilla Hackel, apparently new to Central America, was so tiny that
it took a long time to pick enough for a set. Atwood 2992-3003. Nar-
vaez 3004. Seymour 3005-3021.

Jan. 6 was one of our big days. By the kindness of a member of the
mining staff, Mr. Wilbur, we were taken to Mt. Liveco in Madregava.
Tropical ferns such as Polypodium crassifolium L., Vittaria minima
(Bk.) Benedict, and Adiantum macrophyllum Sw. grew along the gentle
trail thru the rain-forest. Olyra latifolia L. was an impressive wide-
leaved grass. Numerous epiphytic orchids graced the branches of the
trees. Prescottia stachyodes Lindley is one orchid which I think we did
not find elsewhere. Hygrophila guianensis Nees at the edge of a small
pool was new to us. We collected eagerly. Atwood 3022-3053; Narvaez
3054-3067; Seymour 3068-3106.

Jan. 6. Having put our morning's collections into press while in
the field, as was our custom, we were able to take still more plants
right near headquarters in the afternoon. The orchid-laden trees and
bushes yielded numerous species such as Notylia bicolor Lindley. In
a grassy ditch grew Coix lacrima-Jobi L. Atwood 3107-3140; Seymour
3141-3162.

Jan. 7. Siuna. Not to miss one minute of our time, we spent the
forenoon hiking along a trail leading from one farm to another. Among
plants not seen before were Oplismenus hirtellus (L.) Beauv., a
species of Smilax, and-Clematis dioica L. Atwood 3163-3179. Narvaez

1972 Seymour, Second Expedition to Nicaragua U7
3180-3185. Seymour 3186-3209, 3284.

Jan. 7. Mt. Liveco, Madregava, again. So great was our success
yesterday that our presses were soon so full that we had to pass by
many choice plants which we wanted. So we returned to this rain-forest.
Scleria latifolia Sw. var. arundinacea (Kunth) is typical of the locality.
Several species of Hyptis were there, including H. pectinata Poit. and
H. pulegioides Pohl. Perhaps the greatest excitement on this occasion
was furnished not by plants. Running back down the trail, where Sey-
mour was collecting, Atwood called, ''We've found a snake! Do you
want to see it?"’ Seymour hurried up the trail after him. By a tree
festooned with epiphytes, we paused. "It's in there'', breathed Atwood.
A monent before, he had been exploring those epiphytes with his hand.
With a long stick, he pointed and poked, while Narvaez poised with staff
raised, ready to strike or run as occasion demanded, and Seymour
stared at the epiphytes with popping eyes. Suddenly a tiny brown snake
dropped to the ground! We could say that we had seen it; that was
enough. It may have been a coral snake, the deadliest of all, but which
has so small a mouth that it cannot bite a man except on his little finger
or the lobe of his ear. We did not wait for positive identification.
Atwood 3210-3239. Narvaez 3240-3247. Seymour 3248-3283, and later,
on return to Siuna 3284.

Jan. 8. Siuna. Ina few minutes snatched before the plane took off,
Seymour 3285-3287.

Jan. 9& 10. Nocollecting. Back in Managua, caring for recent
collections.

Jan.11. Lake Masaya, Dept. Masaya. Climbing Mombacho was
the order of the day, but we could get no nearer than the city of Granada.
Foiled in that attempt, the day was salvaged by revisiting Lake Masaya
where we had botanized a year ago. The soil at lake level is composed
of voleanic clinkers. Lava cliffs rise abruptly at water's edge, encircl-
ing most of the shore as we saw later from an airplane. To this lake
came women to do their laundry. On leaving, they climbed the steep
cliffs with huge bundles of laundry balanced on their heads. In the clink-
ers grew Tetramerium hispidum Nees. On the cliff was Bromus exalta-
tus Bernh. Wigandia caracasana HBK. throve in the clinkers, a species
characteristic of extremely dry soil. Atwood 3288-3300. Narvaez 3301-
3303. Seymour 3304-3323.

Jan. 12. Managua, Escuela. Only Seymour 3324.
Jan. 13. Santo Domingo, Dept. Chontales, in a rain-forest ona

mountain-side. Coccocypselum hispidulum Standley, new to us grew
in abundance along the trail. Most orchids which we have encountered

48 Pay T Oro cra Vol. 23, oe 5

were epiphytes, but here was Habenaria alata Hooker, a terrestrial
species. Of three species of Hyptis, H. recurvata Poiret, had not
been encountered before. Likewise there were two species of Scleria,
S. pterota Presl var. melaleuca (C. & S.) Standley and S. secans (L.)
Urban. To cite an example of the friendliness and hospitality ot the
people of this country,a friend of Narvaez, Sr. Gracias, received us
into his home and made us his overnight guests with very delightful
courtesy. Atwood 3325-3355. Narvaez 3356-3376. Seymour 3377-
3422.

Jan. 14. No collecting.

Jan. 15. Tipitapa, Dept. Managua. Having but a little time left af-
ter retouching day before yesterday's specimens and putting them onto
the drier, Atwood and Seymour, yet again, looked around nearby for
anything which had been overlooked on previous occasions. An Amaryl-
lis grew in a swamp, too fewto enable us to geta set. Panicum molle
Sw. was common. Atwood 3423-3431. Seymour 3432-3455.

Jan. 16. Boaco, Dept. Boaco. Along a city street grew a Lepidium.
This genus has apparently not become well established, as it was not
seen elsewhere (up to that date). Cruciferae, as a family, are scarce
in Nicaragua in strong contrast to New England. Seymour 3456.

Jan. 16. Camoapa, Dept. Boaco. Changing buses at Boaco, we
rode up hill and down dale again and again, instead of following a valley
as roads usually do. As there were no bridges in this very sparsely in-
habited region, we forded streams thirteen times by actual count. Altho
it was evening when we arrived in Camoapa, we improved the remaining
hour of daylight. The sun rises and sets in Nicaragua at about six o'-
clock the year around. With great courtesy, the proprieter of the hotel,
our host, guided Seymour and Atwood to the most likely place for us to
find flowers. All the land which we saw had been thoroughly grazed.
To find specimens which had not already been partially "collected" by
cattle was one of our most persistent problems. Most plants had had
at least one branch bitten off. As cows are not giraffes, above a cer-
tain height, epiphytes still flourished, such as Catopsis nutans (Sw. )
Griseb. and Brassavola nodosa (L.) Lindley, Epidendrum ionophlebium
Rchb. and E. chinense (Lindley) Ames. Atwood 3457-3473. Seymour
3474-3505.

Jan. 17. Camoapa, Dept. Boaco. Next morning, rising at daylight
to accomplish something before the one bus of the day for Managua left,
we returned to about the same spot, but went a little farther. Again the
best specimens were epiphytes undisturbed by cattle; they included Le-
ochilus scriptus (Scheidw.) Rchb. f. and Jacquinella globosaJacq. Cows
do not climb cliffs, so above a tiny cliff were two fleshy species of Pe-

1972 Seymour, Second Expedition to Nicaragua hig
peromia, still unidentified. Atwood 3506-3521. Seymour 3522-3540.

Jan. 18. No collecting.

Jan 19. Waspan, Comarca del Cabo, one of the northernmost com-
munities in Nicaragua, is separated from Honduras only by the Coco
River. Across the river, rice is extensively cultivated. A precipitous
clay bank drops from the very level land of the town down to the river
below, where it is bordered by very low meadows. Most of our collec-
tions along the river upstream from the town were very ordinary except
Drosera capillaris Poiret which we found rarely. Salvinia in abundance
was floating on a small pool. Dianthera comata L. grew in the swamp.
Only Atwood and Seymour came to Waspan. Atwood 3541-3563. Sey-
mour 3564-3595a.

Dr. Robert M. Hooker, a life-time resident of Bluefields informed
us that this name should be spelled Waspan because in the early days
gold-seekers washed their findings here in pans.

Jan. 20. Waspan, still. Within easy hiking distance was a new type
of habitat which we had only glimpsed before, between Siuna and Limbai-
kan. The pines here, Pinus caribaea Morelet, do not grow close to-
gether like some other species of pine, but are scattered. Later we
were informed that if the land were not burned over frequently, they
would grow close together. The ground was covered with standing wa-
ter, drained a little by a tiny sluggish stream. Sedges were abundant,
such as Eleocharis mitrata (Griseb.) Clarke, new to Central America.
Rhynchospora flourished in four species, namely, R. rariflora (Mx. )
Ell. , new to Central America; R. hirsuta Vahl; R. globosa (HBK.) R. &
S.; R. divergens Chapman, identification somewhat doubtful, but if cor-
rect, a species previously known in Central America only in British
Honduras; also R. barbata (Vahl) Kunth, and R. setacea(Berg.) Btckl.
Burmannia sp., an Eriocaulon and a Xyris; two species of Utricularia;
Hyptis lantanaefolia Poit. and H. conferta Pohl. There were other un-
usual species too numerous to mention. Atwood 3596-3649. Seymour
3650-3706.

Jan. 21. Cororia Bush, 35 km southwest of Waspan. This locality
was sometimes misspelled on our labels; it is correct as given here.
After a long ride by jeep, we reached a point where the jeep could go
no farther and the vegetation made an abrupt change. The pine-savan-
nas ended abruptly to be replaced westward by rainforest. Here was
a geological fault along which ran Kisalaya Creek. The rain-forest was
dense, Far overhead , tree-tops basked in the sunlight but on the ground
daylight was dim; shade was so deep that few plants could survive. On-
ly along the borders of the forest was there much to collect. Scleria
mitis Berg. had not been found before. Two species of Cephaelis, C.

450 PHYTOLOGIA Vol. 23, no. 5

glomerata D.Sm. and C. tomentosa (Aubl.) Vahl, grew not far apart.
This spot yielded another Rhynchospora, R. rugosa (Vahl) Gale, and
Dichronema Watsonii Britt. and D. radicans C. & S. anda scrap of
Scleria bracteata Cav. In the time available we could nowhere near do
justice to this extremely rich and productive area. Atwood 3707- 374 5.
Seymour. 3746-3811.

Jan. 22. Waspan. Exhausted tho we were from the previous day,
Atwood spurred himself to rise early in the morning and gather ina few
more specimens before we had to board the plane to return to Managua.
These were collected in pine-savannas near the town. Atwood 3812-3817.

Jan. 23. No collecting.

Jan. 24. Escuela. While waiting for a bus, Atwood found Asclepias
sp. which we had not found elsewhere. It became more common later
in the season. Atwood 3818.

Jan. 24. Boaco, Dept. Boaco. The good places which we drove
past in Boaco a year ago without a chance to sample them lured us back
on this date for what we feared would be our last trip this year. Climb -
ing a mountain south of the city, we picked a few straggling specimens,
at first meeting nothing but heavily grazed slopes. On our squeezing
thru a barbed wire fence, everything was changed; we were in a rain-
forest near the summit, where the land had not been grazed. Trees
were loaded with orchids such as Epidendrum Schlechterianum Ames
and E. masayensis. One of the memorable discoveries was a fragrant
Adiantum, A. trapeziforme L. Cattleya Skinneri Bateman, Atwood re-
cognized at sight from living flowering plants in his home in Vermont.
Umbelliferae as a rule are scarce in areas where we collected, but
here was a species of Eryngium. The most interesting species are as
yet unidentified. A flock of monkeys were frolicking in the tree-tops
far above. Narvaez did not come on this trip. Atwood 3819-3836.
Seymour 3837-3879.

Jan. 25-26. By this time we were desparate to get our specimens
tied up in bundles and packed for shipping back to Burlington. There-
fore there was almost no collecting on these two days.

Jan. 25. Managua, the Escuela. Seymour 3896, Pennisetum ci-
liare (L.) Link.

Jan 26. Managua, the Escuela. Seymour 3897, Panicum hirticaule
Presl.

Jan 27. Managua, the Escuela. Seymour 3898. Anthephora herma-
phrodita (L.) Kuntze.

1972 Seymour, Second Expedition to Nicaragua us.

Jan. 27. Mombacho, Dept. Granada. To the others, packing
seemed well enough along to warrant one more trip, so Seymour con-
tinued packing while Atwood and Narvaez made a second attempt to
climb this mountain, one of the highest extinct volcanic cones in the
country, which we already knew to be rich in tropical vegetation.
Reaching the mountain was not too difficult, but time was lacking to
attain the summit. Orchids from this mountain included OncidiumCe-
bolleta(Jacq.) Sw. Ferns were plentiful. among them Polypodium plebe Tebe—
jum C.&S. and P. dissimile L. For the return, no vehicle could be
commandeered.With heavily loaded presses, the two men, already very
tired, had to hike six miles to Granada and arrived at headquarters
in the Escuela by bus too tired to speak. Atwood 3899-3924. Narvaez
3880-3895 and 3925-3927.

Jan. 28-29. Managua, the Escuela. Just to mark the fact that we
were still in Nicaragua on these dates, Seymour 3929, Atwood 3930-
3931, including Oncidium ensatum Lindley.

Nos. 3932-3945 have been assigned to orchids which flowered in
Vermont after our return from Nicaragua.

Our first expedition, almost exactly one year earlier was spent
largely in the thickly populated dry strip of land near the Pacific Ocean.
Our special objective on this second expedition was to explore the eas-
tern part of the country which was characterized by rain-forest. Ex-
cept for short sallies to make use of odd bits of time when time did
not permit us to go far, nearly all of the field work was in rainforests.
Nagarote was one of the few exceptions, selected because we had spent
very little time a year ago in Dept. Leon. The other exception is the
trip to Chinandega, undertaken because a year ago we did not botanize
at all in Dept. Chinandega. Now we have explored to a slight extent
every department in the country except Cabo de Gracias a Dios and
have spent the largest proportion of our efforts in that most fasci -
nating and largest department, Zelaya.

The Pringle Herbarium
The University of Vermont
Burlington, Vermont 05401

Reference: Taylor, B. W. An Outline of the Vegetation of Nicaragua.
Journal of Ecology 51:27-54. 1963.

Pohlia annotina (Hedw.) Lindb. var. loeskei Crum,
Steere and Anderson in Guatemala

Dana Griffin, III
Bryophyte and Lichen Herbarium
University of Florida

Bartram (1949) called attention to several genera and species
of northern mosses with disjunct populations in Guatemala. These
forms are found at the upper elevations and are thought to repre-
sent the result of a migration southward by populations into
Mexico and Central America in advance of the glaciers with subse-
quent isolation at the higher elevations upon retreat of the ice.

To this list of disjuncts may now be added the distinctive
taxon, Pohlia annotina (Hedw.) Lindb. var. loeskei Crum, Steere
and Anderson. To my knowledge this is the first report of this
species south of lat. 34° N.

Crum and Miller (1967) summarized the known distribution of
the variety. The plants are fairly common in Europe ranging from
the Iberian Peninsula northward to Greenland and Siberia and
eastward to Asia Minor. In North America they are found from
Alaska to Maine thence southward to Georgia.

Collection data for the Guatemalan plants are as follows:

Pohlia annotina (Hedw.) Lindb. var. loeskei Crum, Steere

and Anderson. GUATEMALA: DEPT. OF JALAPA. Ten kilometers

east of Mataquescuntla along highway 18. Plants on soilbank

along roadside. Elevation 2600 m. leg. F. Dixon. 5 Feb. 1971.

Coll. No. 014323

The Guatemalan plants are indistinguishable from those I have
seen from Europe, including the production of numerous subglobose
propagula in the leaf axils. The gametophytes average 4-5 mm
high. Setae are 3-4 cm long with the capsules ranging in length
from 2.8-3 mm. The neck of the capsule is distinct and slightly
bulging-mamillose. The urn is smooth.

I am indebted to Dr. A.J. Sharp, Department of Botany, Uni-
versity of Tennessee, who has examined this collection and concurs
in the identification.

Literature Cited

Bartram, E.B. 1949. Mosses of Guatemala. Fieldiana 25:1-442.
Crum, Howard and Norton G. Miller. 1967. Three propaguliferous
Pohlias from Michigan. The Bryol. 70(1):118-119.

52

SEVEN UNDESCRIBED SUBSPECIFIC TAXA IN THE ERIOCAULACEAE AND
VERBENACEAE

Harold N. Moldenke

CARYOPTERIS INCANA f. MACROPIIYLLA Moldenke, f. nov.

Haec forma a forma typica speciei foliis plerumque usque ad
11.5 cm. longis )}.5—5S cm. latis recedit.

This form differs from the typical form of the species in hav-
ing the main leaves on flowering branches with petioles 2.5--3 cm.
long and leaf=blades 8.5--9 cm. long and .5-—-5 cm. wide.

The type of the form was collected by an unknown collector
somewhere in Japan on September 20, 1910, and is sheet mmber
1178283 in the United States National Herbarium at Washington.

CARYOPTERIS INCANA var. SZECHUANENSIS Moldenke, var. nov,

Haec varietas a forma typica speciei petiolis uniforme ca.
mm. longis et laminis foliorum uniforme 2.5--3 cm. longis 11.5
cm. latis regulariter grosse antrorse incisis et reticulo venarum
venularumque supra arcte impresso recedit.

This variety differs from the typical form of the species in
having its branches uniformly foliose, the internodes about 2 cm.
long, the petioles uniformly about ) mm. long, and the leaf-blades
remarkably uniform, lanceolate-ovate, 2.5--3 cm. long, 1—1.5 cm.
wide, regularly incised-dentate along practically the whole margins,
the incisions antrorse, 2--3 mm. long, obliquely broad-based,
bluntly subacute, revolute-margined, and the very abundant and
fine vein- and veinlet-reticulation conspicuously and beautifully
impressed on the upper surface.

The type of this variety was collected by Ernest Henry Wilson
(no. 2221) somewhere in western Szechuan, China, in August, 1908,
and is 's deposited in the United States National Herbariun at
Washington,

CARYOPTERIS ODORATA var. INTEGRIFOLIA Moldenke, var. nov.

Haec varietas a forma typica speciei recedit foliis parvioribus
plerumque 3--);.5 cm. longis 1—2 cm. latis integris subtus densi-
ore persistenteque pubescentibus et inflorescentiis in axillis
superioribus terminalibusque dense congestis.

This variety differs from the typical form of the species in
having its leaves smaller, mostly only 3--.5 cm. long and 1—2 cn.
wide during anthesis, usually entire-margined, and mostly more
densely and persistently pubescent beneath and the inflorescence
densely congested in all the upper leaf-axils as well as terminal
on stems and branches.

The type of the variety was collected by Walter Koelz (no. 399)
at Bhadwar, Kangra, Punjab, India, at an altitude of 2000 feet, on
May 5, 1933, and is deposited in the United States National Herbar-
ium at Washington.

453

45h PHY £0100 2a Vol. 23, no. 5

CORNUTIA PYRAMIDATA var. ISTHMICA f. ALBIDA Moldenke, f. nov.

Haec forma a forma typica varietatis corollis albidis et lan-.
inis foliorum ultra medium serrulatis recedit.

This form differs from the typical form of the variety in hav-
ing whitish corollas and the leaf-blades serrulate above the
middle.

The type of the form was collected by Rolando Tun Ortiz (no.
2106) in a high forest at Santa Elena on the west side of the _
highway to Dolores at km. 70, El Petén, Guatemala, on November 20,
1971, and is deposited in the Britton Herbarium at the New York
Botanical Garden. The collector describes the plant as a shrub
3m. tall, the trunk 20 cm. in diameter, and the corollas "blanca
amarillenta",

DURANTA MUTISII f. SERRULATA Moldenke, f. nov.

Haec forma a forma typica speciei laminis foliorum margine
minute obscureque serrulatis recedit.

This form differs from the typical form of the species in
having the margins of its leaf-blades mimutely and obscurely
serrulate.

The type of the form was collected by Antonio Allart (no. 325)
at Colonia Tovar, Aragua, Venezuela, at an altitude of 1800 to
2000 meters, in December, 192), and. is deposited in the Britton
Herbarium at the New York Botanical Garden.

LANTANA MINASENSIS var. HISPIDA Moldenke, var. nov.

Haec varietas a forma typica speciei ramulis plusmimus dense
albido-hispidis recedit.

This variety differs from the typical form of the species in
having the branchlets more or less densely hispidulous with long,
white, often capitate hairs standing at right angles to the
otherwise densely short-pubescent branchlets.

The type of the variety was collected by A. Krapovickas, C. L.
Cristébal, M. M. Arbo, V. Maruflak, R. I. Marufiak, and J. Irigoyen
(no, 17030) on the high part of the barranca of the Rfo Uruguay,
at the edge of a woods, Colonia Garabi, Dept. Santo Tomé, Corri-
entes, Argentina, on December 3, 1970, and is deposited in my per-
sonal "herbarium at Plainfield, New Jersey. The collectors de-
scribe the plant as a shrub 1.8 m. tall.

LEIOTHRIX HIRSUTA f. VIVIPARA Moldenke, f. nov.

Haec forma a forma typica speciei capitulis viviparo-proliferis
recedit.

This form differs from the typical form of the species in hav-
ing some or most of its flower-heads viviparously proliferating
into clusters of miniature leaves.

The type of the form was collected by H. Strang and A. Castel-
lanos (no. 26306) at Pedra Itatina "na rodoc. BR-6", Restinga de
Sernambetiba, Guanabara, Brazil, on March 12, 1967, and is depos-
ited in my personal herbarium at Plainfield, ’ New Jersey.

VARIEGATA ONCIDIUMS
WeW.G. Moir

In Phytologia Vol. 15. No. 1 the author and A.D. Hawkes
had an article on "Studies in the equitant oncidiums". In
Vol. 17. No. 6 of the same publication the author had two
more articles. In the article in Vol. 15 page 10 and 11 is
described a var. majus to Onc. leiboldii. This has proven to
be a natural hybrid between O Onc. leiboldii Rchb.f. and Onc.
variegatum (SW) SW. and in this article it is named Onc.

x cubense, for it arrived in Hawaii before 1950 as a plant
from Cuba without place of origin.

In the American Orchid Soc. bull. 36:220, 1967 Withner
described Onc. osmentii. Just as in the case of Onc. x cu-
bense I have proved by actual breeding that this plant is a
natural hybrid between Onc. haitiense Leonard & Ames arid Onc.
quadrilobum C. Schweinfurth so this name will be recorded
henceforth as Onc. x osmentii (Withner) Moir 1972.

Mr. William Osment of 2435 Cleveland St., Hollywood,
Florida has, in the last couple of years, found some interest-
ing new plants while studying these Variegata oncidiums in
the Dominican Republic. He found, along with Miss Ann Hadder,
a new natural hybrid between Onc. variegatum (SW.) SW. and Onc.
haitiense Leonard & Ames at Sabana Alta in the western part
of the Republic. This is described as Onc. x ann-hadderiae.
In the northwest part of the Republic, Osment found a form of
Onc. desertorum that has a red lip and yellow sepals and
petals. This is described herein as Onc. desertorum var.
aureorubrum. This could possibly be a natural hybrid between
Onc. desertorum and Onc. desertorum var. alborubrun.

In Haiti in 1955 the author collected plants of Onc.
velutinum (Ldl.) when not in flower. When flowering these in
Hawaii one plant turned out to be different. It looked like
a hybrid of Onc. velutinum (Ldl.) and Onc. variegatum (SW.)
SW. A year ago the man made hybrid of these two species was
flowered. It turned out to be identical to the plant from
Haiti and so the Haiti plant has been named Onc. x varvelun.
The all purple form of Onc. velutinum is to be found in both
the D.R. and Haiti on the southwestern portion of Hispaniola.
The flower is larger and its crest has three large horns in-
stead of 5 of the species. This is now designated var. pur-

pureum.

A book has been published in 1970 by the author and his
wife, May A. Moir, on "Variegata Oncidiums" and pictures of
them given along with the prominent points of their character-
istics as to location, growth, flowers, environment, and

455

456 PHY T O.L0'G DA Vol, 23, no. 5

further comments. With those described here there now
totals 30 species, 8 varieties and 11 natural hybrids. Onc.
ibertianum (A.Rich) has proved to be Onc. lemonianum
(Lindl.) so the latter name should now be used. Onc. ten-
ellum listed by Garay on page 450 of "A Reappraisal of the
Genus Oncidium SW." is not to be found in Guyana and its

description reads like Onc. variegatum (SW.) SW. and is not
counted among the 30 species.

There still exists in the Bahamas three types of plants
that have different colors of flowers that are different to
Onc. lucayanum Nash, Onc. bahamense Nash, Onc. sylvestre
Ldl. and Onc. urophyllum Lodd. Plants with purple and white
flowers are three times larger than Onc. lucayanum and have
up to 50 flowers per spike. The yellow flowered form and
the orange red form are plants of intermediate size and have
well spaced flowers but fewer. A form of urophyllum was
found in Hleuthera, Bahamas by John Hall III but it has much
longer rhizomes than the type form in Antigua. The form of
Onc. bahamense Nash (not Onc. variegatum (SW.) SW.) in west
Palm Beach area in Florida is also much more vigorous than
the form from Andros, Bahamas, even after 15 years of grow-
ing side by side in Hawaii.

Many years ago I went to the Dominican Republic and
while dollecting in the area east of Santiago Rodriguez
some plants were collected not in flower. Later one bloomed
that was different to any I had seen before. Later Osment
sent me in 1971 a plant that has flowered in May 1972 that
has the same flower. I thought these had Onc. variegatum in
them but on closer examination it is not so. I know how the
hybrid would be if it was a natural cross of Onc. variegatum
with the yellow flowered species of the area: desertorum,
quadrilobum and haitiense so I have described it as a new
species named it after Dr. Jose de J. Jimenez of Santiago de
los Caballeros, a medical doctor with a hotoy of botany and
author of Supplement No. 1 of Moscosco's Flora Domingensis..

