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PHYTOLOGIA

A cooperative nonprofit journal designed to expedite botanical publication
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Vol. 49 September 1981 No. 2

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BOTANICAL GARDEN
CONTENTS

LOWRY, P. P. II, A floristic survey of the Bear Trap Canyon,
Madison County, Montana, with a discussion of author

citations using the connecting words in or ex............ 81
KASAPLIGIL, B., Past and present oaks of Turkey. PartI .......... 95
WURDACK, J. J., Certamen Melastomataceis XXXIII .............. 147
MOLDENKE, H. N., Additional notes on the genus Priva. X ......... 159
MOLDENKE, H. N., Additional notes on the genus Vitex. XX........ 161
MOLDENKE, H. N., Notes on new and noteworthy plants. CXLIX..... 182

BAF ee hs) OK, FCPRCWISS Foose gn ak din dee RS wie aa bh Be 183

ba

Published by Harold N. Moldenke and Alma L. Moldenke

303 Parkside Road
Plainfield, New Jersey 07060
USA.

Price of this number $3.00; for this volume $12.00 in advance or $13.00 after
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after a volume is closed.

A FLORISTIC SURVEY OF THE BEAR TRAP CANYON,
MADISON COUNTY, MONTANA, WITH A DISCUSSION OF AUTHOR CITATIONS
USING THE CONNECTING WORDS IN OR EX

Porter P. Lowry II
Department of Botany
University of Illinois
Urbana, IL 61801

ABSTRACT: The results of a floristic survey of the Bear Trap
Canyon are presented in the form of an annotated checklist. Two
hundred and forty-two taxa are listed representing 162 genera and 50
families. Habitat and distributional data are provided for each
taxon. The checklist is prefaced by a discussion of the topography
and major plant community types of the area. Problems concerning
author citations using the connecting words in or ex, and the
consequences of their incorrect usage, are discussed.

A floristic survey of the Bear Trap Canyon, Madison County,
Montana, was conducted during the summer of 1979 while the author
was employed by the Butte District Office, Bureau of Land
Management, U.S.D.I. The study was made in order to determine the
total vascular plant flora of the area, as well as to assess the
occurrence, frequency, and distribution of actually or potentially
threatened or endangered plant taxa, in compliance with the
Endangered Species Act of 1973 (Public Law 93-205).

The Bear Trap Canyon, also known locally as the Madison
Canyon, is located on the Madison River 13 km by air (17 km by
road) NNE of Ennis, MT, in an area where the river cuts through the
Precambrian bedrock of the Norris Hills (de la Montagne, 1960).
Here the river flows rapidly to the north, dropping from an
elevation of 1470 m at Ennis Lake to approximately 1420 m at the
Missouri Flats located at the north end of the canyon. Throughout
much of its nearly 16 km length this sharply incised, V-shaped
canyon is in excess of 400 m deep. A small dam, associated with
the Madison Powerhouse, is situated about 3.5 km N of the entrance
to the canyon, and a primitive settlement has been established at
the confluence of Bear Trap Creek and the Madison River. An
undeveloped trail runs along the E bank of the river.

The vegetation differs drastically between the E and W sides
of Bear Trap Canyon. In general, the eastern slope (W-facing) has
large stands of rich, fairly mesic forest dominated by Pseudotsuga
menziesii var. glauca, which often extend down to the shores of
the Madison River. Individuals of Pinus contorta var. latifolia
occur in drier areas. A large number of herbaceous taxa is found
in these forests, including Osmorhiza chilensis, Arnica cordifolia
var. cordifolia, Mertensia oblongifolia var. oblongifolia,

81

82 renunt Olio 22 Vol. 49, No. 2

Smilacina racemosa, S. stellata, Dodecatheon pulchellum ssp.
pulchellum, Actaea rubra, and Clematis occidentalis var.
grosseserrata. Interspersed among these forested areas are fairly
moist to dry, open grasslands and rocky slopes. Four permanent
creeks occur on this side of the canyon, as well as a number of
intermittent drainages.

By contrast, the western slope of the canyon is quite xeric,
with dry, open grassland (often with scattered sagebrush), rocky
slopes, and very little forested area. No continuously flowing
creeks are found on this side of the canyon, and only a few draws
have water in the spring.

A borad diversity of habitat types occurs in the canyon bottom
itself. Many aquatic and semi-aquatic plants are found along the
banks of the Madison River, including large, somewhat scattered
populations of Carex nebraskensis, C. brevior, Scirpus microcarpus,
Juncus balticus var. vallicola, and | Ceratophyllum demersum. The
rocky | banks of the river provide mesic habitats for many species
that appear unable to tolerate long periods of submergence. Fairly
large stands of Prunus virginiana var. melanocarpa, Salix exigua,
and Sambucus racemosa ssp. pubens occur along the banks of the
river, particularly toward the N end of the canyon. Betula
occidentalis and Cornus sericea ssp. occidentalis form dense
thickets along the lower parts of the tributary creeks, and a
number of herbaceous taxa are restricted to the cool, moist shade
they provide. Some of these are: Heracleum lanatum, Mertensia
ciliata var. ciliata, Carex athrostachya, C. Cc. sprengelii, Strept Streptopus
amplexifolius var. chalazatus, Circaea alpina, Ribes hudsonianum
var. petiolare, Viola praemorsa, and Platanthera a dilatata var.
dilatata.

Fairly large, open, seasonally moist meadows occur on the
alluvial fans of the larger tributary creeks where they flow into
the Madison River. These meadows exhibit more floristic diversity
than any other single habitat type in the canyon. Conspicuous
members of these communities include Senecio serra var. serra,
Symphoricarpos occidentalis, Carex spp., Geranium richardsonii,
Agastache urticifolia var. urticifolia, Monarda fistulosa var.
menthifolia, Elymus cinereus var. cinereus, Phleum pratense,
Delphiniun « occidentale, and Physocarpus malvaceus.

Much drier meadows and grasslands, often with Artemisia
tridentata, occupy other level areas and open slopes along the
river. The vegetation here resembles very much that found on the
open areas higher up the canyon walls. Commonly encountered
taxa of these habitats include several species of Lomatium,
Balsamorhiza sagittata, Gaillardia aristata, Lupinus arbustus ssp.
calcaratus, L. burkei ssp. burkei, Oxytropis lagopus var. lagopus,
Agropyron trachycaulum var. unilaterale, Bromus inermis var.
purpurascens, Dodecatheon conjugens var. conjugens, and Delphinium

bicolor.

1981 Lowry , Bear Trap Canyon 83

Dry rock outcrops are fairly common in the canyon bottom, as
well as on its western slopes. These sites support a sparse
vegetation which includes taxa such as Woodsia oregana, Erigeron
caespitosus, Stephanomeria tenuifolia var. tenuifolia, Arenaria
capillaris ssp. americana, Sedum lanceolatum ssp. lanceolatun,
Bouteloua gracilis, Bromus tectorum, Festuca idahoensis, and
Collomia linearis.

A number of weedy and introduced plants are found around the
Madison Powerhouse, the settlement at Bear Trap Creek, and along
the road and parking area at the N end of the canyon. These
include some species which have escaped from cultivation and have
become established in more or less disturbed areas, primarily
along the trail.

Only one plant taxon observed within the Bear Trap Canyon is
of possible significance as a threatened species, Phlox
albomarginata. This plant is known only from the mountains of
western Montana and eastern Idaho (Hitchcock and Cronquist, 1973).
Although P. albomarginata may be fairly common in certain areas, it
was included on a list of plants with restricted distributions in
Montana prepared by R. D. Dorn (pers. comm.). This taxon is in
need of further detailed study to determine whether it should be
considered as a threatened species.

Collections were identified using Hitchcock and Cronquist
(1973), and nomenclature follows Kartesz and Kartesz (1980), with
the following exceptions; Carex (Hermann, 1970), Juncus (Hermann,
1975), Sisyrinchium (Henderson, 1976), Aster (identified by A. G.
Jones) and Lomatium (identified by M. A. Schlessman). Three
additional references were consulted to verify certain
identifications (Booth, 1972; Booth and Wright, 1966; Hitchcock et
al., 1955-1969). Voucher specimens are deposited in MONT, with
many duplicates in MONTU and ILL.

All author citations in which the connecting words in or ex
are used have been verified and agree with Recommendations 46C and
46D, respectively, of the International Code of Botanical
Nomenclature (Stafleu et al., Editors, 1978). Verification of
these author citations was necessary as many of them are
incorrectly given in Kartesz and Kartesz (1980), as well as

elsewhere in the literature.

The distinctions between citations using in and ex are
extremely important, and often not fully appreciated. Abbreviation
of citations in which ex is used incorrectly in the place of in
results in authorship being attributed to the person publishing the
name, rather than to the person actually responsible for both
naming and describing the new taxon. Conversely, abbreviation in
situations where in has been used incorrectly attributes authorship
to the person who supplies only the name, excluding citation of the
authority who actually described it.