Herbarium specimens of the new additions will be sent
to Ames Herbarium, Harvard University, Cambridge, Mass.
For Onc. x cubense, a new label for the specimen Onc. lei-
boldii var. majus will be sent instead of a new specimen.
All specimens have been flowered in Hawaii and checked with
other species. Natural hybrids have been checked with man
made plants of the same crosses. No specimen will be sent
of Onc. velutinum var. purpureum but a photograph will
suffice.

1972 Moir, Variegata Oncidiums 457
ONCIDIUM JIMENEZII, Moir, sp. nov.

Planta grandis. Pseudobulbi nulli. Folia ad 5 sesilis,
lanceolata, acuta, supra profunde canaliculata, crasse cor-
iacea, margine minute denticulata, ad 6 cm. longa, ad 1 cn.
latas virdis. Scapus racemosa, post paniculata, pauci- ad
pluriflora, 15 ad 25 cm. longa; bractae lanceolata, tenui-
membranaceae, ad 1 cm. longa; pedioelli cum ovariis 2 cm.
longi. Sepalum dorsale spathulatum, concavum, rubiginosus;
laterali synsepalum post labellum, linearis, 2 apicem; 1 cn.
longum, 2 mm. latum. Petala 1.2 cm. longa, & mm. lata, ob-
tusa, prima rubiginosa, citrina cum rubiginosa maculata.
Labelli lobi laterales 8 mm. longa, 4 mm. lata, oblongive,
apice rotundati; isthmi augustis; lobus intermedius reni-
formis, apice ungis, laviter crenulatus, 2 cm. latus, totum
labellum ad 1.5 cm. longum, omnia luteas cristi labelli
biseriata, tubercula 3 in serie, toti 6, lutei. Gynostemii
alae magnae, ovalis, acutae, citrinae. Fl. a maius.

Plant large, no pseudobulb. Leaves to 5, lanceolate,
pointed, above deeply grooved, fat and coarse, minute teeth
on margin, to 6 cm. long, to 1 cm. broad. Scape raceme,
later panicle, few to many flowered, 15 to 25 cm. longs
bracts lanceloate, membraneous, to 1 cm. long; pedicle plus
ovary 2 cm. long. Dorsal sepal spathulate, concave, brown
red; laterals synsepal behind labellum, linear with 2 points,
1 cm. long, 2 mm. broad, spotted with brown red. Petal 1.2
cm. long, 8 mm. broad, obtuse, first brown red then pale
yellow with brown red spots near edge. Labellum side lobes
7 mm. long, 4 mm. broad, oblong, rounded end; narrow is-—
thmus; middle lobe reniform, full, apice clawed, large
indent, edge crenulate, 2 cm. broad, yellow, total labellum
to 1.5 cm. long; crest in two series of 3 each, 6 in total,
forward pointing, upper outer ones slightly grooved.

Wings on column large, bottom rounded, upper end pointed.
Flowers in May.

Dominican Republic east of Santiago Rodriguez in shrub-
bery. Collected by Moir in 1955. Kecollected by William
Osment in 1969.

ONC. VELUTINUM var. PURPUREUM Moir, var. nov.

Differt a forma typica flora amplus. Flora color purpura;
cristi 3 partis in vicam 5.

Plant same as species but flower larger and of purple
color with darker purple colors near crest, upper lobes and
at base of sepals and petals, crest, upper lobes and at base
of sepals and petals. Crest has 3 instead of 5 horns that is
the two, one on each side of the large horn of the lower sec-
tion of the crest, are not present so makes lower, large horn
more prominent.

458 PHYTOLOGIA Vol. 23, no. 5

Hispaniola SW. Collected at roadside on citrus trees
along with species and n.h. Onc. x varvelum 1955. Above
Petionville, Haiti.

ONCIDIUM DESERTORUM var. AUREORUBRUM, Moir var. nov.

Planta similis var. alborubrum; petala et sepala lutea
in viceum album.

Strong plant, much greener than type. Sepals and
petals deep yellow and red lip, so very much like var.
alborubrum except for sepals and petals color.

Dominican Republic — near Santiago Rodriguez.
Collected by William Osment. Flowered in Hawaii.

ONCIDIUM x ANN-—HADDERIAE, Moir, nat. hybr. nov.

(Onc. variegatum (SW.) SW. x Onc. haitiense Leonard
and Ames) Habitu inter parentiis intermedium. Flora magna.

Pseudobulb not visable, growth compact to slightly
rhizomatous. Leaves 4 cm. long and 1 cm. broad, olive
green, dull point, upper edge grooved, 5 to 6 leaves.

Scape raceme then later branched. Dorsal sepal 6 mm.

long, 2 mm. broad, concave, brown bars on yellow; lateral
sepals joined to near end, concave, brown, 1 cm. long, 4 mm.
broad. Petals 1.2 cm. long by 5 mm. wide, brown bars part
way on yellow, pointed petals turned upward and slightly
reflexed. Lip 1.7 cm. long 1.9 cm. broad, full, reniform
lower lobe, light yellow, slightly undulating edges; lateral
lobes 5 mm. long by 2 mm wide, turned up. crest large, 6
parts 5 nearly equal in size and all but the middle one of
the lower three are turned down, but the middle one of the
upper three is a pearlike projection. Wings on column
large, rounded base and pointed top. Flowers in May.

Dominican Republic, Sabana Alta near San Juan collect—
ed by Ann Hadder and William Osment of Hollywood, Florida.
Flowered in Hollywood, Florida and Honolulu.

ONCIDIUM x CUBENSE Moir, not. hybr. nov.
(Onc. variegatum (SW.) SW. x Onc. leiboldii Rchb.f.)

Habitu inter parentiis intermedium, excepto folii
longiore. kKhizoma brevis,

Pseudbulb non visigble, plant on 3 to 4 cm. rhizome,
numerous roots. Leaves 6 to 7 cm. long. 1 cm. broad,
upper side longer and curved to a sharp point, large groove.
Scape short and many flowered, bracts small and pointed.
Dorsal sepal 1 cm. long by 3 mm. broad, brown bars on white,

1972 Moir, Variegata Oncidiums 459

concave, erect; lateral sepals joined, brown, concave, two
pointed. Petals 1.2 cm. by 5 mm., undulated, white, with
dark brown bars to 3/4 of way from base. Lip 2 cm. x 2 cm.
white with dark brown bars and spots to mask around crest;
lateral lobes medium sized turned up and rounded; isthmus
about 5 mm3 lower lobe reniform and full. Wings on column
large like variegatum. Stronger grower than either parent
and floriferous.

Cuba, exact spot unknown. kKeceived in Honolulu prior
to 1950. Flowered in Honolulu. Specimen already at Ames
Herbarium as Onc. leiboldii var. majus. Proved by making
the cross.

ONCIDUM x VARVELUM Moir, nat. hybr. nov.
(Onc. variegatum (SW.) SW. x Onc. velutinum Ldl.)

Habitu inter parentiis intermediun.

Pseudobulbs not visible. Growth rhizomatous as both
parents but more lax compared to velutinum, rhizome 10 cm.,
wiry. Leaves to 5 or 6 in number curved and grooved on
upper edge, green, serrated lightly. Scape a raceme at
first then branched later, long, floriferous. Dorsal sepal
8 mm. long x 2 mm. broad, light lavender with brown bars,
concave; lateral sepals jointed together but 2 points, concave,
brown. Petals 1 cm. long x 7 mm. broad at middle, with brown
bars part way, white to pale lavender, and undulated, dull
point, turned inward and upward. Lip 1.7 cm. long to 1.8 mn.
turned up and rounded ends as in velutinum; mask around
crest brown to dotted with brown; crest 6 parts, upper two
large and straight sideways with groove, little pearl like
projection in between large ones, lower part of crest has 1
large part turned up and one little one on each side. Wings
on column are large, lavender, lower end rounded, upper end
pointed.

Haiti about 2500 ft. elevation on citrus trees at road
side, above Petionville. Flowered in Honolulu.

ADDITIONAL NOTES ON THE GENUS VERBENA. XIV

Harold N, Moldenke

VERBENA OFFICINALIS L.

Additional bibliography: Moldenke, Phytologia 23: 377--389.
1972.

Coon (1963) tells us also that "Pliny said 'if the dining cham-
ber be sprinkled in water in which the herb Verbena has been
steeped, the guests will be merrier'. Such a story led to a be-
lief in its efficacy as a cure against the plague, and as a remedy
for almost anything. It even had supposed supernatural powers.
Several Welsh names have meanings such as ‘devil's hate' and
'enchantment herb! ,"

Fior (1969) reports that "Gli antichi attribuirono a questa p.
proprieta meravigliose, la impiegarono per pulire gli altari
prima dei sacrifici e par intessere corone agli eroi ai poeti."

He affirms that the name "verbena" was used by Virgil, which is
correct.

Bertolini (18), as is to be expected, concentrates on the
Roman history of the plant: "Olim laudata apud medicos ob vim
adstringentem, vulnerariam, munc exoleta. Veteres superstitiose
Verbenas adhibebant: 'Siquidem auctores imperii Romani, conditor-
esque immensum quiddam et hinc sumpsere, quoniam non aliunde
sagmina in remediis publicis fuere, et in sacris, legationibusque
Verbenae. Certe utroque nomine idem significatur, hoc est gramen
ex arce cum sua terra evulsam, ac semper et legati cum ad hostes,
clarigatumque mitterentur, idest res raptas clare repetitum, umus
utique Verbenarius vocabatur.' Plin. Nat. hist. lib. 22, cap. 2.
[S8--79 A.D.] Pari ratione utebantur Verbenis in sacris magicis:

"Effer aquam, et molli cinge haec altaria vitta,
Verbenasque adole pingues, et mascula thura
Virg. Eccl. 8. v. 64.65 [37 B.C.]
Sacerdotes eisdem coronabantur:
" eeeeeeeeeealii fontem, ignemque ferebant
Velati lino, et verbena tempore cincti
Virg. AEneid. lib. 12, v. 119.120 [19 B.C.]
Inprimis vero aediculae compitales verbenis ornabantur:
"Flore sacella tego, verbenis compita velo,
Et crepat ad veteres herba sabina focos
Prop. Eleg. lib. h. eleg. 3. v. 57.58." [26 B.C.]

Folkard (188) gives another full account of this fascinating
plant's history, saying "The Vervain, or Verbena, has from time in-
memorial been the symbol of enchantment, and the most ancient na-
tions employed this plant in their divinations, sacrificial and
other rites, and in incantations. It bore the names of Tears of
Isis, Tears of Juno, Mercury's Blood, Persephonion, Demetria, and
Cerealis. The Magi of the ancient Elamites or Persians made great
use of the Vervain in the worship of the Sun, always carrying
branches of it in their hands when they approached the altar. The

460

1972 Moldenke, Notes on Verbena 4,61

magicians also employed the mystic herb in their pretended divin,
ations, and affirmed that, by smearing the body over with the
juice of this plant, the person would obtain whatever he set his
heart upon, and be able to reconcile the most inveterate enemies,
make friends with whom he pleased, and gain the affection, and
cure the disease of whom he listed. When they cut Vervain, it
was always at a time when both the sun and moon were invisible,
and they poured honey and honeycomb on the earth, as an atonement
for robbing it of so precious a herb. -- The Greeks called it the
Sacred Herb, and it was with this plant only that they cleansed
the festival-table of Jupiter before any great solemnity took
place; and hence, according to Pliny, the name of Verbena is de-
rived. It was, also, one of the plants which was dedicated to
Vemms. Venus Victrix wore a crown of Myrtle interwoven with Ver-
vain. -- With the Romans, the Vervain was a plant of good omen,
and considered strictly sacred: —

'Bring your garlands, and with reverence place

The Vervain on the altar.'
They employed it in their religious rites, swept their temples
and cleansed their altars with it, and sprinkled holy water with
its branches. They also purified their houses with it, to keep
off evil spirits; and in order to make themselves invulnerable,
they carried about their persons a blade of Grass and some Ver-
vain. Their ambassadors, or heralds-at-arms, wore crowns of Ver-
vain when they went to offer terms of reconciliation, or to give
defiance to their enemies, a custom thus noted by Drayton: -—-

'A wreath of Vervain heralds wear,

Amongst our garlands named;
Being sent that dreadful news to bear,
Offensive war proclaimed.'

Virgil mentions Vervain as one of the charms used by an enchant—
ress: —

"Bring running water, bind those altars round

With fillets, and with Vervain strew the ground.'
The Druids, both in Gaul and in Britain, regarded the Vervain
with the same veneration as the Hindus do the Kusa or Tulasi, and,
like the Magi of the East, they offered sacrifices to the earth
before they cut this plant. This ceremony took place in Spring,
at about the rising of the Great Dog Star, but so that neither
sun nor moon would be at that time above the earth to see the
sacred herb cut. It was to be dug up with an iron instrument, and
to be waved aloft in the air, the left hand only being used. It
was also ordained by the Druidical priests, for those who collected
it, 'that before they take up the herb, they bestow upon the ground
where it groweth honsy with the combs, in token of satisfaction and
amends for the wrong and violence done in depriving her of so holy
a herb. The leaves, stalks, and flowers were dried separately in
the shade, and were used for the bites of serpents infused in wine.
Another account states that the Druidesses held Vervain in as
great veneration as the Druids did the Mistletoe. They were never
permitted to touch it. It was to be gathered at midnight, at the
full of the moon, in this manner: -- a long string with a loop in

62 P BE O brO Gch Vol. 23, noses

it was thrown over the Vervain-plant, and the other end fastened .
to the left great toe of a young virgin, who was then to drag at
it till she had uprooted it. The eldest Druidess then received
it in a cloth, and carried it home, to use it for medicinal pur-
poses and offerings to their gods. In the Druidic procession, to
the gathering of the Mistletoe, the white-clad herald carried a
branch of Vervain in his hand, encircled by two serpents. The
priests, when performing their daily functions of feeding the
never—dying fires in the Druidic temples, prayed for the space of
an hour, holding branches of Vervain in their hands. Pliny tells
us that the Druids made use of it in casting lots, as well as in
drawing omens and in other pretended magical arts; he also says
that if the hall or dining chamber be sprinkled with water where-
in Vervain lay steeped, all that sat at the table should be 'very
pleasant and make merry more jocundly'.
'Lift up your boughs of Vervain blue,
Dipt in cold September dew;
And dash the moisture, chaste and clear,
O'er the ground and through the air.' -- Mason
"In mediaeval days, the sacred character of Vervain was still
maintained, and the plant was greatly prized, and used in com
pounding charms and love-philtres. Known in our country as Holy
Herb and Simpler's Joy, it was credited with great medicinal
virtues.
'Black melancholy rusts, that fed despair
Through wounds' long rage, with sprinkled Vervain cleared.!
Davenant.
Its juice was given as a cure for the plague, and the plant was
prescribed as a remedy in some thirty different maladies, and was
suspended round the neck as an amulet. Gerarde, however, tells
us that 'the devil did reveal it as a secret and divine medicine';
and R. Turner writes (1687): -- 'It is said to be used by witches
to do mischief, and so may all other herbs if by wicked astrolo-
gers used to accomplish their wretched ends'. But notwithstanding
that it was used by witches and wizards in their incantations and
spells, and was in fact called the Enchanter's Plant, Vervain was
considered to possess the power of combating witches; thus Aubrey
says: —
! ee0eeee0Vervain and Dill
Hinder witches from their will.!
and Michael Drayton writes: —
"Here holy Vervayne, and here Dill,
'Gainst witchcraft much avayling.!
and again —
'The Nightshade strews to work him ill,
Therewith the Vervain and the Dill
That hindreth witches of their will.'
On the Even of St. John (June 23rd), Vervain was for a long time
associated with the observances of Midsummer Eve. Thus we read
in 'Ye Popish Kingdome': —
'Then doth ye joyfull feast of John ye Baptist take his turne

1972 Moldenke, Notes on Verbena 463

When bonfires great with loftie flame in every towne doe burne,

And young men round about with maides doe dance in every

streete,

With garlands wrought of Mother-wort, or else with Vervain

sweete.'
J. White, Minister of God's Word, writes in 162): — 'Many also
use to weare Vervein against blasts; and when they gather it for
this purpose firste they crosse the herb with their hand, and
then they blesse it thus: --

'Hallowed be thou, Vervein,

As thou growest on the ground,

For on the Mount of Calvary

There thou wast first found

Thou healedst our Saviour Jesus Christ,

And staunchedst his bleeding wound,

In the name of the Father, the Son, and the Holy Ghost,

I take thee from the ground.'
In many rural districts, Vervain is still regarded as a plant
possessing magical virtues as a love philtre. It has the reputa-
tion of securing affection from those who take it to those who
administer it. The gun-flint boiled in Vervain and Rue ensures
the shot taking effect. The root of Vervain tied with a white
satin ribbon round the neck acts as a charm against ague. Ver-
vain and baked toads, worn in silken bags around the neck, are a
cure for the evil. — In the northern provinces of France, the
peasants still contime to gather Vervain under the different
phases of the moon, using certain mysterious ejaculations know
only to themselves whilst in the act of collecting the mystic
herb, by whose assistant they hope to effect cures, and charm
both the flocks and the rustic beauties of the village. — The
Germans present a hat of Vervain to the newly-married bride, as
though placing her under the protection of Venus Victrix, the
patroness of the plant. -- Gerarde tells us that in his time it
was called 'Holie Herbe, Juno's Teares, Mercurie's Moist Blood,
and Pigeon's Grass, or Columbine, because pigeons. are delighted
to be amongst it, as also to eate thereof.' -- Astrologers
place Vervain under the dominion of Venus."

This same exhaustive researcher in the field of folklore and
botanical history tells us that the bridal wreaths of the Romans
were usually composed of vervain plucked by the bride herself.

In Brittany the vervain is known as "herb-of-the-cross". The
populace of Madrid was long accustomed, on St. John's Eve, to
wander about the fields in search of vervain, from a superstitious
notion that this plant possesses preternatural powers when gather-
ed at 12 o'clock on St. John's Eve. "The divining Gall-apple of
the Oak, the mystic Mistletoe, the Savin, the Moonwort, the Ver-
vain, and the St, John's Wort are considered magical, and therefore
form part of the witthes' pharmacopoeia — to be produced as oc-
casion may require, and their juices infused in the hell-broths,
philtres, potions, and baleful draughts prepared for their enem-
ies. -- Vervain and St. John's Wort, carried about the person,
will prove a sure preservation against the wiles of Satan and the

64, P Boils Od OcGatek Vol. 23, meen>

machinations and sorcery of witches:

'Gin you would be leman of mine

Lay aside the St. John's Wort and the Vervain.'
The Mistletoe, in addition to its miraculous medicinal virtues,
possesses the power of opening all locks; and a similar property
is by some ascribed to Artemisia, the Mandrake, and the Vervain."

Benchley (1970) says "Supposedly sophisticated Europeans still
splash themselves with exotic oils and drink doses of vervain" as
an aphrodisiac.

Lonicer (1679) speaks in old German of the medicinal virtues of
Verbena officinalis in his time: "Seyn gut zu den feuchten oder
fliessenden Wunden / oder alten Sch¥den. Die Wurtzeln seyn gut /
mit Wein getruncken / ftir die Geelsucht. Eisenkraut heilet aller-
ley innerliche Gebrechen / als die verstopffte Leber / Miltz und
Nieren / in Wein gesotten / und dartfber getruncken. Es hilfft
auch also getruncken / ftir das schwere Athemen / oder Keichen.
Eisenkraut gestossen /als ein Pflaster auf die Wunden gelegt /
heilet und trtficknet sie geschwind. Eisenkraut safft mit Wein ge-
truncken / benimmt des Gifft im Leib. Die Bl&tter und Wurtzel in
Wein gelegt /den Wein getruncken /benimmt das viert&gige Fieber.
Etliche meinen /zum dreytugigen Fieber soll man nemmen drey
Blatter / und zum viertYgigen vier BlXtter und vier Wuttzeln. Die
Blxtter ein Quintlin in Wein gethan /den also vier Tag lassen
stehen / und darnach im Mund gehalten / heilet die Geschwer darinn.
Eisenkraut mit Wasser gesotten / und gentttzt /reiniget den Frauer
ihre Mutter / und bringt ihnen ihre Zeit. Oder siede das Kraut
samt der Wurtzel in Wein / und trincks. Der Saame mit Fenchel-
safft vermischt / und in die Augen gelassen / reiniget sie und
machet klar Gesicht. Eisenkraut ist gut zur Leber / und zum erha-
benen und geschwollenen Miltz / wie gleichfalls zur siechen Lun-
gen. Eisenkraut mit samt der Wurtzel gestossen /darvon getruncken
/oder die Wurtzel pulverisiert und getruncken /vertreibt den Stein.
Die Bl&tter von Eisenkraut in Essig genetzt oder gestossen /und
tfhers heilig Feuer gelegt /ktfhlet und lUschet den Brand.....Das
Kraut und Wurtzel in Wein gesotten /den Mund damit gesptflet oder
geschwencket /ist gur ftir das effen und faule Zahlfleisch. Eisen-
kraut mit Honig vermischt /zeucht die Wunden zusammen. Das Kraut
mit altem schweinen Schmaltz zerstossen /und tibergelegt /leget
nider der Frauen Geschwulst an heimlichen Orten /zusamt den
Schmertzen......£isenkraut soll um S. Johanns Tag / mit aller Sub-
stantz gehackt und gebrannt werden. Eisenkrautwasser Morgens und
Abends / jedesmal drey Loth / sechs oder acht Tag nacheinander ge-
truncken / vertreibt die Geelsucht / ist gut fttr Gifft / dreyt&gig
oder viertugig Fieber / vertreibt die Witrm im Leib / alle Morgen
nffichtern getruncken. Hat die Krafft wie Gamanderlin. Das Wasser
wie obsteht getruncken / ist gut zur engen Brust / und ftir das
schwere Athmen / Geschwer der Lungen / Schwindsucht der Lungen /
sturcket die Leber / und bringt dem Menschen gute Farbe. Das Was-
ser ist auch gut getruncken ftir Schmertzen desz Magens / ftir Ver-
stopffung der Leber und Miltzes / auch Lenden und Blasenwehe.

Mehr ftir Verstopffung desz Eingeweids / Magens und Bauchs / auch
zur Geelsucht. Das Wasser reiniget die Nieren und Blasen vom

1972 Moldenke, Notes on Verbena L65

Gries und zertheilet den Stein in der Blasen. Getruncken dienet
es auch zu den Bl&ttern im Leib / Blutharnen und Grimmen. Eisen-
krautwasser ist das beste Wasser ft{fr Hauptwehe und Schusz desz
Haupts / die Stirn und Schl4ff offt damit bestrichen / und mit
Tiichlin tibergelegt / ist fast gut ftir langwirige Krankheiten / da
man nicht weis wovon sie entspringen. Das Wasser ist gut ftfr
alle Nebel und Geschwer der Augen / sttrckt das bl&Ude Gesicht /
bringt den Schein und Glast wider / alle Tag eimmal oder zwey
darein gethan / und darum gestrichen / gerieben / und Ttichlin
dartfber gelegt. Das Wasser ist auch gut ftir das Essen und Ltch-
er an der Frauen Gem&cht / dieselbige Morgens und Abends damit
gewaschen / auch Ttichlin darinn genetzt und thergelegt. Disz
Wasser vertreibt auch die Feigwartzen / damit gewaschen."

Hubert (1921) discusses the anatomy, chemical composition, and
uses of Verbena officinalis. Crevost & Pételot (193) tell us
that in Indochina "La plante est considérée pas les indigénes
comme amére et aromatique. D'aprés le R. P. Robert, elle passe
pour régulariser les menstrues et pour guérir la 'boule hystér-
ique'. On en fait, pour cela, use purée que l'on fait cuire et
qui se prend avec de l'alcool de riz."

Manning (1956) says that infusions of this plant are used in
reducing temperature in fevers and in the treatment of nervous
disorders. Ward (1967) gives V. hastata L. as a synonym, which,
of course, is ridiculous -- the two species cannot possibly be
confused by anyone who has ever seen them both! He says that the
plant is very bitter to taste, with a slight aromatic odor when
rubbed, and is a nervine, tonic, emetic, and sudorific in medical
usage. He notes that Coffin, writing 90 years ago, said "As an
emetic it ranks next to lobelia; it is also one of the strongest
sweating medicines in nature. It is good for colds, coughs and
pain in the head, and some years ago was highly esteemed as a
remedy for consumption. As an emetic it supercedes the use of
antimony and ipecacuanha to both of which it is superior, since
it not only produces all the good effects ascribed to the others,
but it operates without any of the dangerous consequences that
ever attach the use of antimonial preparations and cramps, and
even death have been known to follow their use.....Vervain will
relieve and cure those complaints in children which generally ac-
company teething; it likewise destroys worms, administered as a
tea it powerfully assists the pains of labour; as a diuretic it
increases the urinary discharge." Ward further states that the
ounce-to-pint infusion is now used and taken in wineglass doses,
and, as a nervine, Skullcap and Valerian are usually added.

Scopoli (1777) poses the question regarding the uses of Ver-
bena officinalis: "Verbenam officinalem ad Cephalaeam etiam in-
veteratam commendat Haenivs?" Smith informs us that even as far
away as the Ryukyu Islands this plant is regarded as medicinal,
being used to bathe babies; it is found abundantly there in sandy
and red earth. Airy Shaw (1966) points out that it was formerly
held in great repute in Europe as a remedy in eye diseases, its
bright corolla, like that of Euphrasia, being supposed, under the

4.66 P BY 20 Eh. 0O,G) D/A Vol. 23, no. 5

old Doctrine of Signatures, to indicate its virtue in that direc-
tion. Rageau (1957) says that in New Caledonia it is regarded as
"amére, Spre et aromatique. La plante entiére est utilisée comme
vulnéraire en médecine populaire frangaise".

Lazaro e Ibiza (1921) asserts that the species is found
throughout Spain, and there is regarded as "Es amarga y astrin-
gente, y fué usada como febrifuga, resolutiva y vulneraria; los
antiguos la consideraban como sagrada." Al-Rawi & Chakravarty
(1964) assert that the plant contains tannic acid, invertin, sap-
onin, verbenalin, verberin, and emulsin, and is used even now as
a corraborant, nervine, antispasmodic, febrifuge, tonic, aphrodis-
iac, antiscorbutic, galactagogue, detersive, emetic, sudorific,
diuretic, and against whooping cough.

Alzate (1968) affirms that Verbena litoralis and V. hispida
can be used as a substitute for V. officinalis where the latter
is unavailable, and have the same properties: "Substituyen a la
Verbena officinalis, comin en Europa y se usan en decocciones o
infusiones a dosis de 10 a 20 grms. por 200 de agua. Todos los
médicos reconocen el gran valor de nuestras verbenas, para cortar
las fiebres de origen gdstrico y tifoidea, administrando la decoc-
cién en lavados y en forma de tisana. Reemplaza a la quinina.
Los antiguos la consideraban como un remedio universal. Ahora
como febribugo, resolutivo y vulnerario. Las hojas y sumidades
bien machacadas suelen usarse en cataplasmas, como vulnerarias, y
el zumo se usa en infusién contra los dolores de cabeza. La
planta fresca machacada y aplicada sobre las sienes y frente,
calma el dolor de cabeza. Hervida en agua aplicada sobre heridas
sucias y de larga supuracién. El jugo de la planta, usado en
fricciones, cura la sarna. El zumo cura la diarrea, la bilis y
voluptuosidad. Las hojas y ramas hervidas curan y resuelven los
tumores internos (tomar copitas). Los romanos y griegos la
llamaban planta VIRGEN y la usaban en cataplasmas de hojas mach-
acadas sobre la vejiga para combatir los derrames seminales noc-
turnos."

Martinez (1969) informs us that in Mexico "El cocimiento de
las inflorescencias se usa para fortalecer el pelo. También se
usa el polvo contra la hidropesfa y la infusién teiforme o el
extracto contra las fiebres intermitentes (en Nay. [=Nayarit]).
En ZitA&cuaro, Mich. [=Michoac4n], usan el cocimiento de las
flores 'contra la biliosidad'. El Prof. Pedro M. Rodriguez dice
citando al Dr. Ricci, que es eficaz contra las fiebres. Se hace
un cocimiento de 5 gramos de hojas en medio litro de agua hirvi-
endo y se deja consumir hasta la mitad, se cuela y se toma en
ayunes durante varios dfas. En Nayarit toman el cocimiento antes
de cada comida contra la fiebre. En Teleoloapan, Gro. [=Guerrero],
lo toman con igual objeto, pero le afladen hojas de fresno.....la
hojas majadas y aplicadas con aceite rosado o con enjundia de
puerco, quita el mal de madre; y aplicdndolas con vinagre apargan
el fuego de San Antén y reprime las llagas corrompidas; mezcladas
con miel sueldan las heridas; su bebida es contra tiricia (icter-
icia) y su cocimiento de sus hojas y raices cura las hinchazones."

1972 Moldenke, Notes on Verbena 67

Gattinger (189) asserts that in his time "The herb in flower".
was used medicinally in Tennessee. Smith (1871) records the
Chinese vernacular name "ma-pien-ts'au" for the plant, roughly
translated as "horsewhip-plant". He affirms that the species is
as common in China as it is in England and that it has received
its common name from the long spiked inflorescence after the fall
of the deciduous corollas. It is confused by the country folk
with a species of Leonurus because of its similarly square stem.
It is said to act on the blood, relieving congestions, obstruc-
tions, dropsical effusions, and haematoceles. It is also cred-
ited with emmenagogue, anthelmintic, and antoscrobutic properties.
The root is said to be astringent.

Ferrdéndiz (1967) notes that the "Planta muy conocida en Europa
y cultivada en todos los jardines, vulgaramente se la denomina la
'Hierba de todos los males' y en verdad que tiene diversas apli-
caciones. Hervida la planta en agua se aplica sobre heridas tér-
pidas de larga supuraci6n. En forma de té, 15 gramos en un litro
de agua, calma la fiebre y en g&rgaras, alivia las afecciones de
boca y garganta. La planta fresca machacada, aplicdndola sobre
las sienes, alivia el dolor de cabeza. El agua destilada de
verbena fortifica el nervio 6éptico y el jugo de la planta en
fricciones, cura la sarna. Buena especialmente para males de
estémabo, por excitar la digestién. Excelente febrifugo."

Verbena officinalis has been studied chemically very exten-
sively. Cheymol (1937) reports the presence of stachyose in the
roots and stems, as well as a verbenaloside which is apparently
identical with cornine. Boudier (1907) isolated a verbenaline
glucoside, while Asano and his associates (1942) found verbena-
lin yielding verbenalol and glucose. Verbenalin and a volatile
oil were also reported by Breitwieser (193), while Boudier
(1908) isolated verbenalin, invertin, and emulsin. Huni and his
associates (1966) report that V. officinalis specifically incor-
porates 2—lic labelled melvalonic acid in the formation of a
glycone moiety of verbalin. Echaust (196) shows that the pre-
viously isolated verbenin and verbenalin are identical glycosides.
He isolated, in addition, adenosin and f-carotene. Kariyone
(1965) reports the same "adenosine" and P-carotene. Biichi & Man-
ning (1960) provide a revised structural diagram for verbenalin.
Reichert & Hoffmann (1935) proved that the glycoside, verbenalin,
from Verbena officinalis, is identical with cornin from Bentha-

midia florida. Stecher (1968) gives its formula as Cy 7H5) P10»

the molecular weight as 388.36, C 52.57 percent, H 6.23 percent,
and 0 41.20 percent. Sakan & Abe (1968) show that verbenalol is
the aglucone of verbenalin, a glycoside. The stereospecific syn-
thesis of racemic verbenalol is presented as one of the most pos-
sible biogenetic precursors of the indole alkaloids and has a
closely related structure to the components from Actinidia poly-
gama and Boschniakia rossica.

Horodysky and his associates (1969) have also studied the
biosynthesis of verbenalin (I) by the administration of acetate-
1-lhc, acetate-2- hig mevalonate-2 » and geraniol-1-l to

,

68 PHYTOLOGIA Vol. 23, noe 5

Verbena officinalis plants. When "I" which had been formed bio-.
synthetically from mevalonate-2—Lhc was degraded, randomization
of the label between carbon atoms 6 and 9 was not observed; carbon
atom 6 retained most of the radioactivity in young and old plants.
Randomization between carbon atoms 3 and 8 varied with the age of
the plant. There was complete randomization in young plants, lin-
ited randomization in older plants, and essentially non-randomiza-
tion in senile plants. The percentage of radioactivity in carbon
atom 3 predominated over that in carbon atom 8 as the plant age
increased. The ratio of radioactivity found in aglucone to that
in glucose is considered to be metabolically significant.

Savage (195) reports that under genus 35 VERBENA in the Lin-
nean Herbarium in London, sheet 15 is labeled "15 officinalis
[spuria delet.]" and has the accepted specific name in Linné's
own handwriting.

Kalm (1770) affirms that he personally saw this species grow-
ing in only one place in North America, shown to him by Bartram
"in a little plain near his house" near Philadelphia, Pennsyl-
vania. It is assumed that the species had been introduced there
from some seeds imported from England by Bartram.

Patzak & Rechinger (1967) give the overall distribution of the
species as "Europa a Britannia et Dania meridiem versus, Regio
Mediterranea, Africa borealis, Asia occidentalis et centralis
usque ad montes Pamir-Alaj et Tien Shan et ad Himalayas occiden-
tales; in America borealem et Africa australem et in regiones
alias introducta."

Martin (1965) describes the species as "rather local" in Eng-
land and Wales, while Paton (1968) says that in Cornwall and the
Isles of Scilly it is "frequent on waste ground during summer,
often with Odonites verna". Perring & Walters (1962) affirm
that it is "casual only" in the northern parts of its range in
the British Isles. Bowen (1968) reports it from Berkshire, Eng-
land, as "native, [in] dry grasslands, roadsides, in decreasing
small quantities". Druce (1897) referred to it as "Native, Viat-
ical. Dry gravelly waysides, pastures, &c. Local. More fre-
quent on calcareous soil in sunny situations [Berkshire, England]
eeee.eefound in all the bordering counties.....Rather common in
{Pang] district.....on chalk ballast near Reading". Dickinson
(1851) says of it "Roadsides and waste ground....Common....fre-
quent......Plentiful" in the Liverpool area,

Erfurth (1867) reports it from "Wegrunder, Gebtisch, Zdune, un-
bebaute Ort in der N&he der St&dte u. Dtrfer" in Germany. Wagner
(1905) says that it is found "An Wegr&ndern, auf Schutthaufen und
wiisten Pl&tzen in Mittel- und Stideuropa und Asien, ntrdlich bis
Stidschweden; im Gebiet ht#ufig." Dietrich (182) found it "In
Dtrfern, auf Schutthaufen, an Zdunen u.s.w. tfberall hdufig. Blttht
vom Juli bis September" in the Berlin area of Germany. He notes
that "Officinell war Herba Verbenae. Die Wurzel giebt den Gtirken
einen angenehmen Geschmack", Martens & Kemmler (1882) found it
"hdufig an Wegen und auf Schutt, doch selten tiber 580 m. auf-
steigend" in Wirttemberg. He also notes that the plant is men-

1972 Moldenke, Notes on Verbena 69

tioned by Pliny. Rechinger (1965) says of it "Gewohnl. E. -—-
Auf wiisten Platzen....bei Bauernhausern" in Austria. Ferguson &
Natzio found it growing on a shingle beach backed by saltmarsh in
Greece.

Montasir & Hassib (1956) state that Verbena officinalis is an
important plant in oases and along the western Mediterranean parts
of north Africa. Lazaro e Ibiza (1921) avers that it occurs in
"Toda Espafia". Sommier & Caruana Gatto (1915) say "Luoghi ruder-
ali e lungo le vie. — Malta e Gozo, abbastanza frequentet" In
Italy Lanfrossi (1827) refers to it as "Comunissima; cresce ne!
ruderi e lungo le strade e fioresce in Giugno, Luglio ed Adosta",
while Tornabene (1891) found it "in ruderatis, ad vias et in
hortis pedemontanae regionisi Catania, Mascalucia, Nicolosi" in
the Mount Etna region. He comments that "Olim apud medicos
laudata ob vim adstringentem, vulnerariam. Nunc exoleta." Ber-
tolini (18) asserts that it is "Vulgaris in viis, ruderatis,
collibus totius Italiae". In Belgium it was found by Sauzé &
Maillart (1880) on the "Bords des chemins, lieux incultes", while
both there and in Luxemburg Paque (1902) found it growing in
"Lieux incultes, bords des chemins",.

Catanzaro (1970) says that it inhabits "coltivati, incolti
coltivabili e aridi, lungo le strade" in the Bivona region. Polu-
nin (1969) reports that it is found in "Waste ground, waysides,
screes.......Much of Eur. (except Is. [=Iceland]): introd. IRL.
N. S. SF. [=Ireland, Norway, Sweden, Finland] Used in herbal
remedies". Bouloumoy (19305 found it in "Lieux incultes; bords
des chemins. Partout" in Lebanon and Syria. Harvey (1868) says
"V. officinalis is common and seems to be truly wild" in South
Africa. In his 1838 work he says "V. officinalis, or a species
much resembling it, is common in the | i cheered of cultivation".
In Lesotho it is said by Guillarmod (1971) to be "widespread in
Southern Africa: often a weed of cultivated ground".

In Afghanistan it is described by Aitchison (1880) as "Common
everywhere from Thal to the Kuram district up to 7000 feet; April
to July". In Nepal, according to Banerjee & Shakya, it is "lo-
cally distributed"; in Bhutan, according to Deb, Gupta, & Malick
(1968), it occurs in open situations; in Pakistan Hussain (1969)
says it grows "in moist ground around ponds". Tingle (1967) re-
ports it from Hongkong. Prain (1903) found it to be "A weed in
waste places. In all the provinces" of Bengal. Clarke (1885)
tells us that it is found in the "Himalaya, alt. 1--6000 ft.,
from Kashmir to Bhotan, frequent. Bengal Plain to the Sundera-
bunds, frequent. —- Distrib. Temperate and subtropical regions...
T. Thomson collected at Kussoor, and Clarke at Chumba, a
monstrous form with proliferous spikes, forming densely branched
panicles, the lower flowers all pedicelled, variously altered and
infertile" [this latter form is probably what is now known as
Stachytarpheta jamaicensis f. monstrosa (Moldenke) Moldenke, due
to a virus infection of Stachytarpheta jamaicensis (L.) Vahl, and
has nothing to do with Verbena officinalis].

Balakrishnan (196) records V. officinalis from Madras and

1,70 PHYTOLOGIA Vol. 23, no. 5

claims that it is the only wild species of the genus in India.
Doubtless he means that it is the only native wild species, since
at least three other species of Verbena have become naturalized
there or have escaped from cultivation in some localities. Rau
(1963) records it from altitudes of 1500--2500 meters in Uttar
Pradesh, India; Maheshwari (1963) found it to be "Common along
canal banks, near temporary puddles and stagnant water channels"
in the Delhi area, where it flowers and fruits from April to Octo-
ber. Puri (1960) says that it occurs frequently in both burned
and unburned areas in Indian forest quadrats and burned Pinus
longifolia forests. Banerji (1965) speaks of it as only "occas-
ional", Dattar & Majumdar (1966) assert that it is "distributed
in the temperate regions". Kapoor (1968) refers to it as "occas-
ional" in Kashmir, while the Banerjees (1969) record it from Bihar.

Ohwi (1965) speaks of it as a "Weed in waste grounds and along
roadsides in lowlands; Honshu, Shikoku, Kyushu. -- A nearly cosmo-
politan weed". Yamazaki (1966) refers to it as "Cosmopolitan in
tropical and subtropical regions". Walker found it to be a weed
in waste places on Okinawa.

Gilkey & Dennis (1967) tell us that "In ballast about Portland
[Oregon] has been found Verbena officinalis L......with...white
(rarely purplish) flowers". Wherry (1967) reports it from Dela-
ware County, Pennsylvania, while Domville & Dunbar (1970) refer to
it as "adventive, rare in rich thickets, Summer" in Ulster County,
New York. Carter & Jones (1968) report it from Forrest County,
Mississippi, but I imagine that their record is based on a mis-
identification of V. halei Small. Gattinger (189) found V. of-
ficinalis "Nearly everywhere about houses and settlements in

sandy soil" in Tennessee. Sudworth, in 1890, found it abundant in
the District of Columbia area. Tatnall (196) records it from New
Castle County, Delaware, Cecil, Sussex, and Worcester Counties,
Maryland, and northern Accomac County, Virginia. Radford, Ahles, &
Bell (196) refer to it as infrequent in waste places in Hyde,
Jones, and Mitchell Counties, North Carolina, and Aiken and Darling-
ton Counties, South Carolina, flowering there from June to October.

Lems (1960) records V. officinalis from Gran Canaria and Tene-
rife in the Canary Islands; Pampanini (1930) records it from Cyre-
naica, Hanson (1969) from Madeira, Porto Santo, Funchal, the Azores,
and the Canary Islands, and Sykes (1970) from Niue island. Peder-
sen (1969) reviews its distribution in Demmark, as well as its
extra-limital distribution in the rest of Europe, the Near East,
and northwestern Africa. Voigt (185) and Hara (1966) assert that
it is cultivated in India. Masamune (1955) records it from Tanega-
sima, Yakusima, Takarazima, Amami-osima, Iheyazima, Isagaki, Irio-
mote, and Yonakuni islands. Perring & Walters (1962) record it
from many localities in Eire, and Gupta (1967) from 2300 meters
altitude in Uttar Pradesh, India. Macbride (1960) avers that it
"probably" occurs in Peru, but he apparently saw no actual speci-
mens to substantiate this supposition, nor to date have I.

Abrams (1951) records this plant from Amador County, California,
and Portland, Oregon. Martinez (1969) reports it from "Hidalgo,

1972 Moldenke, Notes on Verbena 71

Valle de México, Estado de México, etc.", but I have personally
seen it only from Nuevo Leén and suspect that the other references
may apply to the very similar V. menthaefolia Benth., although V.
officinalis is certainly to be expected in and about Mexico City
and some of the other large cities of Mexico, especially ports.

Andersson (1859) records V. officinalis from the Galapagos Is-
lands on the basis of a Charles Darwin collection from James Is-
land, but gives in synonymy "V. caroliniana L. V. polystachya
HBK. var. foliis incisis, laciniis grosse serratis (Hook., l. c.
p. 195)". Examination of the Darwin specimen, however, has re-
vealed that it is actually V. glabrata var. tenuispicata Moldenke.
Hooker's plant and description apply to another Darwin collection
from the same island, which is now the type collection of V. sed-
ula var. darwinii Moldenke . The true V. officinalis L., therefore,
is not known from the Galapagos Islands.

Bouchet & Andy (1966) report that where V. officinalis is re-
garded as an undesirable weed it may be controlled by use of the
herbicides “dichlobenil" and "chlorthiamid", used at the rate of
3 kg. per hectare. It is not affected by buturon, parquat, diquat,
11561RP, nor any mixtures of these.

Hirata (1966) lists the following fungi as infesting the spe-
cies: Erysiphe cichoracearum (in Germany, Russia, and Yugoslavia),
Erysiphe polyphaga (in France, Switzerland, and Italy), Leveillula
taurica (in Iran), Oidium verbenae (in Cdiwica France, Greece, Is-
yael, Japan, Lebanon, Romania, and Switzerland), and Oidiun sp.
(in Spain) . Grove (1935) dds to this list Septoria verbenae Rob.
& Desm,.

I am very grateful to Dr. G. H. M. Lawrence and the Hunt Botani-
cal Library in Pittsburgh, Pennsylvania, for enabling me to see a
copy of the Macer (1477) work listed in the bibliography of this
species. Dr. Lawrence points out that the pages were originally
unnumbered and the Roman numerals now appearing on them were ap-
parently added at a much later date. The volume is discussed in
Hunt Bot. Cat. 1: 5 (1958) and the date of publication is given
as 1477, rather than "187" as given in Pritzel's "Thesaurus" or
as "ca, 1,90" as given elsewhere. I am indebted to Dr. J. J. Wur-
dack of the United States National Museum for the latter informa+
tion. He reports that Agnes Arber also discusses this work in her
book on herbals. This Macer reference is the earliest which I have
as yet seen personally, although the bibliographic history of the
species goes back to 37 B.C.

J. H. Moris, Flora Sardoa, vol. 3, also listed in the biblio-
graphy of V. officinalis in a previous installment of the present
series of notes, is actually inscribed "1858-1859" on the title-
page, but my good friend, Dr. W. T. Stearn, in a letter to me
dated March 8, 1972, says "regarding the publication of Vol. 3 of
Moris, Flora Sardoa, all I can say is that despite much search
the only review found is one of 1860 covering the work as a whole.
I can find no evidence that it was issued in parts and I should
thus simply date vol. 3 as '1859'",

72 P BeyOEsO LY O"G Teak Vol. 23, no. 5

Mr. F. J. Anderson has kindly researched in the library of the
New York Botanical Garden the date of publication of Georgi Jan's
"Hortus Elenchus". He reports that this work is not treated in
Pritzel nor in Stafleu, but according to the Catalogue of the
British Museum Library (2: 92) the plants listed were collected
in 1826 and the work published in 1831. He states, further, that
the Matthioli, Disc. Valgr., references in the same bibliography
of this species are apparently correct. "It is page 1107 of Dis-
corsi ne i sei libri....published by Valgrisi in 1585 at Venice.
No copy of that edition is in the N. Y. B. G. [=New York Botanical
Garden library] but we have the edition of same title, printed by
Valgrisi in 1568 and both the figure and the text relating to
Verbenaca appear on page 1107 of that work." He also notes that
"Johann Schr&tder's Chymic. Dispens. 1669 is actually his Pharma-
copeia Medico-Chymica first published in 161. Our library has
the 1649 ed. and it mentions Verbena on pages 167-168. As to
Orsin. Cap. Opusc. -- No trace of this author or title. It may
be part of a serial journal." The last reference in this memor—
andum is to the "Orsin. in Cap. Opusc. p. 288" citation given by
Bertolini (18),) which I have thus far been unable to verify.

It should be mentioned here that the two Maximowicz (1886) ci-
tations in the bibliography are often mis-dated "1887". The
Robert Brown reprint (1821) is dated as 1821 by Pritzel, but as
"1819" by Barnhart. The Tobe (1969) reference is erroneously
given as page "159" in the index of the work, while the Farns-
worth (1970) reference to item 15121 is erroneously listed in the
index of the work as item "15105". The illustration in Hatton's
work (1960) is said to have been taken from "M. (1565) 1052", but
as yet I have been unable to trace this reference.

The Curtis (1775) publication is often cited as having been
published in 1772, 1773, or 1777, but according to Stafleu, Tax.
Lit. 90. (1967), plate 1 was originally published in May of aly He
1777 is the title-page date for the whole volume, The Muller
(1775) publication is often cited as "1777" also, but here, again,
that is the volume title-page date; the title-page date for fas-
cicle eleven, in which plate 628 appears, is "1775". The Bulli-
ard work (1785) is often cited to vol. "3 & " or "5", but the
volume appears to be unnumbered, at least in the library of the
New York Botanical Garden.

The Sibthorp & Smith (1809) bibliographic reference is often
cited as "1806", but pages 219--l))2 of this work were not issued
until 1809. The Boswell Syme (1866) reference is sometimes
cited as "1863", but volume 6 of this edition was actually not
issued until 1866. The Druce publication (1898), although dated
"1897" on its title-page, was actually not published until 1898
according to a writer in Journ. Bot. 36: 10) (1898). The Gerarde
(1597) work is cited by some authors as "Ger. Em. 718".

It is also worth mentioning here that certain homonyms of names
listed in the synonymy of V. officinalis do not apply to this
plant at all: Verbena foemina Trag. is a synonym of Sisymbrium
officinale L. in the Brassicaceae, Verbena foemina Brunf. is a

1972 Moldenke, Notes on Verbena 473

synonym of Senecio vulgaris L. in the Carduaceae, and Verbena
recta sive mas Fuchs is a synonym of Sisymbrium officinale Le L. The
Verbena sacra ra Gerarde, included in the synonymy of V. officinalis
by me in a previous dhatalinent of this series of notes, appears
actually to be synonymous with V. supina L., as an examination of
the illustration given by Gerarde (1597, 1633) plainly shows.

Of considerable interest is the fact that Tournefort (1719)
lists four "kinds" of Verbena: (1) Verbena communis coeruleo
flore, Common vervain with blue flowers, (2) Verbena lusitanica,
Tatifolia, procerior, Portuguese verwain, (3) Verbena tenuifolia,
iinclenved vervain, and () Verbena urticae-folia, canadensis,
foliis incisis, flore majore, nettle-leaved vervain of Canada,
with cut leaves, and a larger flower. All of these except the
third have usually been relegated to the synonymy of V. officin-
alis — the third being regarded as V. supina L. -- but on this
disposition of these names I have grave reservations. More study
is required here.

Similarly, Haller (1768, p. 96) describes a variety "® Foliis
vix dissectis Hort. Florent. 98 [=Manetti, Virid. Florent. 98.
I751J". This is probably the same variety listed by Bertolini
(18h) as "Eadem foliis non, vel parum dissectis". Haller (1768,

100". It is not clear to me if either of these varietal agetee
tions apply to any of the eleven subspecific taxa at present
recognized in V. officinalis, but the former may be the same as
var. latiloba Sennen.

The Bayliss BS.3045, distributed as V. officinalis, is actual-
ly V. brasiliensis oa E. Contreras 5247 is V. carolina f.

21/35], Tha son. [Austin, 572/35] = sen. wn, (Austin, S/o /asl., me D.
Thomas Pcs and Vi Vollum s.n. [Fort Belknap, 1855] are all V. halei
Small; EC Contreras as 6152 is | is V. litoralis H.B.K.; Hodgkins s.n.
[Herb. Bot. Div. D. S. S. 1. R. 2052] is V. officinalis var. grandi-
flora Hausskn.; Healy 60/6) [Herb. Bot. ,. Div. D. S. I. R. 702h9a]
is V. V. officinalis var. prostrata Gren. & Godr.; T. J. Hale s.n.
[Baraboo, 1861] is xV. perriana Moldenke; and C. "Ritchie ie 57 is | is in
part V. rigida Spreng. and in part some other species. _

Love (1971) cites Murin & Sheikh s.n. from Iraq as the basis for
his report of the chromosome count of on = 1) in this species.
Guillarmod (1971) cites Dieterlen 829, Guillarmod }1) & 1435, and

Laydevant s.n. from Lesotho, deposited in the herbaria at % Capetown,

the University of Basutoland, the Albany Museum at Grahamstown, and
the National Herbarium at Durban. Franchet (1883) cites Franchet

1017 from Turkestan; Banerji (1965) cites Banerji 1069 from Nepal;
Deb, Sengupta, & Malick (1968) cite Sengupta ta 1271, "1275, & 1278 and
Deb 329 from Bhutan. [to be continued]

CERTAMEN MELASTOMATACEIS XIX.

John J. Wurdack
U. S. National Herbarium, Smithsonian Institution

Initial consideration of the Peruvian collections from the
northern outlier of the Cordillera Vilcabamba, Cuzco, and adjacent
regions in Huanuco and Ayacucho has been completed; the first
results on this material, collected under the aegis of the
National Geographic Society and National Science Foundation, were
published last year (Phytologia 21, No. 6. July, 1971). Con-
siderable Vilcabamba specimens remain named only to genus,
awaiting further collections in flower; there surely are more
undescribed species from this region of remarkable endemicity.
Included in the present discussion are ancillary notes on
related species of Miconia, chiefly Bolivian.