84 P BY TO: 0 Git aA Vol. 49, No. 2

For example, there is considerable confusion concerning the
author citation for names supplied by Nuttall and published by
Torrey and Gray in their Flora of North America (1838-1843).
According to Ewan, in the introduction to the facsimile version of
Torrey and Gray's Flora (1969:iii): ''Thomas Nuttall had agreed in
1837 to furnish Torrey and Gray with descriptions of hundreds of
new species which he had discovered in his western travels. These
would carry his name as author."' Clearly the descriptions for
these taxa were supplied by Nuttall and merely published by Torrey
and Gray. The correct author citation for these names is "Nuttall
in Torrey and Gray," in accordance with Rec. 46D of the Code.
Should it become necessary or desirable, for editorial reasons, to
abbreviate this citation, the Code states (p. 40): "The name of
the author who supplied the description or diagnosis is the most
important and should be retained..." This citation is simply
abbreviated as "Nuttall."

However, many authors of floristic and monographic works have
incorrectly cited the authority of these names as "Nuttall ex
Torrey and Gray," with potentially undesirable consequences (e.g.,
Hitchcock et al., 1955-1969; Kartesz and Kartesz, 1980).
Recommendation 46C of the Code states (p. 40): "When an author who
first validly publishes a name ascribes it to another person, the
correct author citation of the name is the actual publishing
author, but the name of the other person, followed by the
connecting word ex, may be inserted before the name of the
publishing author, if desired."" Abbreviation of the author
citation "Nuttall ex Torrey and Gray" would result in the citation
of "Torrey and Gray" as authors of all the names furnished them by
Nuttall. Clearly this is incorrect, as well as being contrary to
the intentions of Torrey and Gray.

Author citations using the connecting word ex are reserved for
situations where only the name is supplied by one person, the
description or diagnosis being prepared by the publishing author.
This is not the case with Nuttall's names, as he supplied carefully
prepared descriptions with his specimens, albeit sometimes modified
by Torrey and Gray. Unfortunately, many botanists have been
careless in their use of the word ex in author citations,
frequently applying it in situations where the word in should be
used. This error has resulted in incorrect abbreviation of author
citations such that only the publishing author is cited, at the
exclusion of the name of the person who named and described the new
plant. Errors of this sort are by no means restricted to Nuttall's
names published in Torrey and Gray's Flora. The contrary
situation, in which in is used improperly, is less common.

In the following checklist the taxa have been arranged
alphabetically to family, genus, and species under the following
categories; Sphenophytina (horsetails), Filicophytina (ferns),
Coniferophytina (conifers), Magnoliopsida (dicotyledons), and
Liliopsida (monocotyledons). The following abbreviations are

1981 Lowry, Bear Trap Canyon 85

used to designate the habitat(s), frequency, and area(s) in which
each taxon occurs:

aq aquatic pf Pseudotsuga forest

cb canyon bottom ra rare

co common rb river banks

da disturbed areas ro rock outcrops

dg dry grasslands sb stream banks

he higher elevations sg sagebrush grasslands

lo local sm seasonally moist meadows
me middle elevations sp sparse

mm moist meadows we weedy

of open forests ws widespread

os open slopes

SPHENOPHYTINA (Horsetails)

EQUISETACEAE
Equisetum fluviatile L. das ens °sp 3° eb
Equisetum pratense Ehrh. nm, ED 5 - Sps “eb
FILICOPHYTINA (Ferns)
ASPLENIACEAE
Athyrium filix-femina (L.) Roth prs ‘sp; eb
POLYPODIACEAE
Woodsia oregana D.C. Eat. ro; sp; cb, me
CONIFEROPHYTINA (Conifers)
CUP RESSACEAE
Juniperus scopulorum Sarg. of arb; ‘Sps "eb
PINACEAE
Pinus contorta Dougl.
var. latifolia Engelm. of; pi; sp 3 me, he
Pinus flexilis James Gf, (pis spss. he
Pseudotsuga menziesii (Mirbel) Franco
var. glauca (Beissn.) Franco pf; co; ws
MAGNOLIOPSIDA (Dicotyledons)
ACERACEAE
Acer glabrum Torr.
var. douglasii (Hook.) Dippell pi, 08; (cb, te
ANACARDIACEAE
Rhus trilobata Nutt. es, ie3 4607 ‘Lo; ‘eb
Toxicodendron radicans (L.) Kuntze os. stbs 103 eb
AP TACEAE
Cicuta douglasii (DC.) Coult. & Rose tbh; sp? cb
Cymopterus bipinnatus S. Wats. os, Sg; sp; me, he
Heracleum lanatum Michx. shee, Lo; cb

Lomatium ambiguum (Nutt.) Coult. & Rose To,V0s; spi eb

86

Par eT OL oe ts

Vol. 49, No
Lomatium cous (S. Wats.) Coult. & Rose 06; Bg; cb,
Lomatium dissectum (Nutt. in Torr. & Gray)

Math. & Const. var. eatonii (Coult. &

Rose) Cronq. TO, O68; sp;
Lomatium foeniculaceum (Nutt.) Coult. & Rose

var. macdougalii (Coult. & Rose) Cronq. SZ; sp;
Lomatium triternatum (Pursh) Coult. & Rose

ssp. platycarpum (Torr.) Cronq. YO: co; cb,
Musineon divaricatum (Pursh) Nutt. in Torr.

& Gray G6, 883 Ta;
Osmorhiza chilensis Hook. & Arn. pL, 08, 805.2).
Osmorhiza depauperata Phil. Sb, pr:..sp3
Perideridia gairdneri (Hook. & Arn.) Math.

ssp. borealis Chuang & Const. sm; sp;
Sium suave Walt. tb5. s05

APOCYNACEAE
Apocynum androsaemifolium L.
ssp. pumilum (Gray) Boivin var. pumilum os, rb; sp;
ASTERACEAE
Achillea millefolium L.

var. lanulosa (Nutt.) Piper Sm, (OS; co,
Antennaria microphylla Rydb. OS). £6: cas
Arctium minus (Hill) Bernh. da, pip sp, “to:
Arnica cordifolia Hook. var. cordifolia G5 PES e035
Artemisia ludoviciana Nutt. Os ,, Sm; Sp;
Artemisia tridentana Nutt. SZ3 co;
Aster conspicuus Lindl. in Hook. sm; sp;
Aster hesperius Gray fb spe
Aster occidentalis (Nutt.) Torr. & Gray sb; sp;
Balsamorhiza sagittata (Pursh) Nutt. ses: Spo 1e-
Brickellia grandiflora (Hook.) Nutt. pi, reg sp
Carduus nutans L. os? lo, we;
Centaurea maculosa Lam. os>10, wes
Cirsium arvense (L.) Scop.

var. horridum Wimm. & Grab. OS; SP, we;
Cirsium undulatum (Nutt.) Spreng. dg: sp;
Cirsium vulgare L. OS, SM; Sp, we;
Crepis acuminata Nutt. ssp. acuminata OS) 825° SP
Erigeron caespitosus Nutt. ra, ‘os > lo;
Erigeron formosissimus Greene $0, OS; sp;
Erigeron speciosus (Lind1l.) DC.

var. macranthus (Nutt.) Crongq. iim, em, pfs cos
Erigeron strigosus Muhl. var. strigosus SZ; Sp, We;
Erigeron subtrinervis Rydb.

var. conspicuus (Rydb.) Cronq. sm; Sp;
Euthamia graminifolia (L.) Cass.

var. major (Michx.) Moldenke rb; sp;
Gaillardia aristata Pursh oS, SM; CO;
Grindelia squarrosa (Pursh) Dunal

var. quasiperennis Lunell dg, sg; co;

me

cb

Lowry, Bear Trap Canyon

Helenium autumnale L.
var. montanum (Nutt.) Fern.
Heterotheca villosa (Pursh) Shinners
var. hispida (Hook.) Harms
Hieracium cynoglossoides Arv.-Touv.
Lactuca serriola L.
Lactueéa-tatarica (L.) C.A. Mey-
ssp. pulchella (Pursh) Stebbins
Leucanthemum vulgare Lam.
Liatris punctata Hook.
Rudbeckia laciniata L.
var. ampla (A. Nels.) Cronq.
Rudbeckia occidentalis Nutt.
var. occidentalis
Senecio canus Hook.
Senecio hydrophilus Nutt.
Senecio integerrimus Nutt.
var. exaltatus (Nutt.) Cronq.
Senecio serra Hook. var. serra
Solidago missouriensis Nutt.
wat. Lascientata Holz.
Solidago multiradiata Ait.
var. scopulorum Gray
Sonchus arvensis L.
ssp. uliginosus (Bieb.) Nyman
Sonchus oleraceus L.
Stephanomeria tenuifolia (Torr.) Hall
var. tenuifolia
Tanacetum vulgare L.

Tragopogon dubius Scop.

BETULACEAE

Betula occidentalis Hook.

BORAGINACEAE

Cynoglossum officinale L.
Hackelia deflexa (Wahlenb.) Opiz var.
americana (Gray) Fern. & I.M. Johnst.
Hackelia micrantha (Eastw.) J.L. Gentry
Hackelia patens (Nutt.) I.M. Johnst.
var. patens
Lithospermum incisum Lehm.
Lithospermum ruderale Dougl. in Lehn.
Mertensia ciliata (Torr.) G. Don
War, €iliatsa
Mertensia oblongifolia (Nutt.) G. Don

var. oblongifolia

BRASSICACEAE

Arabis sparsiflora Nutt, in Torr. & Gray
var. subvillosa (S. Wats.) Rollins
Berteroa incana (L.) DC.

os,
sm,

sm,

sm,

Os,
os,
rb;

pf,

os,
Sg,

Os,

sm,
sm,

sb,

pf;

rb;
os,

ro}
S85

sb;

os;

pf,
OSs;

OS;

sb;

pf;

dg,

SP;

CO}

SP;

ro;
co,

SP>
sm3
SS;

co3

Pe pe

of;
Sg;

SP;
SP;
we;

CO;
SP;
SP;

SP;
SP;
Ea;

SP;

SP;
co:

SP;
cb,

Sp;
lo;

co;

we;
eb,

cb,

we;

SP;
lo;

lo:
SP;
SP;
cn.

cb,

Fras
SP;

me

cb

me

me

88 Pw iT OL OG ir a

Camelina microcarpa Andrz. ex DC.
Cardamine breweri S. Wats. var. breweri
Descurainia richardsonii (Sweet) O.E.

Schulz ssp. viscosa (Rydb.) Detling
Draba nemorosa L.
Erucastrum gallicum (Willd.) 0O.E. Schulz
Erysimum inconspicuum (S. Wats.) MacM.
Lepidium densiflorum Schrad.

var. macrocarpum Mulligan
Lepidium virginicum L.

var. pubescens (Greene) C.L. Hitchc.
Rorippa palustris (L.) Bess.

ssp. hispida (Desv.) Jonsell

var. hispida (Desv.) Rydb.

Sisymbrium altissimum L.

CACTACEAE
Opuntia polyacantha Haw.

CAMP ANULAC EAE
Campanula rotundifolia L.

CAPRIFOLIACEAE
Sambucus racemosa L.
ssp. pubens (Michx,) House

sm,

sm,

SB»

S83

dg

Symphoricarpos occidentalis Hook. mm,

CARYOPHYLLACEAE
Arenaria capillaris Poir,
ssp. americana Maguire
Cerastium arvense L.
Cerastium fontanum Baumg.
ssp. triviale (Link) Jalas
Silene alba (P. Mill.) Krause

CERATOPHYLLACEAE
Ceratophyllum demersum L.

CORNACEAE
Cornus sericea L.
ssp. occidentalis (Torr. & Gray) Fosberg

CRASSULACEAE
Sedum lanceolatum Torr. ssp. lanceolatum

FABACEAE
Astragalus canadensis L,
var. brevidens (Gandog.) Barneby
Astragalus crassicarpus Nutt.
var. paysonii (E.H. Kelso) Barneby
Astragalus lentiginosus Dougl. in Hook.
var. platyphyllidius (Rydb.) M.E.Peck

aq,

os,

Vol.

dg;

C05

SB>

ro,

os,

S8>

ro;

sm,
S83

os,

49,

SB;
$a,

oc*
mm $
SP>
SB;
SP;
Le,

rb;
Os;

S8;

dg;

OS}
pt;

dg;
ro;

OS;

dg;

co,

eo.

Os;

os;

cO3

ro}

SP;
16%

co;
SP;
we;
SP;
cb,
we;

SP;
SP;

SP;

co:

co;
eo;

ed:
SP;

SP>
SP;

ie:

los

co}

SP;

fol oy

SP;

cb

cb

cb
ws

cb

cb

ws

cb

me

cb

1981 Lowry, Bear Trap Canyon

Glycyrrhiza lepidota Pursh var. glutinosa

(Nutt. in Torr. & Gray) S. Wats.
Lupinus arbustus Dougl. ex Lindl.

ssp. calcaratus (Kellogg) D. Dunn
Lupinus burkei S. Wats. ssp. burkei
Lupinus sericeus Pursh var. sericeus

Medicago lupulina L.

Medicago sativa L.
Melilotus alba Medic.

Melilotus officinalis (L.) Pallas
Oxytropis lagopus Nutt. var. lagopus
Oxytropis sericea Nutt. in Torr. & Gray
var. sericea
Thermopsis montana Nutt. in Torr. & Gray
var. montana
Trifolium longipes Nutt. in Torr. & Gray
ssp. reflexum (A. Nels.) Gillett
Vicia americana Muhl. in Willd.
ssp. americana

GERANTACEAE
Geranium richardsonii Fisch. & Trautv.
Geranium viscosissimum Fisch. & Mey.
var. viscosissimum

GROSSULARTACEAE
Ribes aureum Pursh var. aureum
Ribes inebrians Lindl.
Ribes hudsonianum Richards.

var. petiolare (Dougl.) Jancz.
Ribes setosum Lindl.

HYDRANGEACEAE
Philadelphus lewisii Pursh

HYDROPHYLLACEAE

Hydrophyllum capitatum Dougl. in Benth.
var. capitatum

S8>

sm,

sm,
sm,

sb3

sb;

Phacelia hastata Dougl. in Lehm. ssp. hastata

Phacelia linearis (Pursh) Holz.

LAMIACEAE

Agastache urticifolia_ (Benth.) Kuntze
War. urticifolia

Lycopus asper Greene

Mentha arvensis L. ssp. haplocalyx Briq.

Monadra fistulosa L.
var. menthifolia (Graham) Fern.

Nepeta cataria L.

Prunella vulgaris L.

Scutellaria galericulata L.

sm,

S85

SP;

sm,

Os,

sm,
rb;

rhs

cO3

SP;

SP;
SP;

ros

lok

SP;

sm;
sm;
Los

sm;

OSs;

SP;

me,
SP;
Eby
SP;
SP;
we;
we;
Los

SP;
SP;
cb,

SP;

eb,

eb,

cos
rete Fs

cb:,
SP;

ero

SP;
co;

SP;

cos

SP;
lo;

Le;
SP3
SP;

ae are

me

me

ws

90 POY TOD Oo ois

MALVACEAE
Alcea rosea L.
Iliamna rivularis (Dougl. in Hook.) Greene
var. rivularis

ONAGRACEAE
Circaea alpina L.

Epilobium angustifolium L.
Epilobium minutum Lindl. ex Hook.

Gaura coccinea Pursh

PLANTAGINACEAE
Plantago major L. var. major

POLEMONIACEAE
Collomia linearis Nutt.
Gilia tenerrima Gray
Phlox albomarginata M.E. Jones

POLYGONACEAE

Eriogonum umbellatum Torr. var. majus Hook,

Polygonum lapathifolium L.

sm;

sb;

sm,

os,

Rumex crispus L. sb, mm,

Rumex triangulivalvis (Danser) Rech. f.
var, triangulivalyvis

PORTULACACEAE

Claytonia perfoliata Donn
Lewisia rediviva Pursh

PRIMULACEAE
Dodecatheon conjugens Greene

SE,

var. conjugens pr... aly

Dodecatheon pulchellum (Raf.) Merr.

ssp. pulchellum
Lysimachia ciliata L.

Lysimachia thyrsiflora L.

RANUNCULACEAE
Actaea rubra (Ait.) Willd.
Aquilegia flavescens S. Wats.
Clematis occidentalis (Hornem.) DC.
var. grosseserrata (Rydb.) J. Pringle
Clematis lingusticifolia Nutt.
in Torr. & Gray
Delphinium bicolor Nutt. & Wyeth

Delphinium occidentale (S. Wats.) S. Wats,

ssp. occidentale
Ranunculus abortivus L.
Ranunculus acriformis Gray

var. montanensis (Rydb.) L. Benson
Ranunculus macounii Britt.

pi,
pf,

sn,

Vol.

co,

co,

OSs;

rez
Os,

sm,

sm;

pi.
dg;

os,

sm,
rhs

pf,
sm,

of,

OS ;
os,

sm};

rb3

49,

LO,
sm;
lo;

S8;
S83

SP;

SE;

S83
OSs;

CO,
rb;

of;
Os;
Os;

co,
ro;

co,
rb3

sb;
co,

cb,
lo;
co;
SP;

we;

lo;
SP;
ra;

co;
SP 5
we;

SP;

co:

; me,

cb

cb

cb

ws
he

* cb

Oo; ¢b

Lowry, Bear Trap Canyon

Ranunculus uncinatus D. Don in G. Don
var. uncinatus

Thalictrum dasycarpum Fisch. & Lall.

Thalictrum venulosum Trel.

ROSACEAE

Amelanchier alnifolia (Nutt.) Nutt.
var. alnifolia
Fragaria vesca L.
ssp. bracteata (Heller) Staudt
Geum triflorum Pursh var. triflorum
Physocarpus malvaceus (Greene) A. Nels.
Potentilla anserina L.
Potentilla arguta Pursh
Potentilla biennis Greene
Potentilla gracilis Dougl. ex Hook.
var. flabelliformis (Lehn.)
Nutt. in Torr. & Gray

Potentilla pensylvanica L.
Prunus virginiana L.

var. melanocarpa (A. Nels.) Sarg.
Rosa woodsii Lindl.

var. ultramontana (S. Wats.) Jepson
Rubus idaeus L.

ssp. sachalinensis (Levl.) Focke
Rubus parviflorus Nutt.
Spiraea betulifolia Pallas

ssp. lucida (Dougl. ex Greene)

Taylor & MacBryde

RUBIACEAE

Galium aparine L.
Galium boreale L.