MICONIA SUBANDICOLA Wurdack, nom. et stat. nov.

Miconia glandulifera Cogn. var. boliviensis Cogn., Mem.
Torrey Club 6: 38. 1896. Non M. boliviensis Cogn., DC. Mon.
Phan. 7: 901. 1891.

The closest relative of M. subandicola is perhaps M.
aureoides Cogn., which has coarser branchlet pubescence (O.1-
O.2 mm long, rather than 0.05 mm), more conspicuously crenate-
dentate leaf blades, smaller flowers (petals only 5-5.8 m long,
rather than 7-9.5 mm; large anthers 5 mm long, rather than 6.7-
tt mm ) y torus within sparsely to moderately with gland-tipped
hairs (rather than with a fimbriate-ciliolate corona), style
subsparsely with gland-tipped hairs ca. O.1 mm long (rather than
moderately to densely puberulous with barbellate hairs to O.3 mm
long), and ovary cone glabrous or very sparsely glandular-
puberulous (rather than moderately to densely strigulose).
Miconia aureoides is represented by recent material from Colombia
(Putumayo, Schultes & Cabrera 18988), Ecuador (Santiago-Zamora,
Cazalet & Pennington 7709, more depauperate than typical), and
Peru (Loreto, Killip & Smith 28153, distributed as M. glanduli-
fera; Huanuco, Woytkowski 7901; Cuzco, Madison 10020, co-exist-
ing with M. subandicola); Killip & Smith 29528, distributed as
this species is actually of Sect. Miconia near M. smaragdina
Naud. Collections of M. subandicola are known from Peru
(Loreto, Klug 3930, distributed as M. aurea; San Martin, Bel shaw
3396, distributed as M. aureoides; Cuzco, Madison 10030, Dudley
10250) and Bolivia (Beni, 0. E. White 2330, distributed as M.
glandulifera; La Paz, Buchtien 1090, 1092, 1093; Cochabamba,
Bang 1173, type number of M. glandulifera var. boliviensis),
elev. 200-1600 m. I agree with Macbride's tentative synonymi-
zation (Field Mus. Publ. Bot. 13[4]: 414, 433. 1941) of M.
glandulifera under M. muricata (Don) Triana. Miconia muricata
differs from M. subandicola in the much coarser branchlet and

47

1972 durdeck, Certemen uelestoumsetaceis 475

hypanthial pubescence, noticeabiy undulate-denticulate leaf
blades which are rounded to cordulate at the base, somewhat
longer anthers (larger 8.4-8.6 mm long), torus within moderately
glandular-setulose (but without a corona), and ovary apex very
sparsely with clavate glands 0.05 mm long; a good recent match
in floral details for Mathews 1729 is Ferreyra 1623, from
Huanuco. In Cogniaux' monographic placement, the above-
discussed relatives were scattered (spp. 20, 65, 80) among three
sections, but all should be together in Sect. Tamonea.

MICONIA HOSPITALIS Wurdack, sp. nov.

M. aureoidi Cogn. affinis, ramorum pubescentia minore
foliis integris differt.

Frutex vel arbor parva 3-4.5 m. Ramuli paulo nodosi obscure
rotundato-quadrangulati demum teretes cavi paulo infra nodos
perforati in nodis linea interpetiolari obscure armati sicut
petioli foliorum venae primariae subtus inflorescentia hypan-
thiaque sparse pilis granuloso-stellulatis vix 0.05 mm longis
latisque armati. Petioli 1.5-4.5 cm longi; lamina (8-)12-25 x
(2.5-)3.5-6.5 cm oblongo-elliptica apice mucronulato-acuto vel
paulo (usque ad 1.5 cm) gradatimque acuminato basi acuta,
membranacea et integra, utrinque in superficie glabra, breviter
(0.5-1.5 cm) 3-plinervata (pari exteriore marginali neglecto)
nervis secundariis ca. 0.5-0.7 cm inter se distantibus nervulis
subtus subplanis laxe reticulatis (areolis ca. 0.7 m latis).
Panicula 8-11 X 5-8 cm submultiflora; flores 5S-meri (alabastris
maturis solum cognitis), pedicellis 3-4 mm longis et 0-0.2 m
infra hypanthium articulatis, bracteolis mox caducis non visis.
Hypanthium (ad torum) 2.6 mm longum; calycis tubus 0.7-0.9 m
altus intus minute (0.1 mm) et densiuscule strigulosus, lobis
interioribus 0.5 mm altis late ovatis, dentibus exterioribus
callosis non liberis inframarginalibus; torus intus glaber vel
sparse setulosus. Petala (paulo immatura) 4 X 2 mm obovato-
oblonga dense granulosa. Stamina paulo dimorphica; filamenta
sparse glanduloso-puberula; thecae (paulo immaturae) 3.7 mm vel
2.8 mm longae, connectivo basaliter glandulis paucis ornato.
Stylus basaliter sparse strigulosus pilis sparse barbellatis
0.5 mm longis; ovarium 5-loculare et ca. 2/3 inferum, apice
conico glabro 10-costato.

Type Collection: Carlos Schunke A-20 (holotype US
1470416), collected at Schunke Hacienda above San Ramon, Depto.
Junfn, Peru, elev. 1300-1700 m, July 1925.

Paratypes (both Peru, in young bud): Frank Wolfe 12416
(NA, US), from Camp 4 (Peligroso), southwestern slope of Rio
LlullaPichis watershed on ascent of Cerros del Sira, Depto.
Hugnuco, 9° 25' S, 74° hit W, elev. ca. 1585 m, 28 July 1969;
T. R. Dudley 11853 (NA, US), from between San Jose and
Huanhuachayo ca. 23 km SW of Hacienda Luisiana and Rfo
Apurimac, eastern massif of the Cordillera Central, Prov. la
Mar, Depto. Ayacucho, ca. 12° 42" S, 73° 47' W, elev. 1360-
1400 m, 19 Aug. 1968.

Miconia aureoides has pinoid cauline hairs 0.1-0.2 m long,m

476 PHYTOL, 0 Gris Vol. 23, n0a5

infranodal stem perforations, crenulate-denticulate leaf blades,
subpersistent floral bracteoles, probably larger petals, and
style sparsely glandular-puberulous nearly to the apex; the
larger flowers with internal toral corona and strigulose ovary
apices differentiate M. subandicola. The general vegetative
aspect of M. hospitalis is like that of M. flaccida Gleason
(vide infra), which has quadri-alate branchlets and eglandular
anther connectives.

MICONIA FLACCIDA Gleason
In an isotype (US) studied, each of three buds showed 16

stamens (with a minority in each flower smaller), style at the
extreme base sparsely glandular-puberulous (0.1-0.15 mm), and
ovary completely inferior (without a free cone). Ferreyra 4269
(Divisoria-Aguaita, Loreto) has obovate minutely callose-
serrulate leaf blades, 12 stamens in each of two buds, glabrous
style, and ovary with a developed 10-costate cone. Ferreyra 995
Divisoria, Loreto) has flowers (in each of 4 buds) with 10
stamens, style sparsely glandular-puberulous at the extreme base,
and ovary with a well-developed cone. Dudley 11843 and Madison
10239 (both from Huanhuachayo, 12° 43' S, 73° 50° W, La Mar,
Ayacucho) have 10 stamens in each of numerous dissected buds,
smaller hypanthia than in Killip & Smith 25625, glabrous style,
and well-developed ovary cone. Other than these permutations,
the above-cited collections are alike in the narrowly 4-alate
and infra-nodally perforated branchlets, foliage, and floral
details. The variability in this material does not seem further
resolvable without additional collections.

MICONIA REFLEXIPILA Wurdack, sp. nov.

Sect. Amblyarrhena. M. hamatae Cogn. affinis, inflores-
centiarum trichomatibus non crispatis filamentis puberulis
differt.

Arbor parva vel frutex 2-9 m. Ramuli primum sulcato-
quadrangulati demum teretes sicut petioli inflorescentiaque
pilis laevibus paulo vel arcte reflexis plerumque 0.8-1 mm
longis dense armati. Petioli 1-2(-2.5) cm longi; lamina 6-15 X
2.5-5(-6) em late lanceata apice acuto basi late acuta vel
anguste obtusa, rigidiuscula et obscure serrulata, supra dense
appresso-setosa pilis ad basim prominenter expansis (bulla ca.
0.6 mm lata longaque; apex setulifer ca. 0.6 m longus), subtus
foveolata et densiuscule setulosa pilis laevibus gracilibus
usque ad 1 m longis, 5-plinervata (pari interiore 0.5-1 cm
supra basim divergente) nervis secundariis 2-3 mm inter se
distantibus nervulis ultimis obscuris densiuscule reticulatis.
Panicula 5-7 X 5-7 cm submultiflora; flores 5-meri sessiles
interrupto-glomerati (glomerulis 3-5-floris), bracteolis ca.

1 X 0.4 mm ellipticis persistentibus. Hypanthium (ad torum)
1.9-2 mm longum extus dense appresso-setulosum pilis laevibus
gracilibus ca. 0.6-0.8 mm longis; calycis tubus 0.3 mm longus,
lobis interioribus 1 mm longis oblongis vel ovatis (apice
rotundato vel acutiusculo) extus dense strigulosis intus basim

1972 Wurdack, Certamen Melastomataceis 4,77

versus sparse strigulosis (pilis intus ca. 0.1 m longis),
dentibus exterioribus setuliferis inframarginalibus obscuris;
torus intus glaber. Petala 1.1-1.2 X 1.2-1.4 mm suborbicularia
sparse vel sparsissime glanduloso-ciliolata (ciliis 0.03-0.1 mm
longis) alioqui glabra. Stamina isomorphica; filamenta 1.3-
2.3 mm longa sparse glanduloso-puberula (pilis 0.1-0.3 mm
longis); thecae 1-1.3 X 0.5 X 0.4 mm oblongae apice minute uni-
poroso (poro 0.15 mm any, connectivo inconspicue (0.1-0.2
mm) prolongato non appendiculato. Stigma expansum 0.6-0.7 mm
diam.; stylus 3.1-3.4 X 0.3 mm modice puberulus (pilis p. p.
glanduliferis 0.15-0.2 mm longis gracillimis) in ovarii apicem
ca. 0.2 mm immersus; ovarium 5-loculare et 0.6-0.8 inferum, apic
conico setulifero setulis ca. 10 eglandulosis ca. 0.3 mm longis.

Type Collection: TT. R. Dudley 13244 (holotype US 2658940),
collected in dense cloud forest above Camp 3 (Laguna) just below
ridge leading to Camp 4 (Peligroso), southwestern slope of the
Rfo LlullaPichis watershed on the ascent of Cerros del Sira,
Depto. Huanuco, Peru, 9° 20' Ss, 74° 45? Ww, elev. ca. 1330 m,

24 July 1969. "Many-stemmed shrub 6-8 ft. tall; indument bright
burgundy-red."

Paratypes (all Peru): Huanuco: F. Wolfe 12322 (NA, US),
from Camp 3 (Laguna), ascent of Cerros del Sira, elev. 1290 m.
Junfn: Killip & Smith 25908 (US), from Porvenir, Pichis Trail,
elev. 1500-1900 m. Cuzco, Prov. Convencion, Cordillera Vilca-
bamba, ca. 12° 30' S, ca. 73° 30? W: M. T. Madison 10100 (NA,
US) and T. R. Dudley 10436 (NA), from Camp 2 1/2, elev. 1730 m;
T. R. Dudley 10372 (iA, US) and 10345 (NA, US), from between
Camps 2 1/2 and 3, elev. 1730-2000 m; T. R. Dudley 10434 (NA,
US), from Camp 3, elev. 2080 m.

Miconia hamata (Mem. N. Y. Bot. Gard. 16: 20. 1967) has
very fine and much crisped cauline, petiolar, and peduncular
hairs, as well as (from Gleason's dissection notes on the type
at Berlin 1932) glabrous filaments and more numerous (ca. 12)
flowers per capitulum; unfortunately recent topotypical collec-
tions are still unknown. Another relative, M. ruizii Naud., at
least as to the holotype (P), has basally nerved leaf blades,
patent stem hairs 1.5-2 mm long, and larger flowers (hypanthium
plus calyx tube [dry] 2.8 mm long, rather than 1.8 mm; petals
[ary] 1.8-2 mm wide rather than 1-1.2 m). ‘The Junfn collection
of M. reflexipila had been distributed as M. radula Cogn. (cf.
Bull. Torrey Club 58: 244. 1931; the US sheet however shows
puberulous filaments); that species has leaf blades obtusish to
rounded at the apex and much larger (and usually somewhat
pedicellate) flowers. The Cuzco collections of M. reflexipila
are all fruiting.

MICONIA POLYTOPICA Wurdack, sp. nov.

Sect. Amblyarrhena. M. pailasanae Wurdack affinis, foliis
distincte plinervatis bracteolis maioribus differt.

Ramuli primum obscure quadrangulati demum teretes sicut
petioli foliorum subtus venae primariae inflorescentiaque dense
strigosi pilis ascendentibus arcte (vel in inflorescentia laxe)

4,78 Py? 010,61 & Vol. 23, no. 5

appressis laevibus 0.7-1.5 mm longis. Petioli 0.8-3 cm longi;
lamina (2.5-)5-11 X (1-)2-4.4 cm lanceata vel elliptico-lanceata
apice anguste acuto basi late acuta vel obtusa, integra et
tenuis vel subcoriacea, supra modice vel dense strigulosa vel
strigosa pilis arcte appressis laevibus, subtus laxe strigulosa
vel setosa pilis gracilibus laevibus, 5-plinervata pari interi-
ore 0.5-1(-1.7) cm supra basim divergente nervis secundariis

ca. 1.5-2 mm inter se distantibus nervulis subtus planis

areolis 0.3-0.4 mm latis. Panicula 3-9 cm longa lataque sub-
multiflora; flores 5-meri subsessiles (pedicellis crassis
obscuris ) interrupto-glomerati, bracteolis 2.7-3.2 X 0.4-0.7
(-1) mm persistentibus. Hypanthium (ad torum) 2-2.7 mm longum
extus dense subsericeo-strigulosum vel strigosum pilis gracil-
ibus laevibus; calycis tubus 0.2-0.3 mm altus, lobis interiori-
bus oblongo-rotundatis 0.8-1.2 X 1-1.5 mm extus margines versus
glabris intus basim versus glandulis clavatis sparse strigu-
losis, dentibus exterioribus setuliferis obscuris inframarginal-
ibus; torus intus glaber. Petala 1.8-2.6 X 1.3-1.7 mm plerumque
glabra obovato-oblonga apice rotundato-truncato. Stamina iso-
morphica; filamenta 2.1-2.6 mm longa sparsiuscule glanduloso-
puberula pilis 0.1-0.2 mm longis; antherarum thecae 1.6-2 X
0.6-0.7 X 0.45-0.6 mm oblongae poro terminali 0.15-0.2 mm diam.;
connectivum nec prolongatum nec appendiculatum. Stigma expansum
0.5-0.7 mm diam.; stylus 4.4-6 X 0.25-0.35 mm modice glanduloso-
puberulus in ovarii apicem 0.2-0.3 mm immersus; ovarium 3-4-
loculare et 0.7-0.8 inferum, apice conico 0.3-0.5 mm alto
costulato modice pilis eglandulosis 0.2-0.4 mm longis coronato;
semina laevia 0.7-0.8 X 0.4-0.5 m.

MICONIA POLYTOPICA subsp. POLYTOPICA

Folia tenuia supra pilis plerumque 0.3-0.6 mm longis modice
strigulosa subtus sparse vel sparsiuscule laxo-strigulosa pilis
0.3-0.4 mm longis. Hypanthii pili ca. 0.8-1 mm longi.

Type Collection: T. R. Dudley 10346 (holotype US 2626833;
isotype NA), collected in dense cloud forest between Camps 2 1/2
and 3 ca. 15 km NE of Hacienda Luisiana and Rfo Apurimac ,
Cordillera Vilcabamba, Prov. Convencion, Depto. Cuzco, Peru,
12° 35? S, 73° 35? W, elev. 1730-2000 m, 22 June 1968.
"Multiple-stemmed arcuate to lax shrub 6-30 ft. tall, 2-3 in.
DBH; leaves dark green with red venation; peduncles and pedi-
cels burgundy-red; corolla inside white, outside reddish-purple;
anthers golden yellow."

Paratypes: Peru, Prov. Convencion, Depto. Cuzco: C. Vargas
— (US), from near Pintobamba, elev. 2600 m; T. R. Dudle
10438 (105 and 10533 (NA, US), M. T. Madison 10103 (NA, us), all
topotypical, elev. 1730-2000 m. Bolivia: 0. Buchtien 5498
(US), from Hacienda Simaco on trail to Tipuani, Depto. La Paz,
elev. 1400 m.

MICONIA POLYTOPICA subsp. AYACUCHENSIS Wurdack, subsp. nov.
Folia tenuia supra pilis 0.8-1 mm longis modice induta
subtus pilis ca. 1 mm longis paulo appressis dense obsita.

1972 Wurdack, Certamen Melastomataceis 479

Hypanthii pili 1.5-2(-4) mm longi.

Type Collection: M. T. Madison 10296 (holotype US 2626829;
isotype NA), collected in cloud forest in shade above Estera
Rohuana ca. 3 km from Huanhuachayo, Capprichio-Puncu trail on
west side of Apurimac valley, Prov. la Mar, Depto. Ayacucho,
Peru, 129 43" S, 73° 47" W, elev. ca. 2120 m, 18 July 1970.
"Shrub 4 m with pendant foliage; leaves dull green, with scarlet
indument on ribs and margins; fruit lavender, 4 mm diam."

Paratype (near-topotypical): TT. R. Dudley 11920 (NA, US),
ca. 30 km SW of Hacienda Luisiana and Rfo Apurimac.

MICQNIA POLYTOPICA subsp. HUANUCENSIS Wurdack, subsp. nov.

Folia subcoriacea supra pilis ca. 1 m longis densissime
induta subtus pilis ca. 1.2 mm longis paulo appressis densissime
obsita. Hypanthii pili 1.5-2.5 mm longi.

Type Collection: TT. R. Dudley 13521 (holotype US 2626828;
isotype za, collected in open elfin forest between Camp 4
(Peligroso) and Camp 5 (Tabano), southwestern slope of Rfo
LlullaPichis watershed on ascent of Cerros del Sira, Depto.
Huanuco, Peru, 9° 25' S, 74° 44" W, elev. ca. 1680 m, 31 July
1969. "Lax many-stemmed shrub 6-10 ft. tall; indument silvery-
violet."

Paratype (near-topotypical): F. Wolfe 12431 (NA, US), from
ea Camp 4 (Peligroso) and summit of the Sira, elev. ca.
1060 m.

The suggested Ecuadorian relative has leaf blades basally
nerved or barely (2-3 mm) pseudo-plinerved and inflorescence
bracteoles ca. 0.5 mm long. Miconia obscura (Bonpl.) Dc.
(larger leaf blades bullate-setulose above, foveolate and
densely setulose beneath) and perhaps M. lasiocalyx Cogn.
(larger leaf blades, some intermingled stellulate trichomes,
patent hypanthial hairs, larger calyx lobes) are more distantly
related congeners. To M. lasiocalyx have been referred Madison
10278 (Capprichio-Puncu trail, Prov. La Mar, Depto. Ayacucho
and Woytkowski 8196 (Mendoza, Depto. Amazonas} Macbride had
earlier (and correctly) referred Pennell 14053 (Rfo Yanamayo,
Depto. Cuzeo) to this species. Two collections (Killi & Smith
25628 and 25748, from Inehas, Pichis Trail, Junfn) distributed
incorrectly as M. ruizii Naud. represent a taxon closely related
to M. polytopica but with somewhat larger basally nerved or
barely pseudo=plinerved leaf blades (trichomes on the upper
surface basally somewhat expanded) and sparsely stout-ciliate
calyx lobes which are evenly strigulose externally; this
material is also suggestive of M. retropila Wurdack, but the
cauline and inflorescence pubescence is strongly appressed-
ascending.

MICONIA PAEMINOSA Wurdack, sp. nov.

Sect. Amblyarrhena. M. pailasanae Wurdack affinis, ramu-
lorum foliorum hypanthiorumque pubescentia ubique breviore
floribus minoribus differt.

Frutex 0.7-1.3 m. Ramuli primum obscure rotundato-

),80 Pry T 0.1% 0G Tk Vol. 23, nde 5

quadrangulati demum teretes sicut inflorescentia modice pilis
incurvo-patentibus laevibus ca. 0.5 mm longis armati. Petioli
0.8-2.5 em longi graciles; lamina (3.5-)5-10 X (1.3-)2-4 cm
elliptica vel oblongo-elliptica apice hebeti-acuto basi late
acuta vel obtusa, membranacea et obscure denseque ciliolato-
serrulata, supra modice aspero-strigulosa pilis vix O.1 mm
longis ad basim expansis, subtus in venis primariis secundariis-
que sparsiuscule appresso-setulosa (pilis ca. 0.1-0.2 mm longis)
in venulis ultimis superficieque glabra, breviter (0.2-0.5 em)
5-plinervata nervis secundariis ca. 2-2.5 mm inter se distanti-
bus nervulis subtus subplanis densiuscule reticulatis (areolis
0.2-0.3 mm latis). Panicula 4-9 X 3-8 cm pauciflora, ramis
primariis oppositis gracilibus; flores 5-meri plerumque breviter
(0.5-0.8 mm) pedicellati, bracteolis 0.3-0.5 X 0.1 mm subper-
sistentibus. Hypanthium (ad torum) 1.7-1.9 mm longum sparse
pilis laevibus subappressis ca. 0.2 mm longis indutum; calycis
tubus 0.1-0.2 mm longus, lobis interioribus 0.35-0.4 mm longis
oblatis glabris, dentibus exterioribus crassis sparse setulosis
non eminentibus. Petala 1.1-1.2 X 0.7-0.9 mm obovato-oblonga
glabra. Stamina isomorphica; filamenta 1.8-2 mm longa sparse
glanduloso-puberula pilis 0.05-0.1 mm longis; antherarum thecae
1.1-1.3 X 0.35 X 0.3 mm oblongae, poro O.1 mm diam. dorsaliter
inclinato, connectivo nec prolongato nec appendiculato. Stigma
paulo expansum 0.35-0.5 mm diam.; stylus 5 X 0.25 mm sparse
appresso-strigulosus pilis ca. 0.2 mm longis; ovarium 3-loculare
et ca. 2/3 inferum, apice conico 0.3-0.5 mm alto setulis paucis
O.1 mm longis coronato.

Type Collection: T. R. Dudley 13185 (holotype US 2626836;
isotype NA), collected in dense cloud forest between Camp 3
(Laguna) and Camp 4 (Peligroso), southwestern slope of Rfo
LlulaPichis watershed on ascent of Cerros del Sira, Depto.
Huanuco, Peru, 9° 26' S, 74° 45! Ww, elev. ca. 1450 m, 23 July
1969. "Lax to scandent shrub. Inflorescence violet-purple;
anthers golden; fruit bluish.”

Paratypes (both near-topotypical): TT. R. Dudley 13121 (NA,
US), from between Camp 3 and Camp 4, elev. ca. 1400 m; F. Wolfe
12367 (NA, US), from Camp 3, elev. 1290 m.

Miconia pailasana has vegetative pubescence 0.2-0.6 mm
long, hypanthial hairs 0.5-1 mm long, calyx lobes 0.6-0.8 mm
long, petals 1.8-1.9 X 1.4 mm, and ovarial hairs 0.3-0.35 m
long; also the second primary vein pair is only 0.5-0.8 mm from
the leaf margins, rather than 1.5-2.3 mm. Miconia polytopica
Wurdack seems more distantly related, having longer vegetative
and hypanthial pubescence, much larger bracteoles, and larger
flowers.

MICONIA PUNICEA Wurdack, sp. nov.

Sect. Amblyarrhena. De affinitate intima mihi incognita.
In aspectu M. ruizii Naud. reminiscente, ramorum pilis barbel-
latis stigmate non expanso differt.

Ramuli primum subquadrangulares demum teretes sicut petioli
foliorum venae primariae subtus inflorescentiarum ramique modice

1972 Wurdack, Certamen Melastomataceis 481

vel dense pilis erectis vel paulo reflexis laevibus vel sparsis-
sime barbellatis 1-1.5(-2) mm longis et pilis densiuscule
barbellatis paulo reflexis ca. 0.3-0.5 mm longis dense (ramulis)
vel modice (venis primariis) induti. Petioli 2-3.5(-8) om
longi; lamina 9-15.5(-27.5) X 3.5-8(-17) cm ovata apice
longiuscule gradatimque acuminato basi 0.4-0.6(-2) om cordata,
firme membranacea et serrulata (dentibus 0.5-0.7[-1] m profun-
dis et 2[-4+] mm inter se distantibus), supra modice bullato-
setosa bullis 0.5-0.7 X 0.3-0.5 mm setis laevibus 1-1.5 m
longis, subtus foveolata et densiuscule pilis crispulis laevibus
0.5-0.8 mm longis setulosa, 7(-9)-nervata nervis secundariis 3-5
mm inter se distantibus nervulis ultimis subtus planis obscuris
areolis 0.2-0.3 mm latis. Panicula 8-10 cm longa; flores 5-meri
sessiles in ramulis 1-2.5 cm longis racemosi, bracteolis 0.3-
0.5 mm longis persistentibus setulosis. Hypanthium (ad torum)

2 mm longum extus dense setulosum pilis 1-1.3 mm longis laevibus
vel sparsissime barbellatis; calycis tubus 0.2 mm altus, lobis
interioribus 1.2 X 0.8 mm oblongo-ovatis extus basim versus
sparse setulosis intus (glandulis minutis clavatis exceptis)
glabris, dentibus exterioribus setuliferis divaricatis 0.5 m
longis non eminentibus. Petala 2.2 X 1.5 mm obovato-oblonga
glabra. Stamina isomorphica glabra; filamenta 2 mm longa;
thecae 1.7 X 0.5 X 0.5 mm oblongae poro 0.15 mm diam., connec-
tivo paulo (O.4 mm) prolongato non appendiculato. Stigma
paullulo clavato-expansum 0.3 mm diam.; stylus 4.7 X 0.2 m
glaber in ovarii apicem 0.5 mm immersus; ovarium 5-loculare
apice conico 0.6 m alto glabro.