Galium tricornutum Dandy

SALICACEAE

Salix exigua Nutt.

SANTALACEAE

Comandra umbellata CL.) Nabe.
ssp. pallida (A. DC.) Piehl

SAXIFRAGACEAE

Conimitella williamsii (D.C. Eaton) Rydb.
Heuchera cylindrica Dougl. in Hook.

var. cylindrica
Heuchera flabellifolia Rydb.

var. flabellifolia

Lithophragma parviflora (Hook.)
Nutt. in Torr. & Gray

sm,

rb,

pf,

sm,
sm,

sm,

sb;

sm,

sb,
pf;
os,

sm,

os,
Sb?

sm,
sm:
sm,
sm,
sm,

sm,

TB;
of,
os,

rb;

S83

sm }

ra3
rb;
OSs}

sm;
Sp;
S85
OSs;
rb3
EOE
SP;

sm;
OSs;
Co;
Os;
OSs
sb3

on:

sb;
Os;
sb;

co,

co,

SP;
SP,
So,

sb;

co;
SP;

SP;

SP 5
SP;
SP;

lo;

Los

cb,
lo;
lo;

SP;

ws

ws
ws
ws

cb

ws

me

cb

cb

ws

92 P EY DT 0A 6. Tea

SCROPHULARIACEAE
Castilleja hispida Benth. in Hook.

ssp. acuta Pennell

Castilleja miniata Dougl. ex Benth. in Hook.

Var. Miniata

Castilleja pallescens (Nutt. ex Gray) Greenm.

Linaria genistifolia (lise 3.

ssp. dalmatica (L.) Marie & Petitmengin

Penstemon attenuatus Dougl. ex Lindl.
var. pseudoprocerus (Rydb.) Cronq.

Penstemon nitidus Dougl. ex Benth.
var. nitidus

Verbascum thapsus L.

Veronica serpyllifolia L.
ssp. humifusa (Dickson) Syme

SOLANACEAE
Solanum dulcamara L.

URTICACEAE

Urtica dioica L. ssp. gracilis (Ait.) Seland.

var. lyallii (S, Wats.) €.L, Hitche-

VERBENACEAE
Verbena bracteata Lag. & Rodr.

VIOLACEAE
Viola canadensis L.

var. corymbosa Nutt. in Torr, & Gray
Viola praemorsa Doug]. ex Lindl,

LILIOPSIDA (Monocotyledons)

CYPERACEAE
Carex athrostachya Olney
Carex brevior (Dewey) Mack.
Carex douglasii Boott
Carex foenea Willd.
Carex lanuginosa Michx.
Carex nebraskensis Dewey

Carex petesata Dewey
Carex praticola Rydb.

Carex sprengelii Dewey
Carex vernaculata Bailey

Eleocharis palustris (L.) Roemer & Schultes

Scirpus microcarpus Presl

IRIDACEAE
Iris missouriensis Nutt.
Sisyrinchium idahoense Bickn.
var. occidentale (Bickn.) Henderson

Vol.

sm,

sm,
SB»

os,
sm,

os,
os;

1g ie

sb;

os,

pi.

sb;
rb;
sm,
sm,
rb;
EDA
os;

sb;
rb:
5 99
sm;

sm,

49,

OSs;

pf;
dg;

da;
OS;

dg;
eG.

Th

SP,

SP;

aa.

sb;

Sp,

Os;

SP;
SP;

SP;
SP;

SP;
we;

fra;

we;

rel A

SP;

ce:
ra:

Los
Los
SP;

> SP;

le;
as te
los

5 als

Tes
SP;

ios
lo;

ip:

ra:

cb

cb

me

cb

ws

cb

me

1981 Lowry, Bear Trap Canyon 93

JUNCACEAE

Juncus balticus Willd. var. vallicola Rydb. Eb; "ee, Lor eb

guncus filirtormis i

mm, OS3 Sp 3 cb

LILIACEAE

Allium cernuum Roth sii, To; co, lo; eb
Allium textile A. Nels. & J. F. Macbr. TO, OS; sp; cb
Fritillaria atropurpurea Nutt. Dm: COs LOS ‘eb
Smilacina racemosa (L.) Desf. pL. - Of s 2sp* ws
Sui Vacina stellata (U.)- Desf: sb, pes ‘co? ws
Streptopus amplexifolius (L.) DC.

var. chalazatus Fassett sb; ‘co, "fos; cb, me
Zigadenus venenosus §S. Wats.

var. gramineus amineus (Rydb. ) Walsh sm, OS; Sp; ws

ORCHIDACEAE

Platanthera dilatata (Pursh) Lindl.

Var. Gibataeta

POACEAE

sb; sp; cb, me

Agropyron trachycaulum (Link) Malte

var. trachycaulum

sp, “des "co, les; ws

Agrostis stolonifera L. var. stolonifera Lbs: Sis <b
Agrostis thurberiana A.S. Hitchc. pf; sp; cb, me
Bouteloua gracilis / gracilis (H. B.K.) ‘hae; se, de, O83. ca; ws

Bromus anomalus Rupr.
Bromus inermis Leyss.

Di; OLS cos. ws

ssp. " pumpellianus (Scribn.) Wagnon
var. pumpellianus (Scribn.) Wagnon os; sp, lo; ws

Bromus tectorum L.

ass sp, to: ws

Calamagrostis canadensis (Michx.) Beauv.

var. canadensis

Dactylis glomerata L.

mm, sm; sp; cb
tb, sb, sii; sp 7 och

Elymus cinereus Scribn. & Merr.
var. cinereus sm: ‘co; lo; ws
Festuca idahoensis Elmer se, deseo; lo: ws

Phleum pratense L.

sm, OS$ CO; ws

Poa nervosa (Hook.) Vasey
var. wheeleri (Vasey) C.L. Hitchc. sm, OS; SP; ws

Poa palustris L. Le
Poa pratensis L.

rb; sp, we; cb
sm; sp; ws

LITERATURE CITED

Booth, W. E. 1972. Grasses of Montana. Dept. of Botany and
Microbiol., Montana State Univ., Bozeman. 64p.

and J. C. Wright.

Dicotyledons. Ed. 3,
State Univ., Bozeman.

1966. Flora of Montana. Part II.
Dept. of Botany and Microbiol., Montana
305p.

94 PRY 7 0.4.0 6.3% Vol. 49, No. 2

de la Montagne, J. 1960. Geomorphic problems in the Madison
Valley, Madison County, Montana: an introduction and
synthesis. Eleventh Ann. Field Conf,, Billings Geol. Soc.,
pp. 165-169.

Ewan, J. 1969. Editor's Introduction. Torrey, J. and A. Gray, A

Flora of North America. Facsimile Version. Hafner Publ. Co.,
New York and London.

Henderson, D. M. 1976. A biosystematic study of Pacific Northwest
blue-eyed grasses (Sisyrinchium - Iridaceae). Brittonia 28:
149-176.

Hermann, F. J. 1970. Manual of the carices of the Rocky Mountains
and Colorado Basin. U.S.D.A., For. Serv, Agric. Handb. 374.
397p.

1975. Manual of the rushes (Juncus spp.) of the Rocky
Mountains and Colorado Basin. U.S.D.A., For. Serv, Gen. Tech.
Rep. RM-18. 107p.

Hitchcock, .C. L. and A. Cronguist. 1973... Flora af the Pacific
Northwest. Univ. Wash. Press, Seattle and London. 730p.

» M. Ownbey, and J. W. Thompson. 1955-1969.
ees plants of the Pacific Northwest. Parts 1-5. Univ.
Wash. Press, Seattle.

Kartesz, J. T. and R. Kartesz. 1980. A synonymized checklist of
the vascular flora of the United States, Canada, and Greenland.
Univ. North Carolina Press, Chapel Hill. 498p.

Stafleu, F., V. Demoulin, W. Greuter, P. Hiepko, I. A. Linczevski,
KR. MeVaugh, R. D.. Meikle, BR. G. Rollins, BR. Ross, J.. M.
Schopf, and E. G. Voss [eds.]. 1978. International Code of
Botanical Nomenclature. Regnum Vegetabile, Vol. 9/7. Bohn,
Scheltema and Holkema, Utrecht, The Netherlands. 457p.

Torrey, J. and A. Gray. 1838-1843. A flora of North America.
2 Vols. Wiley and Putnam, New York.

ACKNOWLEDGMENTS

I am indebted to J. H. Rumely, Curator of the Herbarium at
MONT, for his assistance in identifying a number of difficult
collections, and for his encouragement. Special thanks go to A. G.
Jones for valuable comments on the manuscript, and for identifying
specimens of Aster. I also thank M, A. Schlessman for determining
material of Lomatium. I am grateful to H. Le Dantec and S. C.
Lowry for their able assistance in the field. Acknowledgment is
due to the Bureau of Land Management, U.S.D.I., for permitting
publication of this work.

*

This research was partly supported by the grant No.

PAST AND PRESENT OAKS OF TURKEY. PART I*

Baki Kasapligil, Biology Department,
Mi lis -Gotltlege, Caktand, Gali£s 94613

Dedicated Eo “the toresiters
Turkey.