Type Collection: T. R. Dudley 10663 (holotype US 2658935),
collected in ceja in rainforest and cloudforest between Camps
21/2 and 3 ca. 15 km from Hacienda Luisiana and Rfo Apurimac,
Cordillera Vilcabamba, Prov. Convencion, Depto. Cuzco, Peru,
elev. 1730-2080 m, 1 July 1968. "Very common shrub 6-15 ft;
indument red."

Paratypes (same general region): T. R. Dudley 10815 (NA,
US, in young bud), from between Camps 3 and 4, elev. 2350 m;

M. T. Madison 10105 (NA; sterile), from Camp 2 1/2, elev.
1730 me

Miconia ruizii differs at least in the smooth cauline
hairs and capitellate stigmas. In petals, stamens, and pistil,
M. hirta Cogn. is similar; that Bolivian species differs in
the longer, stiffer, and sparser large cauline hairs (ca. 3 m
long), the rugose leaf blades with much sparser pubescence,
the esetulose hypanthia, and shorter calyx lobes (0.3 m long)
with tufts of subbasal gland-tipped hairs externally. Another
distant relative (ex char. and notes from the holotype, P) is
perhaps M. staphidioides (Naud.) Triana, with strongly barbel-
late or stipitate-stellate cauline hairs ca. 1 mm long, densely
barbellate hypanthial hairs, and densely setulose ovary apex,
but similar stamens, stigma, and style. The general aspect of
M. punicea, particularly the spicate inflorescence branches, is
like that of M. aeruginosa Naud. (Sect. Miconiea) or some of the
hispid species of Sect. Cremaniun.

482 PHY T 0 L0G 7 4 Vol. 23, no. 5

MICONIA MADISONIT Wurdack, sp. nov.

Sect. Amblyarrhena. M. lasiostylae Gleason et M. modicae
Macbride affinis, ramis arcte quadrangulatis alatis (alis
crassis ca. 0.5 m altis), calycis lobis et petalis maioribus
differt.

Rami arcte quadrangulati sicut petioli laminarum venae
primariae subtus inflorescentia hypanthiaque dense pilis
pinoideo-stellulatis 0.05-0.1(-0.2) mm longis demum caducis
induti. Petioli 1-2 cm longi sparse pilis laevibus 0.3-0.5 mm
longis inconspicue caduceque glanduliferis armati; lamina 12-20
X 4.5-7 em elliptica apice basique acuto, firme membranacea et
integra, obsolete ciliolata ciliolis distantibus crassis
appressis ca. 0.1 m longis, supra glabra, subtus in venulis
superficieque sparse subpersistenterque puberula pilis stellu-
latis ca. 0.2 mm latis sessilibus vel breviter (Coral mm) stipi-
tatis, 5-plinervata (pari tenui inframarginali neglecto) pari
interiore ca. 1-1.5 cm supra basim subalternatim divergenti
nervis secundariis ca. 4 mm inter se distantibus venulis ulti-
mis subtus planis areolis ca. 0.6 m latis. Panicula ca. 14 cm
longa pauciflora; flores 5-meri, pedicellis 1-2(-3) mm longis,
bracteolis ca. 2.5 X 0.5 mm lineari-lanceatis ante anthesim
caducis. Hypanthium (ad torum) 4 mm longum intus paulo 10-
costatum; calycis tubus 0.6-0.7 mm longus, lobis interioribus
2 X 3.5 m oblatis, dentibus exterioribus crassis appressis
inframarginalibus; torus intus sparse glanduloso-puberulus
pilis 0.05 mm longis. Petala 6 X 5-5.7 mm obovato-suborbicu-
laria dense granulosa extus sparse stellulato-puberula. Stamina
isomorphica; filamenta 4.8 mm longa densiuscule glanduloso-
puberula pilis 0.05-0.1 mm longis; thecae 3 X 1.5 X 1.2 m
rectae crasse oblongae poro 0.6 mm diam. dorsaliter inclinato,
connectivo nec prolongato nec appendiculato. Stigma expansum
2.1 mm diam.; stylus 7 X 1 mm dense glanduloso-puberulus;
ovarium 5-loculare et 2/3 inferum, apice conico costato 0.8 m
alto glabro.

Type Collection: M. T. Madison 10295 (holotype US 2626762;
isotype NA), collected in cloud forest on the Capprichio-Puncu
trail above Estera Rohuana (ca. 3 km from Huanhuachayo) on west
side of Rfo Apurimac valley, Prov. La Mar, Depto. Ayacucho,
Peru, 12° 43" Ss, 73° 47? W, elev. ca. 2224 m, 18 July 1970.
"Shrub 5 m. Leaves dull green, with tan scurfy indument below.
Calyx white, with rufous hairs; corolla and stigma white;
stamens bright yellow. Common."

Both suggested relatives have rotund-tetragonal branches
lacking the corneous elevations at the petiole insertions,
sepais only ca. 0.5-0.7 mm long, and petals only 2.5-5 mm long.
Certainly M. madisonii and the two next-described species are
in the general affinity of M. floribunda (Bonpl. ) DOSE qHakshe
species, at least as far as the Bonpland collection (P) is
concerned, has rotund-tetragonal branches, calyx lobes only 0.8
mm long, and anthers bent at almost a right angle at the base
with both thecae and connective well-prolonged ventrally beyond
the filament insertion. Certainly within the Colombian material

1972 Wurdack, Certamen Melastomataceis 4,83
currently ascribed to M. floribunda are several distinct species

MICONIA INCACHACANA Wurdack, sp. nov.

Sect. Amblyarrhena. M. madisonii Wurdack affinis, foliorum
subtus pilis longioribus floribus minoribus ovarii collo modice
glanduloso-ciliolato differt.

Rami arcte alato-quadrangulares (alis crassis ca. 0.6 m
altis) primum sicut petioli foliorum subtus venae primariae
inflorescentia hypanthiaque densiuscule pilis pinoideis ca. 0.1l-
0.15 mm longis induti demum glabrati. Petioli 1-2.5 cm longi;
lamina 10-18 X 4.5-7.5 cm elliptica apice acuto basi obtusa,
firme membranacea et integra, supra glabra, subtus in superficie
sparsiuscule pilis subclavatis 0.2-0.4 mm longis apicem versus
barbellatis investa, breviter (ca. 0.5 cm) 5-plinervata (pari
exteriore tenui inframarginali neglecto) nervis secundariis ca.
4-5 mm inter se distantibus nervulis ultimis subtus planis
areolis ca. 0.5 mm latis. Panicula 19 X 6 cm multiflora ramis
ascendentibus; flores 5-meri, pedicellis ca. 3 m longis,
bracteolis 3.5-5 X 0.2-0.3 mm demum caducis. Hypanthium (ad
torum) 3.4-3.6 mm longum intus paullo 10-costatum; calycis tubus
0.5 mm altus, lobis interioribus 0.7 X 3 mm oblatis, dentibus
exterioribus vix evolutis non eminentibus; torus intus glaber.
Petala 4.8-5 X 4.2 mm obovato-orbicularia dense granulosa extus
sparse caduceque stellulato-puberula. Stamina isomorphica;
filamenta 4 mm longa modice pilis glanduliferis ca. 0.1 m
longis obsita; thecae 2.5 X 1.1 X 1.3 mm crasse oblongae poro
0.5-0.6 mm diam. dorsaliter inclinato, connectivo nec prolongato
nec appendiculato. Stigma expansum 1.6 mm diam.; stylus
densiuscule pilis glanduliferis ca. 0.1 mm longis indutus;
ovarium 5-loculare et 0.8 inferum, collo 0.6 mm alto modice
pilis glanduliferis 0.1 mm longis coronato.

Type Collection: J. Steinbach 9459 (holotype US 1857472),
collected in forest at Incachaca, Prov. Chapare, Depto.
Cochabamba, Bolivia, elev. 2200 m, Feb. 1929. "Baum 8 m."

Miconia madisonii has much shorter foliar hairs, somewhat
larger flowers, and glabrous ovary apices. Miconia modica
Macbride has obtusely tetragonal branches, shorter leaf indument,
and slightly smaller flowers (hypanthium plus calyx tube ca.

2.5 mm long dry, rather than ca. 3.5 mm). Miconia lasiostyla
Gleason has obtusely tetragonal branches, strongly plinerved
leaf blades, and considerably smaller subsessile flowers (but a
glandular-setulose ovary apex).

MICONIA TERBORGHII Wurdack, sp. nov.

Sect. Amblyarrhena. M. madisonii Wurdack affinis, foliorum
subtus pilis minoribus mox caducis petalis stigmateque maioribus
ovarii apice glabro differt.

Frutex vel arbor parva 3-6 m. Rami arcte quadrangulati
(alis crassis 0.7-1 mm altis) ad nodos linea cornea transversali
1-1.5 mm alta armati sicut petioli foliorum subtus venae pri-
mariae secundariaeque inflorescentia hypanthiaque pilis

48h PH er OL (OG. Ta Vol. 23, no. 5

pinoideis ca. 0.03-0.05 mm altis latisque demum caducis sparse
vel modice armati. Petioli (1.5-)3-4(-6) cm longi; lamina
12-22(-26) X 5-10.5 em elliptica vel oblongo-elliptica apice
acuto vel paullo (usque ad J em) acuminato basi late acuta vel
obtusa, firme membranacea vel subcoriacea, integra vel obscure
distanterque calloso-serrulata, ubique primum sparse stellulato-
puberula (pilis 0.05-0.07 mm latis sessilibus) mox glabrata,
breviter (0.5-1.5 cm) 5-plinervata (pari exteriore inframarginmli
tenui neglecto) nervis secundariis ca. 4-5 mm inter se distanti-
bus nervulis subtus planis areolis irregularibus ca. 0.8-1 mm
latis. Panicula 15-30 em longa multiflora, ramis ascendentibus;
flores 5-meri, pedicellis 2-4 mm longis, bracteolis 1.3-2 X 0.1-
0.2 mm linearibus mox caducis. Hypanthium (ad torum) 3.4-3.7 mm
longum intus paullo 10-costatum; calycis tubus 0.1-0.2 mm altus,
lobis interioribus 1.2-1.4 X 3 mm oblatis, dentibus exterioribus
crassis appressis non eminentibus; torus intus glaber. Petala
7-8.5 X 5.8-8 mm obovato-suborbicularia dense minutissimeque
granulosa. Stamina isomorphica; filamenta 5-6 mm longa modice
pilis glanduliferis 0.1-0.15 mm longis obsita; thecae 223 se
1.6-1.8 X 1.6-1.7 mm crasse oblongae poro 0.6 mm diam. ventra-
liter inclinato, connectivo nec prolongato nec appendiculato.
Stigma expansum 2.3-2.4 mm diam.; stylus 5.5-7 X 0.9 mm
densiuscule pilis glanduliferis O.1-0.2 mm longis indutus in
ovarii apicem O.1-0.2 mm immersus; ovarium 5-loculare et ca.

2/3 inferum, apice conico costato ca. 1 mm alto glabro.

Type Collection: T. R. Dudley 10890 (holotype US 2626763;
isotype NA), collected just below Camp 5 ca. 23 km NE from
Hacienda Iuisiana and Rio Apurimac, Cordillera Vilcabamba, Prov.
Convencion, Depuo. Cuzco.) Peru, Meo 3530'S. 730 351. W, See ca.
2700 m, 9 July 1968. "Shrub to 20 ft. ei: aegis red.

Calyx deep purple; corolla purplish in bud, opening whitish-
pink; anthers deep orange."

Paratypes (all near-topotypical): M. 7. ae 10146
(NA, US), from Camp 4, elev. 2663 m; T. R. Dudley 10794 (NA,
us), from between once 4 and 5, elev. EOE m; T. R. Dudley

10772 (NA, US), from just below Camp 5, elev. 2700 m.

Miconia madisonii, M. incachacana, and M. terborghii all
have sharply alate-quadrangular branches, but differ markedly
in pubescence quality as well as (to a lesser extent) floral
details. John Terborgh has been the factotum in the Vilcabamba
explorations as well as the principal zoological collector.

MICONIA MONZONENSIS Cogn. subsp. CUZCOENSIS Wurdack, subsp. nov.
Folia longiuscule acuminata hypanthia paulo minora differt.
Type Collection: M. T. Madison 10147 (holotype US 2626751;
isotype NA), collected in elfin forest near Camp 4, Cordillera
Vilcabamba on east side of Rio Apurimac across from Hacienda
Iuisiana, Prov. Convencion, Depto. Cuzco, Peru, 12° 37' S,
73° 33' W, elev. ca. 2568 m, 26 June 1970. "Small tree 2-3 m;
leaves pleaey light green with red ribs. Corolla and filaments
white; anthers bright yellow. Common."
Paratype (near-topotypical): 1. R. Dudley 10694 (NA, US),

1972 Wurdack, Certamen Melastomataceis 85

from between Camps 4 and 5, elev. ca. 2510 m.

The typical subspecies has leaf apices very shortly and
bluntly acute and hypanthium plus calyx ca. 3 mm long (dry),
while the Cuzco subspecies has leaf apices subcaudate-acuminate
for 1-2 cm and hypanthium plus calyx ca. 2.3 mm long (dry).

MICONIA KOEPCKEANA Wurdack, sp. nov.

Sect. Amblyarrhena. M. monzonensi Cogn. affinis, foliis
minoribus basim versus attenuatis floribus minoribus differt.

Ramuli argute quadrangulati (costis crassis 0.1-0.15 m
altis) primum sparse resinoso-granulosi glabrati. Petioli
liberi 0.3-0.5 cm longi; lamina 4-6 X 1.4-2 cm elliptica apice
hebeti-acuto basi anguste acuta, subcoriacea et integerrima,
glabra, breviter (ca. 0.5 cm) 3-plinervata nervis secundariis
paulo obscuris ca. 2 mm inter se distantibus venulis supra
invisis subtus planis areolis ca. 0.4 mm latis. Panicula 5-10
em longa (pedunculo ca. 3 cm longo incluso) submultiflora;
flores 5-meri plerumque in ramulis terminales plerumque 3-
glomerati, pedicellis 0.3-0.5 mm longis, bracteolis ca. 2 X
0.25 mm caducis. Hypanthium (ad torum) 1.9 mm longum extus
sparse resinoso-granulosum; calycis tubus O.1 mm altus, lobis
interioribus 0.3-0.4 m longis deltoideis remotis, dentibus
exterioribus obscuris omnino adhaerentibus non eminentibus.
Petala 1.5-1.6 X 1.1-1.2 mm obovato-suborbicularia extus glabra
intus minutissime granulosa. Stamina isomorphica glabra;
filamenta 2.5 mm longa; thecae 1.3 X 0.5 X O.4 mm oblongae poro
O.1 m diam. dorsaliter inclinato, connectivo simplici. Stigma
paullulo expansum 0.3 mm diam.; stylus 6.6 X 0.25 mm glaber vel
sparsissime glandulis 0.05 mm longis obsitus in ovarii apicem
0.2 mm immersus; ovarium 3-loculare et ca. 3/4 inferum, apice
conico 0.3 m alto glabro.

Type Collection: TT. R. Dudley 13143 (holotype US 2658937),
collected in elfin forest about halfway between Camps 2 and 3,
ascent of Cerros del Sira, southwestern slope of the Rfo
LlullaPichis watershed, Depto. Hudgnuco, Peru, 9° 27' S, 74°
46? W, elev. ca. 1000 m, 23 July 1969. "Shrub ca. 4 ft.; leaves
glossy green above. Flowers white; anthers yellow."

Paratypes (both near-topotypical): F. Wolfe 12413 (NA,
US), from just below Camp 4, elev. 1480 m; T. R. Dudley 13127B
(naj, from between Camps 2 and 3, elev. ca. 1000 m.

Miconia monzonensis has leaf blades 2.5-4 cm wide with
rounded bases, as well as considerably larger flowers (hypan-
thium 2.2-3 mm long; petals ca. 2 m long) on longer pedicels
(ca. 1 mm) and with more expanded (0.5-0.7 mm) stigmas. The
species epithet commemorates Drs. H. W. and Maria Koepcke, whose
hospitality at Casa Humboldt in Lima was much appreciated by
visiting scientists and whose zoological and ecological works
are landmarks in Peruvian natural history. Maria Koepcke,
tragically killed in an airplane crash, accompanied Dr. Dudley
during his Sira trip.

4,86 PHY TOLOGGIiIA VoL. 23, now Ss

MICONIA WOLFEI Wurdack, sp. nov.

Sect. Amblyarrhena. M. monzonensi Cogn. affinis foliis ad
basim attenuatis calycis lobis longioribus differt. AM.
koepekeanae Wurdack differt foliis plerumque latioribus floribus
maioribus calycis lobis longioribus ovarii apice glanduloso-
puberulo.

Ramuli argute quadrangulati (alis crassis ca. 0.5 m altis)
sicut inflorescentia hypanthiaque primum sparse resinoso-fur-
furacei glabrati. Petioli ca. 0.5 cm longi; lamina 5-7 X (1.5-)
2-3 em elliptica apice hebeti-acuto basi acuta, subcoriacea et
obscure distanterque serrulata, glabra, breviter (0.3-0.5 em) 3-
plinervata (pari exteriore inframarginali tenui neglecto) nervis
secundariis ca. 2-2.5 mm inter se distantibus subtus elevatis
nervulis ultimis subtus planis areolis ca. 0.3-0.4 mm latis.
Panicula 4-7 em longa submultiflora; flores 5-meri, pedicellis
(1.5-)2.2-3 mm longis, bracteolis ca. 1.5 X 0.1-0.2 mm valde
caducis. Hypanthium (ad torum) 1.8-1.9 mm longum; calycis tubus
0.2 mm longus, lobis interioribus 1-1.2 mm longis, dentibus
exterioribus crassis adhaerentibus lobos interiores aequantibus.
Petala 2.1 X 1.8 mm suborbicularia extus glabra intus minutissine
pruinoso-granulosa. Stamina isomorphica glabra; filamenta 2.3
mm longa; thecae ca. 1.4 X 0.6 X 0.5 mm oblongae poro 0.1 m
diam. dorsaliter inclinato, connectivo dorsaliter ad basim
obscure dentato. Stigma paullulo expansum 0.25 mm diam.; stylus
4.7 X 0.2 mm basim versus sparsissime glanduloso-puberulus in
ovarii apicem ca. 0.2 mm immersus; ovarium 3-loculare et ca. 0.8
inferum, apice conico lobulato pilis glanduliferis 0.05-0.1 mm
longis modice coronato.

Type Collection: F. Wolfe 12he9 (holotype US 2626752; iso-
type NA), collected in cloud forest between Camp 4 and summit of
Cerros del Sira, southwestern slope of Rio LlullaPichis water-
shed, Depto. Huanuco, Peru, 9° 25% S, 74° 43" W, elev. ca. 1880
m, 29 July 1969. "Shrub 6 ft. tall. Flowers white."

Paratypes (both near topotypical, in bud): F. Wolfe 12445
(NA, US) and 12447 (NA, US), from near the summit of Cerros del
Sira, elev. 2200 m.

Miconia monzonensis has basally rounded leaf blades, calyx
lobes ca. 0.5-0.7 mm long, and glabrous ovary apices; M. koepckeam
has entire leaf blades only 1.4-2 cm wide, smaller flowers on
much shorter pedicels, and glabrous ovary apices. Miconia
longisepala Gleason does not seem to be an intimate relative of
M. wolfei, having thinner acuminate basally nerved leaf blades,
densely glandular-puberulous filaments and style, and a much-
expanded stigma. Other species in this general alliance recent-

ly described by me (M. car ishana, M. beneolens, M. cajanumana,
M. tachirensis, M. ase differ in other details.

MICONIA ALIGERA Wurdack, sp. nov.

Sect. Amblyarrhena. M. monzonensi Cogn. affinis, foliis
proportionaliter angustioribus ad apicem caudato-acuminatis ad
basim attenuatis venulis subtus laxioribus differt.

Rami prominenter alato-quadrangulati (alis 1.5-2 mm altis)

1972 Wurdack, Certamen Melastomataceis 487

primum sicut folia inflorescentia hypanthiaque sparse resinoso-
furfuracei mox glabrati. Petioli liberi ca. 0.5-0.8 cm longi;
lamina (acumine excluso) 9.5-19 X 3-6.5 cm oblongo-elliptica
apice per 1-1.5 cm caudato-acuminato basi anguste acuta et in
petiolem decurrente, firme membranacea et integra, obscure
calloso-ciliolata, breviter (0.4-0.8 cm) 3-plinervata (pari
exteriore ca. 2-3 mm inframarginali) nervis secundariis plerum-
que 4-5 mm inter se distantibus nervulis subtus planis areolis
ca. O.4 mm latis. Panicula 6-8 cm longa multiflora, ramis
ramulisque breviter (0.5-1 mm) 4-alatis; flores 5-meri sub-
sessiles (pedicellis ca. 0.3 mm longis) in ramulorum apices 3-
4-glomerulati, bracteolis 2.3-2.6 X 0.3-0.4 mm acutis usque ad
anthesim persistentibus. Hypanthium (ad torum) 2.62.7 mm
longum teres; calycis tubus ca. 0.2 mm longus, lobis interiori-
bus ca. 0.2 mm longis remotis, dentibus exterioribus obscuris
appressis non eminentibus; torus intus obscure sparsissimeque
glandulis 0.05 mm longis ornatus. Petala 1.7 X 1.1-1.2 m
obovato-suborbicularia minutissime granuloso-pruinosa. Stamina
isomorphica glabra; filamenta 2.4 mm longa; thecae 1.9-2 X 0.7 X
0.5 mm oblongae ventraliter ad basim per ca. O.1 m steriles
poro 0.1-0.15 mm diam. dorsaliter inclinato, connectivo non
prolongato dorsaliter ad basim obscure (ca. 0.1 mm) hebeti-den-
tato. Stigma paulo expansum 0.6 mm diam.; stylus 6 X 0.3-

0.5 mm apicem versus expansus basim versus sparsissime glandu-
lis 0.05-0.1 mm longis obsitus in ovarii apicem ca. 0.3 m
immersus; ovarium 5-loculare et ca. 3/4 inferum, apice conico
0.4 mm alto sparsissime glandulis sessilibus coronato.

Type Collection: 1. R. Dudley 11261 (holotype US 2658934),
collected in rainforest and low cloudforest at Camp 3 ca. 15 km
NE of Hacienda Iuvisiana and Rfo Apurimac Cordillera Vilcabamba,
Prov. Convencion, Depto. Cuzco, Peru, 12° 30' Ss, 74° 30? w,
elev. 2080 m, 21 July 1968. "Large shrub to small tree 10-30
ft. tall, DBH 3"-6", with dense columnar crown; young leaves
bronze. Corolla white; anthers bright orange. Common melastome
at this altitude."

Paratype (topotypical): 1. R. Dudley 10612 (NA, US).

Miconia monzonensis has narrower branch wings (ca. 0.6 m
high), apically bluntly acute and basally obtuse leaf blades
only about twice as long as wide, and leaf veinlets densely
reticulate beneath (areoles ca. 0.25 mm wide). Other species
of Sect. Amblyarrhena with sharply quadrangular branches (M.
bailloniana Macbride, M. expansa Gleason) are not very closely
related, having 5-plinerved leaf blades and much larger flowers

petals 4-5 mm long). Ome sterile collection, Madison 10285
NA) from the Capprichio-Puncu trail, Prov. La Mar, Depto.
Ayacucho, elev. 1980 m, may also represent M. aligera.

MICONIA ADINANTHA Wurdack, sp. nov.

In aspectu M. elatae (Sw.) DC. affinis, inflorescentiae
ramis in quoque nodo k ovarii apice glabro differt.

Ramuli obtuse tetragoni sicut folia subtus inflorescentia
hypanthiaque densissime stellulato-puberuli pilis ca. 0.c5 m

4,88 PHYTOL OG LA Vol. 23, now &

diam. Petioli 1.5-2.5 cm longi; lamina 13-20 X 5-8 cm obovato-
elliptica apice breviter (0.5-1 cm) subgradatimque acuminato
basi acuta, subcoriacea et obscure distanterque undulato-serru-
lata, supra glabra, 3-nervata (pari exteriore ca. 1 mm infra-
marginali tenui neglecto) nervis secundariis ca. 5 mm inter se
distantibus subtus prominentibus nervulis subtus planis (areolis
ca. O.3 mm latis) ob pilos occultis. Panicula 10-15 X 7-12 cm
multiflora, ramis primariis in quoque nodo 4(-6); flores (5-)6-
meri diplostemoni subsessiles (pedicellis obscuris ca. 0.2 mm
longis) in ramulis conferte interrupto-glomerati, bracteolis
mox caducis non visis. Hypanthium (ad torum) 1.2 mm longum;
calycis tubus 0.2 mm altus, lobis interioribus ca. 0.1 mm longis
dentibus exterioribus obscuris non eminentibus. Petala 1.6-1.7
X 1 mm oblongo-obovata obscure granulosa. Stamina paullulo
dimorphica glabra; filamenta 1.6 mm longa; thecae 1-1.2 X 0.45
mm oblongae late biporosae, connectivo ad basim ventraliter
paullulo (0.1 mm) bilobulato dorsaliter minute (0.1-0.15 mm)
dentato. Stigma paullulo expansum 0.25 mm diam.; stylus 4.2 x
0.2 mm glaber; ovarium 3-loculare et 3/4 inferum, apice conico
O.1-0.15 mm alto glabro.

Type Collection: T. R. Dudley 10384 (holotype US 2626822;
isotype NA), collected in undisturbed cloud forest at Camp 2 1/2
12 km NE of Hacienda Luisiana and Rio Apurimac, Cordillera
Vilcabamba, Prov. Convencion, Depto. Cuzco, Peru, 12° 30' S,
73° 30' W, elev. 1730 m, 23 June 1968. "Tree 50-60 ft. with
dense rounded crown, DBH 6-8 inches; phloem pale pink. Leaves
dark green and glossy above, with dense rufous indument beneath.
Flowers creamy-white."

Despite the plethora of Andean and subandean species with
large discolorous leaf blades, I am at a loss to discover the
really close affinities of M. adinantha. Miconia elata has
distinctly 5-nerved leaf blades, opposite inflorescence branches,
and stellulate=puberulous ovary apices; the only collection
(fruiting) of M. elata known from south of Colombia is C. M.
Belshaw 3472 (US), from Lamas-San Antonio, Depto. San Martin,
Peru. Among the species with four inflorescence branches per
node (variously separated in Cogniaux? system, but surely more
closely related than now indicated), M. crassipes Triana, M.
pavoniana Naud., M. ioneura Griseb., and M. herrerae Gleason
all have larger pleiostemonous (15-24 stamens per flower)
flowers, the first-mentioned also showing stellulate-puberulous
ovary apices. Miconia multiflora Cogn. and M. molybdea Naud.
have petals externally sparsely to moderately stellulate-
puberulous and anther connectives glandular at the base.

Miconia hygrophila Naud., with inflorescence branches 4 per
node, is dioecious and with sparser stipitate-stellulate hairs
on the lower leaf surfaces. My survey for relatives of M.
adinantha did not include the eastern Brazilian species of Sect.
Glossocentrum, but all these differ at least vegetatively.

MICONIA RECONDITA Wurdack, sp. nov.
Sect. Cremanium. M. neriifoliae Triana, M. floccosae Cogn.

1972 Wurdack, Certamen Lielastomataceis 489

et M. biacutae Cogn. affinis, foliis subtus in superficie glabra-
tis inflorescentiae ramis sparse glanduloso-setulosis differt.

Ramuli argute quadrangulati sicut foliorum subtus venae
primariae inflorescentiae ramique primum modice pinoideo-puberuli
demum glabrati. Petioli 1-2.5 cm longi; lamina 7-le X 2.5-3.5
em anguste elliptico-oblonga apice late hebeti-acuto basi obtusa
vel paulo rotundata, coriacea et distanter obscureque calloso-
serrulata, supra glabra, subtus in superficie primum sparsiusaile
pinoideo-puberula (pilis demum caducis) et secus nervos pri-
marios praecipue basim versus sparsissime glanduloso-setulosa,
3-nervata (pari exteriore inframarginali neglecto) nervis
secundariis 2-3 mm inter se distantibus nervulis subtus planis
obscuris (areolis ca. 0.5 mm latis). Panicula 4-7 cm longa
submultiflora, ramulis oppositis pilis laevibus glanduliferis
ca. 0.3 mm longis sparse setulosis; flores 5-meri crasse pedi-
cellati (pedicellis ca. 1 mm longis), bracteolis valde caducis
non visis. Hypanthium (ad torum) 3.4 mm longum extus sparse
(praecipue basim versus) pinoideo-puberulum demum glabratum;
calycis tubus 0.4 mm altus, lobis interioribus 0.5-0.6 m altis
rotundatis, dentibus exterioribus crassis non eminentibus.
Petala glabra 2.6-2.9 X 2.6-2.7 mm suborbicularia, apice retuso.
Stamina paullulo dimorphica glabra; filamenta 2.3 mm longa basim
versus dilatata; antherarum thecae 1.9 X 0.8 X 0.7 mm vel 1.6 X
0.9-1 X 0.7 mm obovato-oblongae late biporosae, connectivo ad
basim ventraliter ca. 0.2-0.3 mm prolongato bilobulato. Stigma
paulo expansum 0.8 mm diam.; stylus 3.3 X 0.6 mm glaber;
ovarium 2(?)-loculare et 2/3 inferum, apice conico 1 m alto
glabro.

Type Collection: M. Bang 2485 (holotype US 1416564; iso-
a. US), collected at Unduavi, Depto. La Paz, Bolivia, Sept.
al .

The three suggested relatives all have leaf blades beneath
and hypanthia completely and persistently covered with pinoid
(long-barbellate, appearing flocculose) hairs and also lack
glandular inflorescence hairs; the flowers of both M. neriifolia
and M. biacuta (the species perhaps doubtfully distinct) are
smaller (hypanthium to the torus 1.8-2.5 mm long; petals 1.5-2 X
1.2-1.7 mm), while M. floccosa (misplaced originally in Sect.
Miconia) has petals (fide Cogniaux) 5 mm long. Bang 2485 was
originally distributed as M. granulosa (Bonpl. ) Naud. (correctly
M. biappendiculata), which has quite different vegetative
pubescence, relatively wider leaf blades, and smaller flowers.

I have not seen the Bolivian collection (by d' Orbigny) cited by
Cogniaux for M. granulosa, but feel sure that the north
Colombian species does not really occur further south.

MICONIA DEMISSIFOLIA Wurdack, sp. nov.

Sect. Cremanium. M. caelatae (Bonpl.) DC. affinis, pubes-
centia compactiore antherarum thecis minoribus connectivo plus
prolongato differt.

Arbor parva vel frutex 2-5 m. Ramuli obtuse tetragoni
demum teretes sicut foliorum subtus venae primariae

4,90 Py Porn oe rk Vol. 23, no. 5

secundariaeque inflorescentiaque dense pilis pinoideis 0.1(-0.2)
mm longis (barbellis ca. 0.01-0.02 mm longis) induti. Petioli
1.5-2.5 (-3) cm longi; lamina (acumine excluso) 6-12 X 2-4(-5)
em apice longiuscule (0.7-1.8 em) subcaudato-acuminato basi
obtusa, coriacea et calloso-serrulata, supra glabra et obscure
rugulosa, subtus in venulis superficieque sparsiuscule pinoideo-
puberula, 3-nervata (pari O.5-1 mm inframarginali tenui neglecto)
nervis secundariis ca. 2-2.5 mm inter se distantibus subtus ele-
vatis nervulis subtus planis laxiuscule reticulatis (areolis ca.
0.7 mm latis) ob pilos occultis. Panicula 8-14 cm longa multi-
flora, ramis primariis oppositis; flores 5-meri, pedicellis ca.
1 mm longis, bracteolis ca. 1.2 X 0.3 mm valde caducis. Hypan-
thium (ad torum) 1.5 mm longum sparse stellulato-pinoideo-
puberulum; calycis tubus 0.2 mm altus, lobis interioribus 0.25
mm longis ovatis, dentibus exterioribus crassis lobos interiores
aequantibus. Petala glabra 1.3 X 0.9-1 mm obovato-suborbicu-
laria apice retuso. Stamina isomorphica glabra; filamenta 1.5-
1.6 mm longa; antherarum thecae 0.7 X 0.5 mm obovato-suborbicu-
lares late biporosae, connectivo ad basim 0.4-0.5 mm prolongato
0.25 mm lato. Stigma (in specimine florifero typico) 0.2 m
diam. non expansum; stylus 1.5 X 0.2 mm glaber vix exsertus in
ovarii apicem 0.3 mm immersus; ovarium 3-loculare et ca. 1/2
inferum, apice conico 0.6 mm alto sparsiuscule glandulis ca.

O.1 mm longis coronato.

Type Collection: M. T. Madison 10168 (holotype US 2626799;
isotype NA), collected in full sun in elfin forest near Camp Gy
Cordillera Vilcabamba across from Hacienda Luisiana, Prov.
Convencion, Depto. Cuzco, Peru, 12° 36? S, 73° 30’ W, elev.
3300-3535 m, 28 June 1970. "Tree 3-5 m; leaves stiffly pendant,
glossy green above, with dense ochre tomentum below. Calyx
green; other floral parts light yellow. Fruit deep violet.
Common."

Paratypes (both Peru): T. R. Dudley 11208 (NA, US), from
Camp 7, Cordillera Vilcabamba, Depto. Cuzco, elev. 3400 m; M. T.
Madison 10356 (NA, sterile), from near Puncu ca. 30 km NE of
Tambo on west side of Apurimac valley, Prov. La Mar, Depto.
Ayacucho, elev. ca. 3380 m.

The suggested Ecuadorian relative has stem hairs ca. 0.5 mm
long, lower leaf surface hairs with longer (0.1-0.15 mm) ena-
tions, pedicels ca. 0.2 mm long, and oblong anther thecae with
the connective only O.1 mm trilobulate at the base. (Among the
Ecuadorian collections [US] currently ascribed to M. caelata,
two taxa are involved, Camp E-4565 and E-4566 probably repre-
senting an undescribed species with the dominant underleaf hairs
quite different from those in the Jameson material). Probably
both species are functionally dioecious, only sporadic anthers
on the Jameson collections of M. caelata being well-formed (but
stigma expanded) and the stigma in buds of Dudley 11208 being
well-expanded (ca. 0.6 mm diam.). The holotype of M.
demissifolia has one fruiting sprig and one in flower (with
well-developed anthers and non-expanded stigma). Other more
northerly relatives of M. demissifolia include M. bracteolata

1972 Wurdack, Certamen Melastomataceis 491

(Bonpl.) DC. and M. biappendiculata (Naud.) Uribe, both with
more discrete foliar hairs (granulose in appearance, the enations
extremely short) and larger flowers with oblong anthers; M.
mutisiana Mef. differs at least in the entire leaf blades with
less dense, more compact, and caducous hairs on the lower
Surface. The general vegetative aspect of M. demissifolia is
like that of M. hygrophila Naud., which has much longer enations
on the lower leaf surface hairs ("cladonioid") and primary
inflorescence branches 4 per node, as well as shorter (0.2-0.3
mm) pedicels, but similar stamens.

MICONIA MICROPETALA Cogn.

Miconia torta Gleason, Phytologia 3: 349. 1950.

Cardenas 3964 (isotype US) fits within the variability
previously cited for M. micropetala (Phytologia 9: 421. 1964),
resembling Wurdack 1440 in the sparser development of simple
hairs on the lower leaf surfaces. A recent series of M.
micropetala from Cuzco (Cordillera Vilcabamba, 12° 35' S, 73°
35" W, elev. 2510-2568 m: Dudley 10695 and 10698; Madison
10150) almost completely lack simple leaf hairs (occasionally
a very few at the extreme blade base beneath). ‘The species
seems to be dioecious, the Jameson type (isotype US) as well
as Weberbauer 1145 being female with expanded stigmas and
abortive anthers; most (or all) of the other collections are
male, with well-developed anthers, unexpanded and non-exserted
stigmas, and abortive ovaries. A recent (male) Ecuadorian
collection is Maguire & Maguire 44367 (NY, US).

MICONIA RUGIFOLIA Triana

Cogniaux had placed this species in Sect. Amblyarrhena.
Actually M. rugifolia is very closely related to M. micropetala
(vide supra), differing in the bullate-rugose upper leaf
surfaces and slightly larger flowers (petals ca. 1.7-1.8 X
0.9-1 mm, rather than 0,.8-1.1 X 0.6-0.7 mm). ‘The two species
share the features of dioecism (Lechler 2196 [K] and Dudley
10705 being female), 2=-pored anthers, and ovary apices with
well-developed flanges to the hypanthial wall. Several recent
Cuzco collections (Pontiacolle, Vargas 19863; Cordillera
Vilcabamba, 12° 35 S, 73° 35 W, elev. 2510-2750 m, Dudley
10907 and 10705) have been referred to M. rugifolia, although
the leaf blades are relatively wider than in Lechler 2196;
here also perhaps belong Madison 10102 (Cordillera Vilcabamba,
Cuzco, elev. 1730 n, fruiting), with less densely bullate leaf
blades, and Wolfe 12427 (Cerros de Sira, Huanuco, elev. 1880 nm,
sterile).

MICONIA CAERULEA (D. Don) Naud. subsp. VILCABAMBAE Wurdack,
subsp. nov.

Ramulorum pili appressi ascendentes ca. 0.8 mm longi.
Foliorum superficies supra sparsissime strigulosae (pilis ca.
0.3-0.5 mm longis) subtus sparse strigulosae (pilis ca. 0.8-

1 mm longis). Inflorescentiae rami ramulique sparse appresso-

492 Pa PO OG a Vol. 23, noes

setulosi pilis ca. 0.6-0.8 mm longis. Calyx extus non setulo-
sus.

Type Collection: TT. R. Dudley 10421 (holotype US 2626776;
isotype NA), collected in montane rainforest between Camps 2 1/2
and 3 ca. 13 km NE from Hacienda Luisiana and Rfo Apurimac ,
Cordillera Vilcabamba, Prov. Convencion, Depto. Cuzco, Peru, 12°
35% S, 73° 35? W, elev. 1460-1730 m, 24 June 1968. "Shrub 6-30
ft., DBH 1/4-1 1/2 in. Flowers white and fragrant. Common."

Paratype (same general region): M. T. Madison 10125 (NA,
US), from below Camp 2 1/2, elev. ca. 1622 m.

The typical subspecies has branchlets and inflorescence
densely setulose with somewhat reflexed hairs 1-1.5 mm long,
leaf blades above sparsely to moderately loose-strigulose and
below moderately appressed-setulose, and calyx sparsely fine-
setulose at the toral zone outside. The type of M. caerulea is
female, a good recent match (male ) being Vargas 334 (Us) from
Amaibamba-Ceilan, Cuzco. Both collections of subsp. vilcabambae
are male. Killip & Smith 24758 (US), from San Ramon, Junin, is
Somewhat intermediate between subsp. caerulea and subsp.
vilcabambae, with the patent cauline and inflorescence pubescence
and lower leaf surface hairs of the typical subspecies but with
trichome density attenuated. The cauline and inflorescence
hairs of M. caerulea are smooth and in part caducously and
inconspicuously gland-tipped.

MICONIA DASYCLADA Wurdack, sp. nov.

Sect. Cremanium. M. caeruleae (D. Don) Naud. affinis,
ramulorum pilis sparse barbellatis folia subtus in superficie
glabra differt.

Ramuli sulcato-quadrangulati demum teretes densiuscule
pilis sparse barbellatis gracilibus paulo reflexis ca. 2 mm
longis p. p. inconspicue glanduliferis armati et glandulis
0.03-0.05 mm longis obsiti. Petioli 1-2(-3.5) cm longi; lamina
9-15(-18.5) X 3.5-7(-8) em oblongo-elliptica apice breviter
(1-2 em) gradatimque acuminato basi rotundata, membranacea et
integra, breviter (ca. 0.5 mm) distanterque appresso-ciliolata,
supra sparsissime strigulosa pilis 0.3-0.5 mm longis, subtus in
venis primariis secundariisque sparse pilis gracilibus laevibus
0.2-0.7 mm longis sparsiuscule induti in superficie glabra, 3-
nervata (pari exteriore inframarginali tenui neglecto) nervis
secundariis ca. 4 mm inter se distantibus venulis subtus planis
laxe reticulatis (areolis ca. 0.8 m latis). Panicula 8-11 X
7-13 cm multiflora, ramulis modice setulosis pilis laevibus vel
sparsissime barbellatis 1-2 mm longis p. p. glanduliferis;
flores 5-meri essentialiter sessiles (pedicellis ca. 0.2 m
longis) in ramulis interrupto-glomerulati, bracteolis ca. 0.8 mm
_longis subulatis persistentibus. Hypanthium (ad torum) 1.4-1.6
mm longum glandulis ca. 0.05 mm longis modice obsitum alioqui
glabrum vel sparsissime setulosum; calycis tubus 0.1 mm longus,
lobis interioribus 0.4 mm longis oblongis, dentibus exterioribus
acutis non eminentibus. Petala 0.8 X 0.6-0.7 mm obovata
minutissime granulosa. Flores masculini ignoti; stamina in

1972 Wurdack, Certamen Melastomataceis 493

floribus femineis sterilia, filamentis 1.1-1.2 mm longis glabris,
thecis 0.3-0.4 X 0.2-0.4 mm late obovatis, connectivo paullulo
(0.1 mm) prolongato non expanso. Stigma in floribus femineis
peltatum 0.8-0.9 mm diam.; stylus 3.3 X 0.3 m sparse pilis
glanduliferis 0.05-0.1 mm longis puberulus; ovarium 3-loculare
et 0.8 inferum, apice conico 0.2 mm alto sparse pilis 0.1 m
longis setuloso.

Type Collection: TT. R. Dudley 11829 (holotype US 2626773;
isotype NA), collected just below campsite at Huanhuachayo ca.
25 km SW of Hacienda Luisiana and Rfo Apurimac, eastern Massif
of Cordillera Central opposite Cordillera Vilcabamba, Prov. la
Mar, Depto. Ayacucho, Peru, 12° 43" S, 73° 50' W, elev. ca.

1570 m, 18 Aug. 1968. "Shrub or small tree 15-20 ft. Inflores-
cence branches pinkish.”

Paratypes: M. T. Madison 10097 (NA, US, in young bud) and
T. R. Dudley 10378 (NA, sterile), both from Camp 2 12,
Cordillera Vilcabamba, Prov. Convencion, Depto. Cuzco, Peru, ca.
12° 38° Ss, 73° 38" W, elev. ca. 1730 m.

Miconia caerulea has smooth trichomes throughout and leaves
beneath evenly appressed-setulose with hairs ca. 1 mm long.
Miconia dasyclada is rather a bridge to the species of Sect.
Cremanium with densely roughened pubescence and does not seem
intimately related to M. brachyanthera Triana nor M. peruviana
Cogn. From the description and type photograph, M. herzogii
Cogn. of Bolivia differs at least in the densely stellate-
furfuraceous branchlets (Gleason’s notes indicating pinoid hairs
ca. 0.3 mm long) and relatively broader acute-based leaf blades.
As seems true also in both M. caerulea and M. cyanocarpa Naud.,
M. dasyclada is dioecious, the type collection being female.

MICONIA POLYCHAETA Wurdack, sp. nov.

Sect. Cremanium. M. polygamae Cogn. affinis, foliis pro-
portionaliter angustioribus pilis subtus (venis primariis
exceptis) simplicibus bracteolis latioribus calycis lobis
distincte evolutis extus sparse setulosis differt.

Frutex vel arbor. Ramuli paulo nodosi obtuse sulcato-
quadrangulati demum teretes densiuscule glandulis ca. 0.05 m
longis induti; nodi sicut petioli foliorum subtus venae
primariae inflorescentiarum ramique dense pilis stipitato-
stellulatis (stipite 0.3-1 mm longo, radiis gracillimis) obsiti.
Petioli (1-)2-3(-4) em longi; lamina 7-18(-23) X 3-6(-9) cm
elliptico-oblonga apice acuminato basi obtusa vel rotundata,
membranacea et irregulariter ciliolato-serrulata, utrinque
sparsiuscule pilis laevibus gracilibus laxiusculis ca. 1 m
longis induta, (3-)5-nervata vel paulo (usque ad 0.4 cm)
plinervata nervis’ secundariis ca. 4-5 mm inter se distantibus
venulis subtus planis laxe reticulatis areolis ca. 0.8 mm latis.
Panicula ca. 8 cm longa multiflora; flores 5-meri subsessiles
(pedicellis ca. 0.2-0.3 mm longis), bracteolis ca. 1-1.2 X 0.2-
0.3 mm oblanceatis ciliolatis usque ad anthesim persistentibus.
Hypanthium (ad torum) ca. 1.5 mm longum extus sparse glandulis
minutis obsitum alioqui glabrum; calycis tubus 0.2 mm altus,

holy PHYTOLOGIA Vol. 23, no. 5

lobis interioribus 0.2-0.4 mm longis dentibus exterioribus
ecrassis sparse setulosis paullulo eminentibus. Petala 1.5 X
0.9-1 mm oblongo-obovata glabra. Stamina fertilia (in floribus
masculinis) paullulo dimorphica glabra; filamenta 2.4-2.5 mm
longa; thecae 1-1.1 mm longae 4-loculares late incompleteque

4 -porosae , connectivo 0.25-0.3 mm prolongato paullulo expanso.
Pistillum in floribus masculinis abortivum, stigmate puncti-
formi, stylo ca. 0.3 mm longo; ovarium fertile (in floribus
femineis ) 3-loculare, apice conico glabro, stylo glabro, stig-
mate expanso ca. 0.4 mm lato.

Type Collection: T. R. Dudley 10430 (holotype US 2626777,
male; isotype NA), collected in cloud forest between Camps 2 1/2
and 3 ca. 15 km NE from Hacienda Iuisiana and Rio Apurimac,
Cordillera Vilcabamba, Prov. Convencion, Depto. Cuzco, Peru,
12° 35" S, 73° 35% W, elev. 1730-2000 m, 24 June 1968.
"Multiple-trunked tree 10-40 ft., DBH 2-6 in.; leaf indument
red to brownish.”

Paratypes (all Peru): San Martin: R. Ferreyra 1024 (US;
male), from Margarita, Divisioria. Cuzco (all near-topotypical):
M. T. Madison 10184 (NA, US; female, in bud), from near Camp 2,
elev. 1565 m; Madison 10179 (NA; male) and 10112 (NA, US; in
very young bud), Dudley 10380 (NA; sterile) and 10626 (NA;
male), all from near Camp 2 1/2, elev. 1622-1730 m.

Miconia polygama has leaf blades ca. 1.5 (rather than 2.5-
3) times as long as wide with all the hairs beneath stipitate-
stellate and the simple hairs above denser and somewhat shorter,
setiform bracteoles, and barely evolved esetulose calyx lobes;
M. vitiflora Macbride has relatively wider leaf blades glabrous
above and with only roughened pubescence beneath, as well as
glabrous calyces. In anther form, M. polychaeta is like M.
paradisica Wurdack, which however has different vegetative
features. The general vegetative facies, but not the repro-
ductive features, of M. polychaeta is rather like that of M.
erioclada Triana (Sect. Miconia).

MICONIA THAMINANTHA Wurdack, sp. nov.

Sect. Cremanium. M. apricae Gleason affinis, foliis supra
essentialiter glabris differt.

Frutex vel arbor parva (1-)2-9(-18) m. Ramuli primum
obtuse quadrangulati demum teretes paulo nodosi in petiolorum
insertionibus ca. 0.5-1 mm callose elevati et caduce barbellato-
setulosi sicut petioli foliorum subtus venae primariae inflo-
rescentiaque pilis irregulariter stipitato-stellatis ca. 0.3-
0.5 mm longis laxissimis (ut videtur arachnoideis) sparsiuscule
vel modice puberuli. Petioli 0.8-1.5 cm longi; lamina (6-)8-
13 X (2)3-4(-5) cm oblongo-elliptica apice gradatim breviterque
acuminato basi rotundata, firme membranacea et integra vel
obscure serrulata, supra in superficie glabra (vel in zona 1 m
inframarginali sparse obscureque scabrido-strigulosa pilis
ecrassis ca. 0.1-0.2 mm longis), subtus in superficie glabra et
epunctata in venulis sparse glandulis 0.03 mm longis sparse
induta, 3-nervata (pari exteriore tenui inframarginali neglecto)

1972 Wurdack, Certamen Melastomataceis 95

nervis secundariis ca. 2-3 mm inter se distantibus nervulis
subtus planis areolis ca. 0.5 mm latis. Panicula (3.5-)6-9 cm
longa submultiflora, ramis primariis oppositis; flores 5-meri
subsessiles (pedicellis 0.2-0.5 mm longis) glomerati, bracteolis
ca. 1.2-1.5 X 0.3 mm oblongis ciliolatis persistentibus.
Hypanthium (ad torum) 1.3-1.5 mm longum sparse glandulis minutis
indutum alioqui glabrum; calycis tubus 0.1-0.2 mm altus, lobis
interioribus 0.4 mm longis semicircularibus, dentibus exteriori-
bus crassis non eminentibus interdum setulis 1-3 brevibus orna-
tis. Petala 1.3-1.4 X 0.8-1 mm obovato-oblonga retusa glabra.
Flores masculini: stamina paullulo dimorphica glabra; filamenta
2-2.1 mm longa; thecae 0.8-1 X 0.4 X 0.5 mm oblongae late (0.25-
0.35 mm) biporosae, connectivo ca. 0.2-0.25 mm prolongato
crasso; ovarium abortivum, cono ca. 0.4 mm alto, stylo 0.2 m
longo incluso, stigmate punctiformi. Flores feminei: stamina
sterilia, filamentis 1.5-1.7 mm longis, thecis ca. 0.7 X 0.2 X
0.2 mm, connectivo ca. 0.2 mm prolongato; stigma capitellatum
0.5 mm diam.; stylus 2.3 X 0.15 m glaber; ovarium 3-loculare

et 2/3 inferum, apice truncato-conico 0.4 mm alto glabro.

MICONIA THAMINANTHA Wurdack subsp. THAMINANTHA

Foliorum subtus venulae sparse pilis 0.2-0.4 mm longis
simplicibus indutae; foliorum cilia marginalia O.1 mm longa
eaduca, marginibus ut videtur minute crenulatis.