AB Ss) “he R 2A eve

The 450 "spp: Gf Quercus are restricted
to diverse environments of the north-
ern hemisphere. One of the ancient
angiosperm genera, it was abundant
during the Cretaceous associated with
spp. of Magnoliaceae and Lauraceae.
Tentatively, eighteen fossil taxa are
recognized, mostly from the upper Mio-
cene diatomite sediments along the
Gurcti Valley some 90 miles north of

Ankara. From western Anatolia, addi-
tional leaf remains were collected from
open strip mines of lignite. Most cor-

iaceous leaf impressions are of ever-
Green faxa »wWHth "at Fant tres “be texbane
Caucasian, Himalayan and east asiatic
spp. Presently Quercus is represented

by -20\ spp. in Turkey, most “oF which are
deciduous trees of economic importance.
Distributional maps indicate the geo-
graphical extent of these spp. Q.troyana
and Q.pontica are very rare Tertiary
relic spp. which merit protection from

danger of extinction. Interspecific
hybridization is common; 31 hybrids
have been recognized. No attempt has

been made to describe the hybrids of
the fossil spp. although the magnitude
of foliar and cupular-variation sug-
gests a high degree of hybridization.

My interest in the genus Quercus started in 1937
when I was a student of Kurt Krause,

DEB

of

one of the collabor-

76-06883 from the National Science Foundation and partly

by a Faculty Research Grant from Mills College,

California 94613.

55

Oakland,

96 Pat Y TOL 0:65. A Vol. 49, No. 2

ators of Adolf Engler. Amentiferae, so-called primitive
angiosperms, were exemplified by the oaks and oak allies
under the order Fagales. Since then, I have been collect-
ing oaks during my botanical excursions as well as during
my leisure trips in Turkey and elsewhere. My collections
grew bigger and bigger and tempted me to study them for
my doctoral dissertation at the University of California,
Berkeley. Unfortunately, my specimens were shipped to a
wrong address and it took me many years to locate them.
Meanwhile as students of Herbert Mason at U.C. Berkeley,
we questioned the primitive status of Amentiferae with
simple flowers versus the complex flowers of the Ranalian
groups. Most of us inclined to believe that the simpli-
city of the Amentiferae resulted from a series of abor-
tions of the floral appendages and therefore, Amentiferae
could not be primitive. In fact, we were pleased to note
that the heterogenous group of Amentiferae occupied ad-
vanced levels in the monophyletic system of Charles Bessey.
I postponed my studies of oaks and studied some of the
members of Lauraceae instead.

Many years later, I discovered quite incidentally
the Tertiary fossil beds of Gircti Valley near Gluven Vill-
age some 90 kilometers north of Ankara during a picnic
outing. During my first excavations in 1968, I collected
125 specimens of impressions, mostly belonging to the oaks,
maples, chestnuts, elms and Zelkova, Magnolias and Mahonias.
Among these broad-leaved trees, Quercus was the most
common element, represented by leaves, catkins, and cupule
impressions. Glyptostrobus, Sequoia, Taxodium and five
species of pines were the most common species of gymno-
sperms. When I showed the collections to the late Profes-
sor Ralph Chaney, he was most impressed with the redwoods,
of couse, since they had never been reported from Asia
Minor. Dr. Chaney identified the age of the Giiven deposits
as Upper Miocene while several other colleagues in Paleo-
botany and Geology suggested the Pliocene epoch. With the
support of the National Science Foundation, I made three
additional visits to the Giirctii (Georgian) Valley and with
the collaboration of Daniel Axelrod and the paleontologists
of the Mineral Research and Exploration Institute of Tur-
key (M.T.A.), we collected more than a thousand specimens
tay L375) 1916 .anad Lasse. The duplicate or counterpart spe-
cimens were deposited at the Turkish Museum of Natural
History M.T.A. in Ankara, but all the plant and animal
impressions were loaned to me for identification. With
the assistance of Professor Axelrod, we also selected rock
samples of andesite, rhyolite and welded tuffs for absol-
ute age determination through K/Ar isotopes in biotites
contained in these rocks. The age of the fossil deposits
in Gurctti Valley was 14.1 + 1 million years, corresponding

198i Kasapligil, Oaks of Turkey 97

to the Upper Miocene. This ancient coniferous forest,
rich in Arcto-Tertiary elements, occupied a mesophytic
environment around a large fresh-water lake as evidenced
by the abundance of frustules of pennate fresh-water dia-
toms, Cyprinid fish fossils, frogs, salamanders, mosquito
larvae, Nematoceran flies, dragon flies, bees and beetles
as well as by aquatic plants such as Salvinia, Egregia,
cattails,etc.. Apparently, repeated volcanic activities
took place around this fresh-water lake pouring lava and
hot volcanic ash into the lake which I believe was a rem-
nant of the Sea of Paratethys which consisted of brackish
water. The rapid sedimentation of diatomaceous frustules
associated with volcanic ash and the accumulation of fine
silt through the streamsflowing into the lake formed the
laminated diatomites and paper shales which contain the
fossil impressions. Unfortunately, most of the organic
matter was decomposed and I had no compressions to study
the vfotrar cuticies. From the sediments, we extracted a
great variety of pollen grains, most of which belong to
Quercus spp.. However the morphology of the pollen grains
under the resolving power of the light microscope was not
a useful tool for the species distinction. On the other
hand, the foliar venations of the oaks were well preserved
including the minor venations in many specimens. The ex-
tant species of the oaks of Turkey were the main tools of
comparison with those of the Miocene oaks. With the gen-
erous help of the Turkish Forestry Service, I built up a
sizable collection of the living oaks. Serveral leaves
from each of the twenty living species were cleared in
sodium hydroxide and Chloral hydrate solutions. After
having been stained with safranin, these cleared leaves
were mounted permanently in large-size glass plates. For
the description of the foliar vasculations, I followed the
standardized terminology outlined by Hickey and Dilcher.
Some Miocene oaks of Turkey exhibit strong affinities to
North American and Far Eastern taxa as I will point out

in this paper. The cleared leaves of the American and
Asian oaks were borrowed from the U.S. Geological Survey
in Menlo Park through the courtesy of Jack Wolfe and from
the slide collections of the U.C. Paleontology Department
in Berkeley through the courtesies of Wayne Fry and Howard
Schorn. The Miocene oaks of Turkey display an enormous
variation with respect to the foliar, floral and cupular
structures. Obviously, the Tertiary oaks exhibited an
enormous amount of interspecific hybridization and intro-
gression as is characteristic of the extant species of
oaks. Although I could recognize the possible hybrid oaks
among the extant species, it was impossible to make such
an attempt for the Miocene oaks since not a single speci-
men was collected with leaves, buds, catkins, or fruits

98 PHYTOLOGTIA Vol. 49, No. 2

attached to a branch.

As we know now, the oaks are not as recent as we
thought. Quercus, Quercophyllum and Dryophyllum were re-
ported from the Early Cretaceous of the northern hemi-
sphere. Approximately 135 million years ago, the oaks
were widely distributed together with the several members
of Magnoliaceae and Lauraceae. Unfortunately, we have no
information concerning their occurrence in Asia Minor
during the Cretaceous period. During Oligocene and Mio-
cene, the oaks were widely distributed together with Tert-
iary relics such as Metasequoia and Cercidiphyllum. Some
Cretaceous and Early Tertiary oaks were confused with the
genus Fagus since the foliar remains resemble each other.
Quercus miopontica reported in this paper is very similar
to Q.pontica native to the mountains of N.E.Turkey and
Abkhasia in Caucasus. Q.pontica in the high elevations
of the Caucasian range (1400-2000 m.) and Q.sadleriana
in northern California and southern Oregon (1300-2300m.)
represent the most primitive taxa of the genus as pointed
out earlier by Schwarz. In spite of a disjunct distribu
tion, these two living taxa are very similar with regard
to their habits, vegetative and reproductive structures,
foliar venation and their epidermal ultrastructure; hence
they deserve to be reduced to a single taxon.

I am very grateful indeed to my colleague Hicri Aksoy,
former Undersecretary of the Turkish Ministry of Agricul-
ture who collected excellent specimens of fossil oaks from
the Glven locality and Gergek Sarag, a paleontologist of
the Mineral Research and Exploration Institute (M.T.S.) of
Turkey for accompanying and helping me a great deal during
my expeditions to Giiven of Ankara and to the open lignite
mines of western Anatolia.