Type Collection: T. R. Dudley 10531 (holotype US 2626816,
male; isotype NA), collected in cloudforest just below Camp 3
ca. 15 km NE from Hacienda Luisiana and Rfo Apurimac, Cordillera
Vilcabamba, Prov. Convencién, Depto. Cuzco, 12° 30' S, 73° 30!
W, elev. ca. 2000 m, 26 June 1968. "Slender tree 10-30 ft.
tall; leaves brownish below. Flowers creamy white. Common and
weedy locally."

Paratypes (all near-topotypical): T. R. Dudley 10677 (NA,
fruiting), from Camp 2, elev. 1460 m; M. T. Madison 10186 (NA, male;
elev. 1892 m), T. R. Dudley 10341 (NA, US, female; elev. 1730-
2000 m), 10344 (NA, US, female; elev. 1730-2000 m), and 10541 (NA,
male; elev. ca. 2000 m), all from between Camps 2 1/2 and 3.

MICONIA THAMINANTHA subsp. HUANUCENSIS Wurdack, subsp. nov.

Foliorum subtus venulae esetulosae; foliorum cilia margi-
nalia 0.2-0.3 mm longa persistentia, marginibus ut videtur
serrulatis.

Type Collection: 1. R. Dudley 13187 (holotype US 2658939,
male), collected in dense cloud forest between Camp 3 and Camp
4, southwestern slope of the Rfo LlullaPichis watershed on the
ascent of the Cerros del Sira, Depto. Huanuco, Peru, 9° 26" §S,
7h° 4st W, elev. 1450 m, 23 July 1969. "Many-stemmed shrub
8 ft. tall. Flowers white.”

Paratypes (all near-topotypical): F. Wolfe 12368 (NA, US,
male?, in young bud), T. R. Dudley 13226 (NA, US, in very young
bud) and 13225 (NA, US, fruiting), all from near Camp 3, elev.
1280-1290 m.

Miconia aprica has leaf blades above sparsely asperous~

4,96 Py Heo D0) Die Or Girl, A Vol. 23, no. 5

strigulose with stout hairs, marginally more obviously ciliate,
and beneath sparsely punctulate on the actual surface, as well
as cauline hairs irregularly stipitate-stellate (and with a
more robust stalk) and more developed styles in the male flowers;
in other floral details, the two species are very similar.
Miconia peruviana Cogn., with leaves strigulose above, shorter
pinoid pubescence on the leaves beneath, and primary inflo-
rescence branches 4 per node, is another relative (and very
similar to M. aprica); a recent collection (female, in flower)
matching well Lechler 2066 (K) is Vargas 6872, from Ollachea,
Carabaya, Puno. Both M. aprica and M. peruviana are dioecious;
the type collection of M. aprica is male (as is also Soukup
3426), female collections (with abortive anthers and expanded
stigmas) being Ferreyra 3639, Killip & Smith 24316 and 24381,
all from Junin. Miconia galactantha Naud. (isotype FI) differs
at least in the more closely serrulate leaf blades which are
very sparsely scurfy on the veins beneath, as well as the
essentially glabrous inflorescences and setuliform (ca. 0.3 mm
long) early-caducous bracteoles. Miconia saxatilis Macbride
differs at least in the sharply quadrate branchlets and larger
plinerved basally acute leaf blades; M. coelestis (Don) Naud.,
M. crassistigma Cogn. (with longer pedicels), and M. brevistylis
Cogn. all have leaf blades puncticulate on the surface beneath
and less thickened anther connectives.

Miconia lugubris Cogn. (also, more distantly, M. atrofusca
Cogn.) is related to the M. aprica-M. thaminantha alliance, but
with smaller leaves and more abundantly setulose calyces. A
relative of M. lugubris (in young bud; with much sparser simple
hairs on both sides of the leaves) has been collected on the
summit (elev. 2200 m) of the Cerros del Sira in Hudnuco (Wolfe
12430, 12443); collections at anthesis are needed for descrip-
tion. Previously I referred Asplund 13125 (S, US), from Carpish,
Huanuco, to M. lugubris, but this collection essentially lacks
the dense simple hairs on the lower leaf surfaces and is at
least subspecifically distinct (but not the same as M.
thaminantha subsp. huanucensis).

MICONIA RUFIRAMEA Wurdack, sp. nov.

M. dumetosae Cogn. affinis, ramulorum pubescentia longiore
foliis proportionaliter latioribus ad basim rotundatis differt.

Ramuli obtuse tetragoni demum teretes sicut petioli inflo-
rescentiae axis densissime pilis minutissime denseque barbella-
tis 0.5-1.2 mm longis patentibus armati. Petioli 0.3-0.5 em
longi; lamina 1.8-2.7(-3.8) X 1.3-2(-2.7) cm, late elliptica
vel paullulo ovato-elliptica apice late acuto vel brevissime
(usque ad 0.3 cm) subabrupteque acuminato basi rotundata, rigide
coriacea (marginibus plerumque recurvis) et obscure distanterque
ciliolato-serrulata (ciliis crassis appressis 0.1-0.2 mm longis),
supra glabra et paulo rugosa, subtus in venis primariis sparse
caduceque pinoideo-furfuracea (pilis ca. 0.1-0.2 mm longis) et
in venulis superficieque sparsiuscule glandulis minutis 0.02 mm
longis obsita alioqui glabra, 3-nervata nervis secundariis ca.

1972 Wurdack, Certamen Melastomataceis 497

2 mm inter se distantibus sicut venulis supra insculptis
nervulis subtus planis areolis plerumque 0.6-0.9 mm latis.
Panicula 2.5-4 X 2-3 cm submultiflora, ramulis oppositis
sparsiuscule pinoideo-furfuraceis (pilis 0.1-0.2 mm longis);
flores 5-meri breviter (in fructibus 0.5-1 mm) pedicellati,
bracteolis 0.3-0.5 X 0.1 mm valde caducis. Hypanthium sparse
glandulis minutis obsitum alioqui glabrum. Petala immatura
minuta glabra. Antherarum thecae immaturae ca. 0.6-0.65 X O.4-
0.45 mm oblongo-ovoideae incomplete 4-loculares ut videtur
biporosae, connectivo paulo (immaturo ca. 0.2 m) prolongato.
Fructus ca. 2.5 X 2.5 mm, calycis lobis interioribus 0.4 m
longis, dentibus exterioribus crassis lobos interiores aequanti-
bus, ovarii apice glandulis paucis ornato.

Type Collection: 2. R. Dudley 10919 (holotype US 2626810,
in young bud, male?; isotype NA), collected just below Camp 5
ca. 23 km NE from Hacienda Iuisiana and Rfo Apurimac, Cordillera
Vilcabamba, Prov. Convencion, Depto. Cuzco, Peru, 126 35> 8,
73° 35' W, elev. ca. 2750 m, 10 July 1968. "Dense shrub to
8 ft.; leaves dark green, glossy above, cucullate; stem indument
rufous-tomentose. Flowers whitish yellow."

Paratypes (near-topotypical): M. T. Madison 10152 (NA,
US), from Camp 4, elev. ca. 2663 m (fruiting); T. R. Dudley
11143 (NA, US), from below Camp 6, elev. ca. 3000 m (fruiting).

Miconia dumetosa has rather scurfy-pinoid cauline pubes-
cence 0.1-0.3 mm long and acute-based leaf blades with length/
width ratio 2.2-2.9 (rather than 1.5-1.7). Vegetatively
M. rufiramea rather resembles M. secundifolia Cogn. subsp.
malcabalensis Wurdack, which has thicker clavate cauline hairs
0.2-0.3 mm long and much larger bisexual flowers with pinoid-
puberulous hypanthia.

MICONIA AYACUCHENSIS Wurdack, sp. nov.

Sect. Cremanium. M. cremophyllae Naud. affinis, foliorum
ciliis longioribus floribus maioribus differt.

Ramuli obtuse sulcato-quadrangulati demum teretes in nodis
linea interpetiolari paullulo (0.2 mm) elevata armati sicut
folia subtus inflorescentia hypanthiaque sparse glanduloso-
punctati alioqui glabri. Petioli 0.8-1(-2.5) cm longi; lamina
5-6.5(-9.5) X 2.2-3.3(-4.7) om elliptica vel oblongo-elliptica
apice acuto vel paulo (ad 0.5 em) subgradatimque acuminato basi
late acuta, coriacea et integra, conspicue appresso-ciliata
eciliis 1-2 m longis, 3-nervata nervis secundariis plerumque
3-4 mm inter se distantibus nervulis subtus planis laxiuscule
reticulatis (areolis ca. 1 mm latis). Panicula ca. 5 X 3 cm
submultiflora; flores 5-meri, pedicellis crassis 1.5-2 mm
longis, bracteolis ca. 0.7 X 0.1 mm valde caducis. Hypanthium
(ad torum) 3.7 mm longum; calycis tubus 0.7 mm altus, lobis
interioribus oblongo-triangularibus 1 mm longis, dentibus
exterioribus crassis inframarginalibus. Petala 2.5-2.6 X 2.2 m
obovato-oblonga extus glabra intus minutissime granulosa.
Stamina paullulo dimorphica glabra; filamenta 2.7-2.8 mm longa;
antherarum thecae 1.7-1.8 X 0.6 X 0.8 mm late biporosae;

498 Pie MOds/ OG Ta Vol. 23, no. 5

connectivum O.7-1 mm prolongatum ad basim dorsaliter dente
hebeti vel paullulo (0.15 mm ) bilobulato et ventraliter appen-
dice 0.2-0.3 mm bilobulato armatum. Stigma expansum 1 mm diam.;
stylus 4 X 0.65 mm glaber in ovarii apicem ca. 0.6 mm inmmersus;
ovarium 3-loculare et 2/3-3/4 inferum, apice conico 1 mm alto
sparsissime glandulis 0.1 mm longis coronato.

Type Collection: T. R. Dudley 11992 (holotype US 2658938),
collected above timberline above Punccu campsite ca. 50 km SW
from Hacienda Luisiana and Rfo Apurimac, eastern Massif of the
Cordillera Central opposite Cordillera Vilcabamba, Prov. La
Mar, Depto. Ayacucho, Peru, ca. 12° 45 S, 73° 53" W, elev.
3400-3600 m, 23 Aug. 1968. "Dense rounded shrub; calyx red;
petals white; fruit purple."

Paratype (near-topotypical): J. W. Terborgh 10381 (NA,
sterile), from elfin forest near Punccu, elev. 3380 m.

Miconia cremophylla has somewhat larger leaf blades with
margin ciliation only ca. 0.2 mm long, as well as considerably
smaller (hypanthium ca. 2 mm long, petals 1.8-2 X 1.6-1.8 mn,
anther thecae 1.3-1.5 mm long) subsessile flowers with barely
(0.1-0.15 mm ) appendaged anthers. Other already-described
relatives include M. tovarensis Cogn. and M. turgida Gleason
(cf. Phytologia 22: 409. 1972), as well as (more distantly)

M. coelestis (Don) Naud. and M. malatestae Macbride (both with
much larger leaf blades with shorter cilia).

MICONIA VARGASII Wurdack, sp. nove

Sect. Cremanium. M. ayacuchensi Wurdack affinis, floribus
minoribus calycis dentibus exterioribus eminentibus differt.

Ramuli obtuse sulcato-quadrangulati demum teretes in nodis
linea interpetiolari crassa 0.2-0.3 mm elevata armati sicut
folia subtus inflorescentia hypanthiaque sparse glanduloso-
punctata alioqui glabri. Petioli O.7-1.7 cm longi; lamina
4-9.5 X 1.8-4.5 em elliptica vel oblongo-elliptica apice hebeti-
acuto basi acuta vel obtusa, coriacea et integra, appresso-
ciliata ciliis 1-1.8 mm longis, 3-nervata nervis secundariis
plerumque 2-3 mm inter se distantibus nervulis subtus planis
laxiuscule reticulatis (areolis ca. 1 mm latis). Panicula 6-8 X
3-4 cm submultiflora; flores 5-meri, pedicellis 1.7-2.5 mm
longis, bracteolis ca. 1-2 X 0.1-0.3 mm fimbriato-ciliolatis
mox caducis. Hypanthium (ad torum) 3 mm longum; calycis tubus
0.5 mm altus, lobis interioribus 0.6-0.8 mm altis ovatis, denti-
bus exterioribus crassis 0.2-0.5 mm eminentibus. Petala 1.9-
2 X 1.3-1.4 mm obovato-oblonga ubique minutissime granulosa.
Stamina paulo dimorphica glabra; filamenta 1.8-2.2 mm longa;
antherarum thecae 1-1.1 X 0.4-0.45 X 0.5-0.6 mm paulo curvatae
late (0.35-0.4 mm) biporosae; connectivum 0.35-0.7 mm prolonga-
tum dorsaliter dente hebeti vel acuto et ventraliter appendice
paulo bilobulata armatum. Stigma capitellatum 1.2-1.3 mm diam.;
stylus 5 X 0.5 mm glaber in ovarii apicem 0.4-0.7 mm immersus;
ovarium 3-loculare et 2/3-3/4 inferum, apice conico 0.6-1 mm
alto sparse glandulis 0.1 mm longis coronato.

Type Collection: C. Vargas 2942 (holotype US 1830121),

1972 Wurdack, Certamen Melastomataceis 99

collected at "Puyupatamarca," Prov. Urubamba, Depto. Cuzco,
Peru, elev. 3400 m, 6 Aug. 1947. "Arbusto 3-4 m; pétalos
blancos."

Paratype: R. D. Metcalf 30774 (US), from near Wenner Gren
ruins, Prov. Urubamba, Depto. Cuzco, Peru, elev. 3400-3600 m.

Miconia ayacuchensis has considerably larger flowers with
inframarginal external calyx teeth, but similar leaf ciliation
and anther connectives. The other relatives discussed under
M. ayacuchensis (vide supra) all have shorter leaf ciliolation
and non-projecting external calyx teeth. Vargas 2942 was
distributed as M. alpina Cogn. and Metcalf 30 i as M. andina
Naud. (a synonym of M. latifolia [Don] Naud.); both these are
kemerous species of Sect. Chaenopleura. The type and paratype
localities for M. vargasii are in the group of Inca ruins just
south of Macchu Picchu (Fejos, P. Archaeological Exploration in
the Cordillera Vilcabamba southeastern Peru. Viking Fund Publ.
Anthrop. 3. 1944; Vargas, C. Homenaje Botanico a Machu Picchu.
Rev. Mus. Inst. Arqu. 19. 1961).

MICONIA CHORIOPHYLLA Wurdack, sp. nov.

Sect. Cremanium. M. brevistylae Cogn. affinis, foliis
tenuioribus petiolis gracillimis floribus bene pedicellatis
differt.

Ramuli teretes sicut folia inflorescentiaque primum sparse
vel modice resinoso-granulosi glabrati alioqui glabri. Petioli
1-2.5 cm longi, ca. 0.7-0.9 mm diam.; lamina oblongo-lanceata
vel oblongo-elliptica apice gradatim vel subgradatim per 1-1.5
(-2) cm acuminato basi acuta vel obtusa, membranacea et integra,
obscure (0.05 mm) distanterque appresso-ciliolata, 3-nervata
vel inconspicue (ad 0.2 cm) 3-plinervata nervis secundariis 2-3
mm inter se distantibus nervulis subtus planis laxiuscule retic-
ulatis (areolis ca. 0.5-0.7 mm latis). Panicula 3-12 X 3-9 cm
submultiflora vel multiflora, ramis oppositis divaricatis
quadrangularibus gracilibus; flores 5-meri unisexuales, pedi-
cellis 0.5-1(-1.5) mm longis juxta hypanthii basim vel paullulo
infra articulatis, bracteolis ca. 0.7 X 0.1-0.15 mm mox caducis.
Hypanthium (ad torum) 0.9-1.5 mm longum extus sparse granuloso-
resinosum; calycis tubus 0.15-0.2 mm longus, lobis interioribus
0.1-0.15(-0.2) mm altis inconspicuis, dentibus exterioribus
subobsoletis inframarginalibus. Petala 0.8-l1 X 0.7-0.9 mm
suborbicularia vel obovato-suborbicularia apicem versus paullulo
erosula inconspicue granulosa. Flores masculini: stamina
paullulo dimorphica glabra; filamenta 1-1.2 mm longa; thecae
0.6 X 0.35-0.45 mm obovato-oblongae late biporosae incomplete
4eloculares, connectivo 0.2-0.4 mm prolongato angusto; stigma
non expansum; stylus ca. 0.4 mm longus inclusus; ovarium aborti-
vum. Flores feminei: stamina abortiva, filamentis 1.2 m
longis, thecis 0.35-0.4 xX 0.2 mm, connectivo 0.3 mm prolongato;
stigma capitellatum 0.7-0.8 mm diam.; stylus 2.6 X 0.35 m
glaber exsertus in ovarii apicem 0.1-0.2 mm immersus; ovarium
3-loculare et 2/3 inferum, apice glabro alis 5 cum hypanthii
pariete connexo.

500 Pi XT 10.0 .G EA Vol. 23, noges

MICONIA CHORIOPHYLLA var. CHORIOPHYLLA

Foliorum lamina plerumque 8.5-14.5 X 1.7-3(-3.8) cm.
Linea interpetiolaris non evoluta.

Type Collection: JT. R. Dudley 10405 (holotype US 2626786,
male; isotype NA), collected in cloud forest at Camp 2 ca. 10 km
NE of Hacienda Iuisiana and Rfo Apurimac, Cordillera Vilcabamba,
Prov. Convencion, Depto. Cuzco, Peru, 12° 30' S, 73° 30? w,
elev. 1460 m, 24 June 1968. "Shrub 6-15 ft. tall; petioles and
venation on upper leaf surface reddish. Flowers white."

Paratypes (all Peru): Cuzco (near-topotypical): T. R.
Dudley 10433 (NA, US, male), M. 7. Madison 10191 (NA, sterile)
and 10192 (NA, male), all from near Camp 2; T. R. Dudley 10613
(NA, sterile) and 10388 (NA, sterile), Madison 10111 (NA,
sterile), all from Camp 2 1/2, elev. 1730 m; T. R. Dudle ———
(NA, US, male in bud), 10445 (NA, US, female in flower), 10540
(NA, fruiting), all from between Camps 2 1/2 and 3, elev. 1900-
2000 m. Ayacucho (Prov. la Mar, Cordillera Central opposite
Cordillera Vilcabamba): T. R. Dudley 11828 (NA, US, fruiting),
from Huanhuachayo, elev. 1570 m; T. R. Dudley 11918 (NA, male),
from between Huanhuachayo and Punccu, ca. 120 45" S50 (seep

MICONIA CHORIOPHYLLA var. BREVIFOLIA Wurdack, var. nov.

Foliorum lamina plerumque 4.5-6(-7) X 1.8-2.5 cm. Linea
interpetiolaris obscure evoluta.

Type Collection: T. R. Dudley 13520 (holotype US 2626791,
male; isotype NA), collected in open elfin forest between Camps
4 and 5, southwestern slope of Rio LlullaPichis watershed on
ascent of Cerros del Sira, Depto. Huanuco, Peru, 9° 25! S, 7h°
4k? W, elev. 1680 m, 31 July 1969. "Dense shrub ca. 6 ft. tall.
Inflorescence and flower buds deep purple."

Miconia brevistyla has firmer leaf blades with appressed
cilia 0.2-0.3 mm long and sparse inconspicuously barbellate
hairs 0.2-0.3 mm long basally along the costa beneath, as well
as subsessile (pedicels 0.1-0.2 mm long) flowers. ‘The type
collection of M. brevistyla is male and the species may yet
prove synonymous with M. coelestis (Don) Naud. (type collection
female). Another relative is M. elongata Cogn. (type collection
female), with quadrangular branchlets, much firmer and shortly
plinerved leaf blades on stouter petioles, somewhat larger
hypanthia, and more distinctly (ca. 0.3 mm) lobed calyx; to
M. elongata I have referred several recent Peruvian collections

Puno: Vargas 14825. Cuzco: Vargas 6235 and 14914). Cogniaux
originally placed M. elongata in Sect. Amblyarrhena, but the
type clearly indicates placement with other dioecious species
of Sect. Cremanium. Miconia choriophylla also generally
resembles M. glaberrima (Schlecht.) Triana var. australis
Macbride, which differs from both the new varieties at least in
the longer furfuraceous indument along the primary leaf veins
beneath (paratype US, with galled inflorescences); I doubt that
Macbride's variety is conspecific with the Central American
species and indeed the La Merced population may well be includ-
able in M. choriophylla (subspecifically distinct), but detailed

1972 Wurdack, Certamen Melastomataceis 501

examination of Macbride 5769 is needed. One somewhat anomalous
collection (Dudley 10457, female; material incomplete) has not
been included in the paratypes of M. choriophylla, having foliage
as in var. choriophylla (and collected at the same locality as
Dudley 10445) but with interpetiolar cauline lines developed and
calyx more lobed than typical.

MICONIA LACHNOCLADA Wurdack, sp. nov.

M. bullatae (Turcz.) Triana affinis, foliis maioribus
distincte 5-nervatis inflorescentia multiflora filamentis gla-
bris ovarii apice glabro differt.

Ramuli sicut petioli (apicem versus) pilis patentibus
minutissime barbellatis 2-3 mm longis sparse induti et sicut
petioli inflorescentiae pedunculusque pilis dense barbellatis
1-1.5 mm longis patentibus dense armati. Petioli (0.4-)0.7-1.2
em longi; lamina (2.2-)3-5.7 X (1.3-)2=3.7 em ovato-oblonga vel
elliptica apice late acuto basi late obtusa, rigida et minute
serrulata (dentibus ca. 0.2 mm calloso-ciliolatis), supra
bullata bullis mucro crasso ca. 0.2 mm longo armatis alioqui
glabra, subtus in venis primariis secundariisque modice pilis
parbellatis crispulis ca. 1 mm longis et in venis primariis
densiuscule pilis pinoideis ca. 0.2 mm longis induta, 5-nervata
nervis secundariis ca. 2 mm inter se distantibus nervulis subtus
planis densiuscule reticulatis (areolis ca. 0.3 mm latis).
Panicula 3-5 cm longa multiflora terminalis, ramulis modice
pilis pinoideis ca. 0.3 mm longis armatis; flores hemeri, pedi-
cellis 1.2-1.7 mm longis, bracteolis angustis ca. 0.5 mm longis
mox caducis. Hypanthium (ad torum) 2.5 mm longum extus sparse
stellulato-puberulum et minutissime glandulosum; calycis tubus
0.2-0.3 mm altus, limbo paullulo (0.35 mm) 4-undulato, dentibus
exterioribus crassis non eminentibus. Petala 1.7-1.8 X 2.3 m
late obovata paulo retusa minute granulosa. Stamina isomorphica
glabra; filamenta 1.9-2 mm longa, basim versus 0.8 m lata;
thecae 1.5 X 1.1 X 0.8 mm obovatae late biporosae et 0.3 m
emarginatae, connectivo ad apicem 0.1-0.2 mm eminenti ad basim
dorsaliter 0.15 mm hebeti-dentato ventraliter paulo (0.15 mm )
bilobulato-prolongato. Stigma capitatum 0.9 mm diam., 0.5 mm
altum; stylus 2.3 X 0.5 mm glaber in ovarii apicem ca. 0.2 m
inmersus; ovarium 4-loculare et 0.8 inferum, apice truncato-
conico 0.4 mm alto glabro.

Type Collection: M. T. Madison 10378 (holotype US 2626800;
isotype NA), collected in full sun along trail near Cusimachay
cae 25 km NE of Tambo on west BLOpe of Rfo Apurdmac valley, Prov.
La Mar, Depto. Ayacucho, Peru, 12° 49" S, 73” 50" W, elev. ca.
3587 m, 28 July 1970. "Tree if m tall, with dense crown; leaves
stiff, dull dark green, rugose, with scarlet-tipped hairs below.
Calyx light green; corolla and stamens pale lemony-white."

Miconia bullata has 3-nerved leaf blades only up to 2.4 X
1.8 om (usually smaller) with short pee? O.1 mm long) sparse pin-
oid has on the primary veins beneath (as well as longer simple
hairs), inflorescences mostly 3-5-flowered, glabrous (except for
the sparse glands) hypanthia, sparsely (especially apically)

502 PRY TO ro £5 Vol. 23, now

glandular filaments, and sparsely glandular-setulose ovary
apices; apart from the collections listed in the synonymization
of M. trichocaula Macbride (Phytologia 11: 395. 1965), M.
bullata is now known from Huancavelica, Peru (iisengealnae
Marcavalla, elev. 3000-3200 m, 0. Tovar O46). Miconia
miles-morganii Macbride is more distantly related, having only
short (0. 1-0.2 mm long) pinoid vegetative pubescence, longer and
relatively narrower anthers, and sparsely glandular-setulose
ovary apices (but 4 -nerous flowers, despite the original
description). ‘The type Soltection: of M. miles-morganii (isotype
US) shows styles sparsely glandular-puberulous as well as
stellulate=puberulous; a more northern ,population of this
species a & Wright 5595 and Lopez, Sagastegui &
Collantes 4438, both from Cerros Calla-Calla, Amazonas) shows
styles without stellulate hairs (very sparsely glandular), but
otherwise does not differ from the Hudnuco material.

BOOK REVIEWS

Alma L. Moldenke

"BEEKEEPING - THE GENTLE CRAFT" by John F. Adams, 182 pp., illus.,
Doubleday & Co., Garden City, New York 11530. $5.95.

This is so enthusiastically and delightfully written a guide
for the amateur beekeeper that non-bee-raising readers like my-
self find or will find themselves musing about watching these
fascinating creatures at their highly diversified labors and
about enjoying their delicious product first-hand. For the
apiary hobbyist there is much valuable guidance in this book.
There is a useful bibliography that is interestingly annotated.