I particularly wish to thank my colleague Ismail
Karakan, former Chief of Turkish Forestry Service who pro-
vided a large collection of herbarium specimens of oaks
collected from Turkey. These specimens will be deposited
in the herbaria of the University of California, Davis and
California Academy of Sciences in San Francisco upon the
completion of this work. Since the Turkish Forestry Ser-
vice in Ankara has a complete set of these collections,
the surplus specimens will be distributed to the herbaria
and other institutions or individuals upon the priorities
of the written requests I receive. I would like to thank
Dr. Robert Ornduff, Director of the Univ. Herbarium in
Berkeley, Dr. Daniel Axelrod and Dr. John Tucker (Botany
Dept., U.C.Davis), the staff members of the Botany Dept.
of the California Academy of Sciences and the curators of
the Paleobotanical collections at the Paleontology Dept.
of U.C.Berkeley who kindly allowed me to use their collec-
tions and libraries for comparison and identification

1981 Kasapligil, Oaks of Turkey 99

of my specimens. The hand drawings were prepared by Anne
Crocker, Elizabeth Fall, Anne Jones, Shannon Parr, Lee Ann
Tegart and Angela van Patten, all former students of Mills
College. Credit is given to every artist in the legends
of the illustrations. The photographs of the fossil spe-
cimens and cleared leaves were taken by David Kasapligil
with the assistance of Howard Schorn, by Margret Mukai
and Robin Rickensrud. The microslides of the cleared
leaves were prepared by Joan Amorogo, Margret Mukai and
Ellora Ong. Several other students of mine such as Eliz-
abeth Fall, Sylvia Hsi, Sue Kirkbride, Keli Ryan and Eliz-
abeth Varnhagen were most helpful in preparing the speci-
mens for this study and classifying the research material.
My humble role in the course of this work was being the
conductor of an orchestral team and assuming all responsi-
bilities regarding the factual and speculative statements.

For the sake of convenience and simplicity, partly
due to my own ignorance, the taxa included in this study
are arranged alphabetically covering the extant and fossil
species as well as the hybrid oaks of Turkey. However,
the classical systems of classifications proposed by
Camus and Schwarz are presented synoptically.

Last but not least, I would like to thank my friends
Lita Clapper ‘and Syivia Hsi ‘for a diftiveult task of typing
the manuscript and Dr. Rimo Bacigalupi (Curator Emeritus
of the Jepson Herbarium, U.C. Berkeley) for reading this
introduction.

100 Yom st Oe Oe Loe

Vol. 49, No. 2

A KEY FOR THE IDENTIFICATION OF THE EXTANT SPECIES

OF GENUS QUERCUS IN TURKEY

1.Acorns ripening within the first year
2.Leaves are deciduous
3.Leaf margins lobate
4.Petiole is grooved and twigs are
glabrous
5.Peduncle long, usually longer
than petiole
6.Blades with 9-12 pairs of
lobes, 1/6.te0.1/10, the, breadth
of the blade; no intercalary
Veins; petiole. 10—20mm... cnsic%
6 Blades swith, four to six pabrs
of lobes 1/3 to 1/5 the bread-
th of the blade; has inter-
calary veins; petiole 5-7mm. .
5.Peduncle short or absent

7.Scales of cupule are pubescent

and not tuberculate; inter-

calary veins are not present..
7.Scales of cupule are glabrous

and tuberculate; intercalary

VEINS are. DTECSOCR Es = a's Seuss sie ovene
4.Petiole is ungrooved and twigs are
tomentose
8.Has persistent stipules; peti-
OLeS. 2— SMe MONG « ub écic wdc s «0 «6 = »

8.Stipules not persistent
9.Blades with 4-7 pairs of lat-
eral veins; sinuses 1/3 to 1/5
the breadth of the lamina
10.Petiole 15-25mm; broad obo-
vate leaves up to l6cm. ...
10.Petiole 5-12mm; leaves obo-
vate-oblong, 4-10cm. ......
9.Blades with 7-12 pairs of lat-
eral veins; sinuses 1/6 to 1/5

1.Q.-hartwissiana

2-0.robur

3.Q.petraea

4.Q.polycarpa

»>.9.frainetto

6.Q.vulcanica

7.Q.pubescens

the breadth of the lamina. ...8.Q.macranthera
3.Leaf margins entire or serrate
11.Singly or doubly toothed serrate
Margins; leaves glabrous above,
eliptic-obovate; 10-20 pairs of
PAE al Nd as 6 Je Aang aatas aay ee ----9.90.pontica

3.Leaf margins lobate or serrate or
entire
12.Shallowly lobed or serrate blades
with entire margins at base;
glabrous above, pubescent below

1981 Kasapligil, Oaks of Turkey 101

S= Ll vpairsvot, baterad: veins)... sLOjOeboussaéeri
2.Leaves are semi-decidous
13.Entire at leaf base and lobed to-
wards tip; leaves glabrous, 4-6cm;
peduncle vis .shortor -sessilew /ui11.O.infectoria

l.Acorns ripening within the second year
14.Leaves are evergreen
15.Lower leaf surface is tomentose;
leaf margins are entire or irreg-
ularly serrate
16.Ledves.1.5.to 3 em in length,
oblong-obovate; iregularly

POO CME Gia! «. atutaneccnenshal af ecatyite ee <a e 6) os 12.Q.aucherii
16.Leaves 3-7 cm in length, oblong
=OVaECy TO. FanGeOlate. is cae sdcce £3.QO.3 Lex

15.Lower leaf surface is glabrous;
leaf margins are undulated
17.Spinespoint toward tip of leaf;
leaf margin not cartilagenous;
cupule scales are recurved. ...14.Q.calliprinos
17.Spines spread outward; leaf
Margins are more or less car-
tilagenous; cupule scales are
CONDAGE she aie Ai > wun 0 at a aietal obi ele. = ee 15.9.coccifera
14.Leaves are deciduous
18.Leaves variably lobate
19.Stipules are persistent; 4-7
pairs of lobes; petiole 8-15mm
LONG wate sated elec «. oe. zpaepewane os LG cOncerrius
18.Leaves simple with serrate margins
20.Leaves tomentose with short
DFA SELES «On (ECCEN s) Laneisi cate ose oe L7 36 .beaneri
20.Leaves glabrous
21.Long bristles on teeth; 7-12
pairs of acuminate teeth;
Leaves,.5=10 cm in length. ...18.0.libandi
21.Short bristles; 8-14 pairs
of subaristate teeth; leaves
SET OE PERG EIY on | <a o0) oie aye ara e 19.Q.trojana
14.Leaves are semi-deciduous
22.Leaves are tomentose beneath, ob-
long-ovate, 6-10 cm in length;
margins have triangular aristate

TOBDESS ie iodo 6.5 sae eS wie le ee eee ele leer 20.Q.macrolepis

102 Pen YT, 0°LO'S. DA Vol. 49, No. 2

A synoptic classification of the Turkish oaks according
to the system of A. Camus (1936-38):
[Subgenus Cyclobalanopsis Schneider (cupular scales
concrescent & concentric 77 spp. in East Asia
(Malaysia) not reported from W. Asia, but some
oaks from Turkey with concentric cupular
scales ]
Subgenus Euquercus Hickel et Camus
Section Cerris spach

subsect.: Cocciferae Schott. (Q. coccifera L., Q.
calliprinos Webb., Q. aucherii Jaub. et
Spach.)

subsect.: Macrolepides A. Camus (Q. libani Oliv.,

Q. aegilops ssp. brantii Lindl, Q. aeg.
ssp. macrolepis ky., Q. aeg. ssp.
vallonea ky., Q. trojana Jaub. et Spach)

Ssubsect.: Eucerris Oersted (Q. cerris L.)
Section Mesobalanus A. Camus

subsect.: Ponticae A. Camus (Q. pontica Koch)

subsect.: Macrantherae A. Camus (Q. macranthera

Fisch. et Mey.)
Section Lepidobalanus Endl. (= Subgenus Quercus)
subsect.: Ilex Liebmann-Oersted (Q. ilex L.)

subsect.: Galliferae (Spach) Gurke (Q. infectoria
Of ee

Ssubsect.: Hartwissianae A. Camus (Q. hartwissiana
Steven)

subsect.: Sessiliflorae A. Camus (Q. sessilis Ehrh.

= Q. petraea Liebl., Q. dschorochensis
Koch, Q. pinnatiloba Koch, Q. iberica
Steven, Q. lanuginosa Lam.)
subsect.: Pedunculatae A. Camus (Q. robur L. =
Q. pedunculiflora Koch = Q. haas ky.)
Lsubsect.17: Sadlerianae Trelease (Represented by
Q. sadleriana R. Br. in northern Calif.
& S.W. Oregon, but closely related to
Q. pontica Koch classified under Sect.
Mesobalanus by A. Camus). ]

[subsect.18:

Prinoideae Trelease (Includes Q. prinus
L. = Q. montana Willd., Q. muhlenbergii
Engelm., Q. prinoides Willd. Related to
subsect. 17 and both subsects. related
to subsect. Ponticae under Section
Mesobalanus |]

1981 Kasapligil, Oaks of Turkey 103

The classification of the Turkish oaks in the subgeneric
system proposed by O. Schwarz (1934, 1936-37, 1964).
Subgenus Quercus (= Lepidobalanus (Endl.) Oersted)
Section Roburoides Schwarz
subsect.: Castaneiformis Schwarz
ser. Ponticae (Stef.) Schwarz (Q. pontica K. Koch)
[ser. Sadlerianae Trel. (Q. sadleriana R. Br. in
Caiz€..]
subsect.: Roburiformis (Q. petraea (Mattuschka)
Liebl., Q. p. ssp. iberica (Stev.)
Krassiln., Q. polycarpa Schur., Q. dale-
champii Asch. et Gr.)
Section Robur Rchb. (Q. hartwissiana Stev., Q. robur
Be) :
Section Dascia Kotschy (Q. macranthera Fisch. et Mey.

= Q. syspirensis K. Koch, Q. frainetto Ten. =
Q. conferta Kit., Q. vulcanica Boiss. et

Heldr., Q. pubescens Willd.)
Section Gallifera Spach (Q. boissieri Reut., Q.
infectoria Oliv.
Subgenus Cerris (Spach) Oersted
[Section Suber Spach (Q. Suber L. only under cultiva-
tion in Turkey) |

Section Vallonea Schwarz (Q. brantii Lindl., Q. macro-
lepis Kotschy)

Section Erythrobalanopsis Oerst. (Q. libani Oliv.,

Q. trojana Webb)

Section Eucerris Oerst. (Q. cerris L.) [Q. castanei-
folia C.A. Mey not recorded from Turkey, but
present in Upper Miocene of Ankara |

Subgenus Sclerophyllodrys Schwarz

Section Ilex (Endl.) Oerst. (Q. ilex L.)

Section Coccifera spach (Q. calliprinos Webb., Q.
coccifera L. Q. aucherii)

[Erythrobalanus (Oerst) Schwarz, with 175 spp. from North
and Central America (Q. rubra L, Q. coccinea Muench,

Q. palustris Muench. in cultivation only. |]

104 P Roe TOL. OC LA Vol. 49, No. 2

The Hybrid oaks of Turkey (Based on the available collec-
tions examined and the hybrids reported by Bean, 1976,
Camus, 1936-1954, Karamanglu, 1976, Kasapligil, 1947,
Kotschy, 1867, Menitsky, 1968, 1972;* Rechinger, 1938;
Schwarz, 1934, 1936-1937, 1964; Zohary 1961):

QO. xX auzendi Gren. et Godr. (= Q.coccifera x Q.ilex)
from S.France, introduced into cultivation

in Europe. Possibly, the same as
0. x auene 2a)

Q.boissieri x Q.pubescens
Sterile specimen from a denuded coppice.

Er zincan Prov. |, Arapkirt dasteu, -Datcluca
Subd@ist®r; abt. cas” 300m. (Ahmet Ipictiruk
No. 5 and 34 Adiyaman, Tasliyazi, alt.850m.
of Turkish Forestry Service); Hakkari Prov.
Semdinli Distr., Alt. 1450m in pure stands,
(Cavit Araz NO. $35... Turkish Forestry ser
vice) Malatya-Hekimhan, Akpinar Locality.
(B.. ‘Kasapligil No/'s097 ex A. Tuncer, T.Flaee

0. brantii x Q.boissierd

[Fertile specimens growing in pure stands.
South eastern Anatolia, Diyarbakir Provin-
ce, Lice Distr:|, Hani Subdistr. Takyanos
series, alt. ca. 950m: (Cavit-Araz No. -s
Turkish Forestry Service) ]

O.brantii x Q.vulcanica

(Diyarbakir Prov.:hivee Distr: >) abt. 950m.
(Cavit’Araz ,-T Fis)

Q.calliprinos x Q.coccifera

[B. Kasapligil,(a very common hybrid be-
tween the two closely related spp. in
western and southern Asia Minor; Mersin,
Kuzuncubelen (Kasapligil No. 5107 ex
E.H. Bozakman) |.

O.cerris x Q.infectoria

[B. Kasapligzl No .'5030;, N. Anatelia;
alt. 600m. ]

Q.cerris x Q.libani ‘
[Not to be confused with Q.x libanerris

Boom from Netherlands. Maras Prov. :
Akerda§ (E.K.Balls No. 994) ].

1981 Kasapligil, Oaks of Turkey 105

Q.cerris x Q.pubescens ssp. anatolica
[Common in Anatolia (B.Kasapligil No. 4874,
UC. Davis: No. 6515) 7 "Konya. Prov. :Kadin. Hani,
Yukdur Dastzr. ((B.. KaSapligidl No... 4874)).

Q.frainetto x Q.brachyphylla
[=Q.conferta x Q.brachyphylla tomassinii,
Schwarz, E. Thrace:Istrancadag(Mattfeld
Noe.. (53549). (39Ss Sas 7y04

O.fraanetto x Q.polycarpa
[=QO.conferta x dschorochensis,Shwarz,
E. Thrace: Istrancada§ (Mattfeld No.3657)].

Q.frainetto x Q.pubescens

[=Q0.conferta x Q.pubescens, Shwarz in vic-
inity of Istanbul: Beykoz(DinglerNo. 111),
Uskidar (Krause No. 3200); B. Kasapligil
No. 3384, Zonguldak; Istanbul Prov:
GCatalca, Dumissa 4. one 111i No.-~32)
Zonguldak Prov.: aycuma, (M. G:. Na. 32);
Caycuma, Kilimli eae (Mi Cx Noe Sai:
E.Thrace: Istanbul, Gatalca, DurUusu, alt.
L60: nm. ;. CVedad= Y6nerts. No: 32)'.1

Q.frainetto x Q.vulcanica
Ankara Prov.: ubuk distr., near Karag6ol,
alt. ca. 1500m., on the western slope of
the dormant volcano,(S.Erik No. 479,
Sepec( So; 1973)

Q-hartwissiana x Q.petraea

[Sterile specimen from Trabzon Prov.,
southern slopes of Zigana Pass, Alt. ca.
2000m., (Kasaptigit No. 5281, Seppe. 8,29 78)%

Q.-hartwissiana x Q.petraea ssp.iberica
[Q.armeniaca x Q.iberica, Schwarz,
N.Anatolia: Trabzon (Ky.No.385a); Trabzon:
Zigana Pass, Southern Slopes, Alt. 2000m.
(B.« Kasaphbigéhl, 19 76))<

Q.hartwissiana x Q.polycarpa
[=Q.armeniaca x Q.dschorochensis,Q.pseudo-
dschorochensis, -sessiles var. dschoro-
chensis, Schwarz, N.E. Anatolia:Gimugane
{Schnell in Ky.Rev. No.39)].

106

Q.infectoria

Q.infectoria

Q.infectoria

Q.infectoria

x

Pu ST OL Ove Tui Vol. 49, No. 2

Q.boissieri

Sterile specimens from pure stands of de-
nuded coppice. Eastern Anatolia :Adiyaman
Prov., Tagliyazi village, Harmandere Forest,
Pie. S50) Mm. , mnnet Ipigtrtk No. 34 and 36
Turkish Forestry Service,Oct. 4, 1974.

Q.polycarpa

{Istanbul Prov.:Belgrad Forest (0.Schwarz

in Rechinger)].

Q.pubescens

[Schwarz, B.Kasapligil, ex M.Posat No. ll,

Kocaeli Prov.: izmit; B.Kasapligil No.4875,

Konya Prov.: Kadinhan; B.Kasapligil No.4733,

Izmir Prov.: Bergama, Kapikaya Village].
-robur

[Istanbul Prov.:Ortak6y and Beykoz along
the Bosphorus (O.Schwarz in Rechinger)].

Quercus x libanerris Boom.

A cultivated hybrid between Q.libani and
Q.cerris reported from Rotterdam,
Netherlands

Q.libani x 0. brantii

Fertile specimens from a denuded coppice,
associated with Fraxinus. Hakkari Prov.:
Uzimct Village, Gimen Community forest,
Zapsuyu series, alt. ca. 1200m. (Cavit

Araz No. 8) Turkish Forestry Service,Bing6dl,
Agageli Forest,Alt. ca. 1250 m.(Cavit Araz
No.’9) Turkish Forestry Service.

Q.libani x Q.boissieri

(Kasapligil No. 5090, ex I.Bozakman, Konya
Prov.: Aksehir, Dereli, alt. 500m.)

Q.macranthera x Q.pubescens

Sterile specimen from Hakk@ari Prov.,
gemdinli Distr. ,: ALt. 1750n8., Associated.
with Sorbus. (Cavit Araz No.6)

Q.macrolepis x Q.cerris

[=Q.aegilops x Q.cerris, Schwarz 1934,
N.W. Anatolia: Erenkéy, (Sintenis Nos. 1883

1981 Kasapligil, Oaks of Turkey 107
1196); Troy (Virchow).]

Q.petraea x Q.cerris
Sterile specimen from E. Thrace: Kirklareli
Prov. ¢ “Vize-DisEers', Kizilaga¢ village,
Kiremithane series, Alt. 200m., exp.:East,
in pure stands, 20m. high, @ 30cm.

Q.petraea x Q.pubescens

[Ankara Province, Gubuk Prstr.;') in ene
Vicinity of Karag6él,; alt’ “ca."1550n: en

the volcanic Western slopes, associate with
Populus tremula. (S.Erik No. 480); Sinop
Proyv.: Ayanei2&,; (M- Sahin No. 24); Istanbul.
(Schwarz in Rechinger)].

Q.petraea x Q.robur

[=Q.rosacea Bechst., Kocaeli Prov.:
Adapazari, Ugurlu Village (M. Posat,
Turkish Forestry Service No. 7)].

Q.polycarpa x Q.brachyphylla
[=Q.dschorochensis x brachyphylla tommasinii,
Schwarz, E. Thrace: Istirancadag(Mattfeld
Ho.” 3373, s55F)-¥.