"PHOTOSYNTHESIS : METHODS IN ENZYMOLOGY Volume XXIII Part A"
edited by Anthony San Pietro with editors-in-chief Sidney P.
Colowick & Nathan O. Kaplan, xix & 743 pp., illus., Academic
Press, London & New York, N. Y. 10003. 1971. $29.50.

The editor's short preface gives a good idea of the nature of
this book: "The discovery of the chloroplast reaction by Hill in
1937, commonly known as the Hill reaction, heralded the concerted
enzymological attack on the mechanism of photosynthesis. In the
ensuing three and a half decades, a tremendous but diverse wealth
of knowledge has accumulated from the multiplicity of experimen-
tal approaches used to study this problem. The aim of this (and
the succeeding) volume is to provide as comprehensive as possible
a coverage of these methodological approaches, namely, biochemi-
cal, biophysical, genetic, and physiological.

"The presentations in this volume consider isolation and cul-

1972 Moldenke, Book reviews 503

ture techniques of algae, bacteria, and diatoms; plant tissue cul-
ture; the preparation and properties of mutants; cellular and sub-
cellular preparations from algae, bacteria, and plants; and the
purification and properties of components of the photosynthetic
systems."

All this is covered with amazing thoroughness in the sixty-nine
component papers, each with its own bibliography. Many new ques-
tions are asked and answered, and the inevitable still newer ones
recognized and often analyzed. The whole work is far removed from
that which is simple, yet there is clarity in the authors' explan-
ations that makes for reasonable comprehension.

The detailed author and subject indexes as well as the table
of contents make access to the material facile. What a storehouse
of information lies within the covers of this book!

Index to authors in Volume Twenty-three

Barkley, F.A., 323, 327 Moldenke, A. L., 197, 250, 316,
Boghdan, K. S., 323, 327 502

Boivin, B., 1 Moldenke, H. N., 180, 181, 210,
Cuatrecasas, J., 350 212, 257, 314, 366, 413, 453,
Feddema, C., 201 460

Gibson, D. N., 33h Ogimi, C., 155

Griffin, D., {52 Reveal, J. Le, 163

Hale, M. E., dro, 179, 343 Robinson, H., 149, 151, 153, 305,
Titis, H. H., 28 307, 310, 312, 390, 393, 395,

King, Re M., 151, 153, 305, Se bs 05, 09
310, 312, 393, 395, 397, 405, Rogers, C. M., 39

ho9 Rudd, V. E., 11, 321, 402
Rome hs Po, 155 Seymour, F. C., 40
Mildner, R., 39 Sieren, D. J., 30)

Moir, W. W. G., 455 Wurdack, J. J., 47h

Index to supra-specific scientific names in Volume Twenty-three

Abies, 158, 228, 297 Aecidium, 292, 293

Acacia, 187, 190, 221, 302 Aegiphila, 31), 31 an
Acantholippia, 433 ean 127. Ae) Ne ee
Aceraceae, 255 Aegiphyla, 27

Achillea, 31, 81-83, 87, 138 Aeschynomene, 321

Actinella, 80, 133 Agalinis, 134, 138

Actinidia, 67 Agastache, 138

Adenocaulon, 7, 65, 138 Agave, 187

Adiantum, 6, 50 Ageratina, 153
Adoxa, 138 Agnus, 27

50) PE Y TOP vOrGeren Vol. 23, no. 5
Agoseris, 9, 118, 119, 126, Austroeupatorium, 393

27S L315. 150 Avicennia, 16, 17, 19, 421,
Agrianthus, 312 ~ 422, 425, 427
Agrostis, 237, 264 Avicenniaceae, 197, 317, son
Aira, 287 Bahia, h, 79, 138
Ajuga, 37 Bahianthus, 312, 313
Aldama, 205 Balisaea, 321, 322
Allocarya, 138 Balsamorhiza, h, 70, 71, 138
Alloplectus, 334-336 Barrosoa, 307, 312

Alnus, 18), 228

Aloysia, 387. N19, 22,825,
eT a 433, 437

Amaranthaceae, 255

Amaranthus, he

Amaryllis, 8

Amasonia, 18, 31

Amblyarrhena, 76, 477, 479,

i680, 482, 483, 185-187, 192

Ambrosia, eh 66, 67, 138, 22

Amphigymnia, 3hh, 36

Amsinckia, 138

Anaphalis, 7, 63, 138

Anaplanthus, 138

Androcera, 138

Antennaria, 6, 7, 55--63, 138

Anthemis, 3, 4, 81, 83-85, 138

Anthephora, 50

Anthopogon, 138

Antigonon, ))3

Aplopappus, 22, 23, 128

Arbutus, 328, 302

(eeten, Gl Glas wer 18

Arnica, 2, 6, 93-99, 138

Artemisia, 8, 87-91, 138

Arundo, 216

Asclepias, 264, 50

Ascomycetes, 263

Ashmeadiella, 286

Asperugo, 136

Aster, (Zo le 1h, 22 2h-h43, h6,

126, 138, 26h
Asteraceae, 151, 153, 305-307,
Foo 393, 395-397, 05,
09
Astragalus, 17), 235

Bartsia, 138

Begonia, 323-325, 327-333
Begoniaceae, 323, 330
Begoniastrum, 331
Benthamidia, 67

Besseya, 138

Betulaceae, 255
Bicornuta, 33

Bidens, 3, 6, 75-77, 138
Bifusella, 155-162
Bifusepta, 156, 161, 162
Boehmeria, 26)

Boltonia, 3, 2h, 138
Bombus, 286

Borago, 138

Borreria, )))
Boschniakia, 138, 67
Botrytis, 27)

Bouchea, 210, 318, 14, 416, 25
Brassavola, ))8

Brassicaceae, 213, 436, h7i2
Brickelia, a £2, 1138
Bromeliaceae, hho, hs

Bromus, 7

Bryophyta, 19

Buchloé, 65, 138

Burmannia, )9

Bursera, 232

Butia, 215

Cadiae, 01

Caesalpiniaceae, 01

Calamagrostis, 26)

Calendula, 5, 108, 109, 138

Callicarpa, 210, 318, lh-l16,
2h, 425, 27, 28

Calliopsis, 26)

1972 Index 505
Campanula, 138 Clematis, 6
Cardiospermum, 2);2 Clerodendrin, 28

Carduaceae, 213, 36, 473
Carduus, 5, 111-116, 139
Carex, 26)

Cartholinum, 138

cargopteris, 422-25, 28, 29,
438,
Au36

Casselia,
Castilleja, 135, 138
Catopsis, Catopsis, lS, ins
Cattleya, 150
Cecidomyia, 367

Cellicarpa, 28
Cenchrus, 235

Centaurea, 5, 7, 8, 116, 117,
138
Cephaelis, 6, 49, 450
Cephalocereus, 18)
Cerastium, 225
Ceratina, 286
Ceratoides, 16)
Chaenactis, 80, 138
Chaenopleura, li99
Chaenorrhimm, 138
Chamaesaracha, 138
Chamomilla, 6, 138
Cheilothela, 390, 392
Chelone, 138
Chelostoma, 286

Chelostomopsis, 286
Chenopodiaceae, 255

Chenopodium, 118, 317
Chondrophylia, 138

Chromolaena, 09
Chrysanthemum, 3, 8, 85, 86, 138

Ghrysopsis, 2, 1h, 21, 138
Chrysothanmus, bo 2h, "138

Cibotium, 317

Cichorium, 8, 117, 138
Cirsium, 5, 109, 111, 116, 138
Citahrexylun, 4,28
Citharexylum, 180, 318, 1h,

415, 417, 418 1,28
Clandularia, 1,26

Clerodendron, 428, 29, 38
Clerodendrum, 180, 199, 315, 318,
pe i, 127, 120-1126, i,28-

Pec a
Climaciacese, 150
Cmelina, 130
Cnicus, 11), 138
Cnidoscolus, 153
Coceocypselum, L7

Coelocentrum, 331
Coelomycetes, 253

Coix, 6

Collandra, 138

Collinsia, 138

Collomia, 138

Colocasia, 3L7

Columnea, 33h, 335
Compositae, 1-9, 140, 201, 255,
304, 3 396, 12
Congea, 1,23, 2h, 426, 430
Congeae, 130

Connatae,

coor Ph 307-309
Conoclinium, 307, 308

Conopholis, 138

Convolvulus, 138
Conyza, 54, 138
Cordia, 226
Coreopsis, 3, 75, 138

Cornus, 226, 295
Cornutia, 319, 41h, 415, 430,

45h
Cota, 81, 138
Cowania, 235

Cremaniu, 481, 489, 492-l9h,
~ 497-500

Crepis, 9, 118, 119, 127-129,
138

Critonia, 05

Critoniopsis, 358

Cronquistianthus, 09-11

Cruciferae, 301, 48

Cryphaea, 149, 150

Cryphaeaceae, 150

506

Cryphidium, 19
Cryptantha, a

Cunninghami » 155, 156, 158-162

Cuphea, Ta

Cupressineae, 2
Cupressus, 187
Cuscuta, 133, 1

Cyclachaena, 66,

Cyclolobium, 32
Cynoglossum, 13

Cyperaceae, 161, WhO, hh5, bh6

PHY T OL Oxygen Vol. 23, no. 5

, 150

55

38
138
1

8

Cyperus, Alley lil

Cyptodon, 119
Dasystephana, 1
Datura, 13
Daucus, 265
Decachaeta, 397
Dendrocryphaea,

38

149

Desmodium, 10, 42, bh)

Dianthera, ))9
Dichronema, ),50
Dicranella, 390

Dicrastylidaceae, Wis BLT LB

113
Dimorpholepis,
Diototheca, 730

Diplopappus, ez,
Diplostephium,

305, 306

138
350-353

Doellingeria, 21, 1, 138

Downingia, 138

Dracocephalum, 138

Drosera, ),)9
Drymaria, 226
Drymontia, 335,
Dudleya, 226

Duplicaria, 156,

Duranata, 30

Duranta, 319, 413, 15-17, 426,
nan

30,

336

161

Echinacea, By 0,130

Echinops, 6, 10
Echium, 138
Eichornia, )),2
Elaeagnon, ),30

Eleocharis, 5,

Ellisia, 138

9, 138

Lhé, Lg

Elscholtzia, 137

Elsholtzia, 137, 138

Emilia, )))6

Epeolus, 286

Epidendrum, 8, 50

Equisetum, 251

Eragrostis, 6

Erechtites, 93, 138

Erigeron, 1, 2, 7, 3-55, 138,
265

Erimus, 282, 30

Erinus, 318

Eriocaulaceae, 197, 317, 318,113

Eriocaulon, line lh, 116-128,
eee? 42h, 425, 430, 432,

Eriodictyon, 235

Eriogonum, 163-178

Erodium, 138

Eryngium, 109, 138, 450
Erysiphe, 2655 27h, wale 298,471
Erythrina, 237

Espeletia, 350, 358, 360, 362,
ee

Euchlaena, 218, 29

Eulonchus, 286

Eupatorieae, 151, 153, 305-307,
309-312, 393, 395-397, 05,
09, a2

Eupatorium, 6, 10-12, 138, 152,
15h, 26h, 305-308, 311-313,
393-395, 405, penta:

Euphrasia, 138, 6 2

Eurotia, 16
Buthamia, 30h,
Fabaceae, )01
Fagaceae, 255
Faradaya, 25
Feramea, )27
Fleischmannia, 05, 06
Fontinalaceas, 150
Franseria, 67, 138
Fucaceae, },31
Fuirena, ),))

Fungi Imperfecti, 253
Gaillardia, 35h, °80, «138, «235

Galeopsis, 138

L972

Galinsoga, 3, 77, 78, 138
Gentiana, 138
Gentianella, 138
Geranium, 136, 138, 370
Gerardia, 13, 138
Geron, 286

Gesnera, 31

Gesneria, 131
Gesneriaceae, 33)

Geum, 26)

Ghinia, 16, 431

Gilia, 138

Gireoudia, 331

Index

507

Heliconia, 45
Heliopsis, 3, 68, 139

Heliotropium, 139
Helipterun, 305
Helogyne, 09
Heriades, 286
Hesperapis, 286
Heterodera, 27)
Heuchera, 226
Hieracium, 9, 10, 130-132, 137,
139
Hierobotana, 370
Hillebrandia, 323

Glandularia, 182, 186, 211, 222, Hippia, 432

225-227, 234, 239, 2hh, 278,
279, 299-301, 303, 370, 372,
3135°h31
Glechoma, 138
Glondularia, 195
Glossocarya, 23, 32
Glossocentrum, },88
Glycine, 12
Gmelina, 319, 417, 422-26, 432
Gnaphalium, 7, 55, 64, 65, 138
Goyazia, 322
Gramineae, 2)8
Gratiola, 138, 387
Grazielia, 305
Grindelia, ), 13, 1h, 138
Gutierrezia, , 1h, 138
Gymnopsis, 201, 205
Gyptidium, 310, 311
Gyptis, 312, 313
Habenaria, 8
Hackelia, 138
Halenia, 138
Haplopappus, 2, 22-2, 42, 138
Hastingia, 32
Hatschbachiella, 393, 39h
Hebeclinium, 151, 05-08
Hedeoma, 139

Hedwigiaceae, 150

Helenium, , 80, 81, 139
Heliantheae, 201

Helianthus, ), 68, 70-75, 80,

81, 139, 26

Hippodamia, 286
Hoffmannia, 27

Holmskioldia, 16, 17, 423, 425,
26, 4432
Holmsktldia, 32
Homopappus , 23, 239
Hookeriaceae, 150
Hoplitis, 286
Hydrocharitaceae, 317
Hydrophyllum, 139
Hygrophila, 46
Hylaeus, 286
Hymenachne, 3
Hymenofyranus, 32
Hymenopappus, 8, 78, 139
Hymenopyramis, 432
Hymenospytanis, 32
Hymenoxis, 79, 139
Hymenoxys, i
Hyoscyamus, 139
Hypericum, 287
Hypochaeris, 9, 119, 139
Hypoderma, 158, 160, 162
Hypodermataceae, 155, 162

Hypodermateae, 155, 161
otrachyna, 3h)
Ryptis, kh, 4h7-Lh9
Hyssopus, 139, 387
Hysteriales, 162
Imbricaria, 179, 3h)
Impatiens, 139, 26h
Indigofera, 302

508 PHY TO L:0°G 1k Vol. 23, no. 5

Ipomoea, 232, 317 Leveillula, 71
Isthmiella, 158, 162 Liatris, 6, 12, 13, 139
Itonida, 367 Lichenes, 179, ” 31,3

Iva, 5, 7, 65, 66, 139 Limosella, 135, 139
Jacquinella, Ls Linaceae, )39
Jacquinia, },)2 Linanthus, 139
Juglandaceae, 255 Linaria, 139

Juncaceae, 161 Linum, 139, 39
Junellia, 194, 270, 18, 32, Lipocarpha, ))6

De aGes Lippia, 318, 413-18, 420, 421,
Juniperus, 235, 302 426, 432, 433
Kalaharia, 210, 21 Liquidambar, 225
Knautia, 139 Lithospermum, 136, 139
Knesebeckia, 331 Litsea, )33
Koanophyllon, 395, 396 Tiaees, 225

Kohleria, 336, 337 Lobelia, 139, 226
Krigia, 9, 119, 139 Lomatogonium, 139
Kuhnia, 312, 313 Lophiosphaera, 160, 162
Kyllingia, 6 Lophodermium, 160, 162
Laciniaria, 13, 139 Lourteigia, 307, 309, 09, 412
Lactuca, 9, 123-125, 130, 139 Luehea, 433
Lallemantia, 139 Lycium » 139

Lamiaceae, 37 Iycopodiacese, 251
Lamium, 139 lycopsis, 139

Lantana, 180, 210, 319, 387, Lycopus, 139, 36)
Li3-h21, 123, 126, 32, Loh lygodesmia, 10, 120, 125, 126,

Lappula, 139 139
Lapsana, 8, 118, 139 lysiloma, 232
Larrea, 187, 270 Lythrum, 26)
Leersia, 26) Machaeranthera, L, 6," Toneeene
Leguminosae, 141, 187, 321, 01 Macrophomina, 27h.
Leiophron, 265 Madia, ], 7, 78, 139
Leiothrix, 18, 4S Wangifera, 43
Leochilus, ))8 Marrubium, 139
Leontodon, 128, 139 Matricaria, 3, 8, 81, 83-85, 139
LeOMUrDs, a9. 167 Maurandya, 226

Lepac Lepachys, 70, 139 Megachile, 286
Lepidanthrax, 286 Megalodonta, 77, 139
Lepidiun, Te Melampyrum, 139
Leptilon, 54, 139 Tee eeeee oT

elanco es,

weptodens 150 Meliola, 22)

Leptolobium, 03
Lestodiplosis, 367
Leucanthemum, 85, 139
Leucodontaceae, 150
Leucophysalis, 139

Melissa, 139
Melissodes, 286
Meloidogyne, 27)
Mentha, 137, 139

1972 Index 509

Menyanthes, 139 Odontites, 139
Mertensia, 136, 139 Oenothera, 265
Mesanthemun, 13) Oidium, 274, 283, 293, 471
Metazoa, 251 Oleaceae, 255
Miconia, 7-502 Oligoneuron, 19, 139
Wicronycetes, 162 Olyra, 46
Microseris, 8, 9, 118, 126, 139 Oncidium, 451, 455-459
Microsphaera, 27) Onosmodium, 139
Mikania, 312, 313 Ophryosporus, 397-00, 09
Mimosa, 238, 302 Oplismems, 6
Mimulus, 139 Opun untia, 187, 192, 238, 2h1, 302
Mirabilis, 235 Orehidacese, se 40, L4s
Moldavica, 139 Oreocarya,
Molinadendron, 295 mie “350, 354, 356
Wonarda, 139 Orobanche, 136, 139
Moraceae, 255 Orthocarpus, 139
Moussonia, 336 SHyzOpSES, 16H
—— strya
Yusei, 119, 150 Pachythamnus, 153, 154
lyosotis, 139 Padus, 184, 226

carpus, )01-l0) Paeonia, 323
iyrospermm, 1,01 Paepalanthus, 181, 211, 417, 418,

lon, 01 434

Panicum, 45, 448, 50
My hiza, 1
Sse ra =H om Parmelia, 179, 343-39
ee - Pedicularis, 135, 139
Napeanthus 337 pte eet

‘ Pennisetum, 50

Navarretia, 139 she
Neckera, 149, 150 Penstemon, 133, 139, 17h, 225,

2
Hemophilia, 239 ctretatta) be
Less anon hone Mh So Peperomia 8
Neocabreria, 151, 152 Ere ok. 29°
Neogreenella, 153 Petasites, 6, 65, 71, 91, 92, 139
Nepeta, 139 Petraea, 318, 13h
Niphaea, 337 Petraeovitex, 43h
Nonea, 139 Petraevitex, 43)
Nothocalais, 118, 139 Petrea, 318, 319, 415, 416, 418,
Nothofagus, 179 26, “13h
Notyiias Lh6 Petunia, 318, 43h

ctanthaceae, 187, 317, 318, Phacelia, 139

a, 43 Phlox, 139, 26h
Nyctanth[acleae, 3h Phoradendron, 302, lj
Nyctantheae, 3) Phryma, 2)2
Obletia, 43h Phycioides, 286

Odonites, 68 Phyla, 319, 413-16, L19, 422,

510 PnY 10 i) OfGer ak

423, 426, 430, 432-L3h, 436,
4,38
Phylla, 43h
Phyllosticta, 265
Phymatotrichum, 27)
Physalis, 139
Physostegia, 139
Picea, 158
Picradeniopsis, 79, 139
Picris, 9, 119, 120, 139
Pinguicula, 139
Pinnatella, 150
Pinus, 158,
297, 302, 375, hh9, L70
Piqueria, 326
Pittosporum LT
Plagiobotrys, 139
Plantaginaceae, 255
Plantago, 139
Platanaceae, 255
Platycentrum, 331
Pleurocarpi, 150
Pleurogyne, 139
Pohlia, 52
Poiretia, 141-13, 145-148
Pelangstn! 139
Polites, 286
Polygonaceae, 163, 178, 255
Polypodium, 6, 451
Potentilla, 78
Premna, 20-23, 425, 126, 3k
Prenanthes, TO, E291 30), 139
Prescottia, mY
Prionodontaceae, 150
Pritzelia, 331
Priva, 415, 116
Proboscidea, 139
Prosopis, Prosopis, 187, 221, 21, 302
Prunella, 139
Psilocarphus, Ge OS, 1359
Psilotum, 251
Psoralea, 16
Pteridophyta, ))2
Puccinia, 188, 265, 293
Pycnanthemum, 26),

Pyrethrum, 6h, 86, 139

167, 18h, 228, 295,

Pyrrocoma, 23, 2h, 139

Quercus, 18), 187, 225, 226, 228,

232, 295, 302, 375
Ratibida, i, 10,239
Reichenheimia, 331
Reseda, 226
Rhacocarpeae, 150
Rhinanthus, 139
Rhizoctonia, 274
Rhus, 237
Rhynchospora, hus, 449, 450
Rhytisma, 162
Rhytismataceae, 155
Romanschultzia, 225
Romanzoffia, 139
Rubiaceae, 27
Rudbeckia, 4, 69, 70, 139
Ruta, 307.

Sucittaria! hs
Salicaceae, 255
Salix, 43h, lh3
Salvia, 139
Sargassum, 31
Saussurea, 6, 110, 111, 139
Scaphyglottis, )h5
Schizaea, 251
Scirpus, 26)

Scleria, 7-50
Sclerocarpus, 201-209
Sclerotium, 27)

Scrophularia, 139
Scrophulariaceae, 36, 437

Scutellaria, 139

Sezegleewia, 3)

Senecio, 2, 6, 91, 93, 99-108,
139, 153, 169, 213, 436, Uiz3

Septoria, 265, 293, {71

Sherardia, 3)

Sideranthus, 22, 23, 2, 139

Silybum, 5, 116, 139

Sisymbrium, 213, 436, 472, 73

Smilax, tho

Solanum, 10, 2h2

Soleslia, 158, 162

Balesecnera. 337-31

Vol. 23, no. 5

72

Solidago, 2, 14-22, 26, 99, 10,

265, 304
Sonchus, 9, 122, 123, 10
Sophoreae, 4,01
Spartina, 22, 26),

Spartothamnella, 435
Sphaerioideae, 253
Sphaeropsidales, 253
Sphaerotheca, 27),

Sphenodesma, 35, 1,38
Sphenodesme, 23, 42h, 430, 35,

38
Spiridentaceae, 150
Stachys, 10
Stachytarphaeta, 35

Stachytarpheta, 15-13, 21,
423, 25, 435, h69

Stenotus, 23, 10

Stephanomeria, 9, 120, 10

Stilbaceae, 197, 317, 318, 413

Stilbe, 35

Stomatanthes, 393

Stygnanthe, 33)

Stylodon, 18)

Stylotrichium, 212

Sympharema, 35

Symphorema, 133-135

Symphoremaceae, 197, 317, 318,

413
Symphorena, 35
Symphyopappus, 312, 313
Symphytum, 1h0
Syngonanthus, 417, 418, 435
Syrrhopodon, 390-392

Tanacetum, 8, 86, 87, 10

Index

511

Thelesperma, 6, 76, 10

Thielaviopsis, 27)
Tiliaceae, 33

Timotecia, 35

Timotocia, 318

Tithonia, 42

Tonella, 10

Townsendia, 2, 3, 2h, 25, 10,
17

Tragopogon, 9, 120, 121, 10

Trimorphaea, 52, 53

Tripsacum, 218

Trychinolepis, 397-399
Tuberostyles, 06

Turpinia, lhl, 142

Typha, 26)

Ulmaceae, 255

Umbelliferae, 50

Unamia, 21, 10

Urbania, 35

Urolepis, 310, 311

Utricularia, 10, 9

Vaccinium, 156, 161

Valeriana, 1)0

Valerianaceae, 11,0

Variegata, 55

Veraena, 212, 282, 35

Verbascum, 11,0

Verbena, 181-196, 199, 211-2h5,
2h7, 257-271, 273-296, 317,
318, 366-379, 381, 383, 385-
388, 413-15, 417-19, 421-23,
426-128, 431, L34-L37, 60,
461, 463-73

Verbenaca, 377, 378, 437, 472

Verbenaceae, 197, 199, 316, 318,

Taraxacum, 9, 121, 122, 126, 140 13, h2

Tecoma, )),3

Tectona, 17, 423, 426, 435
Tetradymia, 16)

Tetramerium, 17

Tetrandra, 377, 435
Tetraneuris, 79, 140
Tetraphila, 331

Teucridium, 26, )35
Teucrium, 10, 265, 302, 387

Verbenna, 20, 437

Verbenoxylun, 418, 428

Yerbina, 156, 378, 137

Vermicularia, 37

Vernonia, 7, 10, 12, 140, 350,
357, 350

Veronica, 134, 140, 265, 36,

437
Veronicastrum, 1,0, 26)

512 PHY T 0) 4, 0:05 Vol. 23, no. 5

Veronicena, 27

Vervena, 257, 37

Viter, 37

Vitex, 211, 315, 13-16, )18-
421, 423-25, 427, 430, 43h,

437, 438
Vittaria, 6

Volkameria, 38
Volkameriae, 38
Weilbachia, 331

Wigandia, 7

Xanthium, 5, 67, 68, 10
Xanthoparmelia, 36
Xanthoxalis, 10

Xyris, hh9

Yueca, 190, 235
Zaluzania, 238

Zapania, 38

Zea, 248, 2h9

Publication dates for Volume Twenty-three

No. 1 -—- March 2, 1972
No. 2 — April 7, 1972
No. 3 -—- May 20, 1972
No. -- June , 1972
No.5 -— July , 1972

Bree
Si