Q.polycarpa x Q.petraea ssp.iberica
[=QO.dschorochensis x iberica, Schwarz;
Q.-pubens Ky. in Sched., Q.pubescens var.
pubens Wenzig, N. Anatolia: Giresun and
Trabzon (Schnell in Ky. Rev. No. 37, 37a.)]

Q.polycarpa x Q.pubescens
[=Q.dshorochensis x pubescens, Schwarz,

in vicinity of Istanbul:Tarabya, Bosphorus
(Dingler No.99), Anadoluhisari (Krause No.
2991), N.W. Anatolia: tnegdl1 (Dingler No.
818)].

Q.polycarpa x Q.robur
[=Q.dschorochensis x pedunculiflara ? ,

Schwarz, E. Thrace: Belgrad Forest in
Istanbul (Mattfeld No. 3172)].

108 Pon YY 20.5.0 6 ik Vol. 49, No. 2

Quercus aucherii1. Jaub et. Spach, Iii. plant. ors, i%
p. 113, t. 58 (1842-1843). Syn.: Coccigera aucherii?
Gandogar, Flora Europae, 21, p. 64 (1890).

Trees up to 14 m. high; trunk diameter up to 60
cm.; crown shape oval. Terminal buds oval, pubescent,
2-4 mm long and stipules deciduous. Leaf blade 4 cm
long, 1-2 cm wide, upper surface pale green-grayish to
green and subglabrous, lower side grayish and pubescent;
shape: whole lamina symmetrical; apex obtuse, mucronate;
base normal obtuse to rounded; margins entire or sinuate
and irregularly toothed with both kinds of leaves some-
times occuring on the same tree; if serrate: apical
side concave, basal side concave to somewhat widely
acuminate; no glands on teeth; sinuses rounded; spacing
regular; serration simple; ours, on upper part of lamina;
cuspidate tip; outline oval-oblong=elliptical; texture
coriaceous, petiole yellowish-brown, pubescent, 2-6 mm
long and not grooved; midrib wavy; pedicel of fruit very
short; fruit, cylindrical, 325, emriong, L0=1L6 am-thick,
purplish brown; cupule bell-shaped, covering 1/3 or 1/4
of acorn, cupular bracts triangular, appressed at base,
free at tips, densely pubescent.

Venation: if margin entire, brochidodromous; if
margin serrate, semicraspedodromous; primary vein stout,
straight to slightly sinuous; secondary veins not par-
allel, directed towards teeth; moderate acute angle to
wide acute toward base of lamina; angle of divergence
40-65 degrees; uniform; moderate width; if margin entire,
curved abruptly (brochidodromous condition): if margin
serrate: curved uniformly (craspedodromous condition) ;
loop-forming branches mostly at right angles, rarely
acute, seldom, composite intersecondaries present; sec-
ondary veins both opposite and alternate along midrib;
tertiary veins: lower ones at right angle or subright,
upper ones at right angle; percurrent: mostly forked,
sometimes simple, sinuous, oblique; angle of tertiaries
decrease towards apex, mostly alternate, often opposite;
higher order of venation: the highest order of vein
5th degree; no excurrent branching, secondaries branch
to form loops; quaternary veins at moderate width,
random; ultimate marginal venation: fimbrial, of
tertiary size; areoles: mostly rounded quadrangular
and triangular shape, small; veinlets none or simple
linear, sometimes curved, rarely branched once.

Distribution: Mainly southwestern Anatolia (see
the map), associated with Pinus brutia, Juniperus and
Ceratonia along the coastal regions.

Localities: B. Kasapligil I. Batat #5075, ex.
Meyer and Pesmen, S.W. Turkey, Aydin Province, Akg¢akonak,
along the road to Pirianne; Kenan Alpacar #9 Turkish
Forestry Service, S.W. Turkey, Antalya Province, Finike
District, Belen Village, altitude 600 m.; K. Alpacar #27

1981 Kasapligil, Oaks of Turkey 109

Turkish Forestry Service, S.W. Turkey, Antalya Province,
Elmali District, altitude 1350 m.70K. Alpacar $54*Turkish
Forestry Service, S.W. Turkey, Antalya Province, Fenike
District, Belen Village, altitude’ 550 mu; Bi Kasapligil
#5076, S.W. Turkey, Mugla Province, between Karabdgirtlen
and Mugla.

Quercus aucherii may represent a hybrid species
between Quercus ilex and Q. calliprinos since foliar
characteristics are intermediate between these two ever-
green species. However, the fruit and cupular charac-
teristics are quite different from those of the possible
parental species as seen in the illustrations at the end.

Quercus boissieri Reut. ex Boiss. Diagn. Ser. 1, 12, p.
119 (1853). syn: Q. infectoria ssp. boissieri (Boiss.)
Gurke) Ris Eurt. 2," py 69" (i897) -r Orbis tani carsspe
orientalis DC., Prod. 16 (2):18-19 (1864); Q. infectoria
Tchihatchef, Asie min. Bot. 2:18 (1864);°Q. petioloris
Boise ee Diagn oil (12): 120° (1853)su0.. tauricolatKotehy

Die Eichen tab. 10 (1862); Q. inermis Kotchy in Unger and
Kotschy, Cypern. 215 (1865); Q. pfaeffingeri Kotschy Die
Eichen tab. 23 (1862); Q. boissieri spp. petiolaris
(Berss.)Schw2. Notizbh! Bot. 7 Gart.. 232177 (2956)47 Oe
araxina: (Trautve) iGrossh. Fi. eaucs.. 22238 7(1930)<

Deciduous trees up to 12 m in height with a dia-
meter of 60 cm. Young twigs glabrescent; at maturity
covered with white, solitary hairs. Terminal buds ovoid,
brown, glabrous, bud scales oblong and stipules per-
sistent. Leaf blade oblong-elliptical, size: 1.5-10 cm
long and 1.5-4.3 cm wide, grayish green and glabrous
above, yellowish green and puberulent below and micro-
phyllous to mesophyllous; shape: whole lamina symmet-
rical, base mostly symmetrical; normal oblong to narrow
obovate, sometimes narrow to wide; apex: acute to
obtuse, apical lobe acute to obtuse; base mostly rounded,
sometimes auriculate; margin: entire to widely crenate
or serrate, 1/3-1/2 to apex; apical side of teeth
straight, basal side convex; no glands on teeth apices;
sinused rounded; spacing regular; simple venation; extent
of serration variable; apex cuspidate. Petiole yellowish
brown, 5-10 mm long and not grooved. Midrib straight.
Fruits almost sessile, cups obovate or hemispherical,
bracts appressed and prominently tuberculate, densely
pubescent; acorns cylindrical-ovate, halfway enclosed
within cupule, mucronate at apex, approximately 4 cm in
length.

Venation: if margin entire of crenate-brochi-
dodromous, if margin serrate-craspedodromous; primary
vein; moderate to stout; straight, sometimes sinuous
toward apex; secondary veins; if margin entire: wide
acute to right angles; if margin not entire: narrow to

110 P8272 0.1.0.8 es Vol. 49, No. 2

moderate acute; upper secondaries more acute than lower
secondaries; moderate width; if margin entire or crenate:
curved abruptly, if margin serrate: straight; where
loop-forming branches exist, joined at right to obtuse
angles; rare, composite intersecondaries from midvein in
serrate condition, frequent, simple intersecondaries in
entire margin; secondary veins not parallel; directed
toward teeth with an angle of divergence of 31-61 degrees;
6-12 pairs secondary veins either alternate or sub-
opposite along midrib; tertiary veins from lower to
right angles, upper ones at right angle; percurrent:
mostly forked, sometimes simple straight; if margin
entire: tertiaries close to parallel to midvein, if
margin serrate: oblique, tertiary angle decreases up-
ward; alternate; higher order of venation: the highest
order -- 5th degree, sometimes 6th degree; no excurrent
branching; quarternary veins--moderate, orthogonal;
quinternary veins--moderate to thick, orthogonal to
random; ultimate marginal venation fimbrial, or tertiary
size; areoles: irregular, often quadrangular; small;
veinlets usually linear to branched twice, often more
than one veinlet per areole; sometimes none.
Distribution: Mainly S.E. Anatolia, but some in
S.W. Anatolia as well as along the Mediterranean coasts
of Southern Anatolia. Mostly in denuded coppices
associated with Pinus brutia, Juniperus spp., Pistacia,
Daphne, Sorbus, Platanus, Ulmus, Quercus calliprinos,
On cerris, 0.) Liban, Acer hyrcanum, A. obtusifolium,
Ostrya carpinifolia, Fraxinus ornus, Pyrus syriaca,
Ostrya carpinifolia. Q. boissieri usually occupies semi
arid and arid regions of the Middle East extending
southward to Lebanon, Syria and Israel and from eastern
Anatolia through northern Iraq, it reaches Transcaucosus
and northern Iran. It also occurs in western Cyprus.
hocalitiess’ Cc. Araz #21) Torkach Fus., EE. funkey;
Bing6l Province, Agaceli Village, Yayladiizti Arg Forest,
altitude 1250 m: pAw Ipictiruk. 47 T.F.Si,) B.) Turkey,
Malatya Province, Arapkir District, Dutluca Subdistrict,
altitude’ 1200 m.; A. Ipi¢ctirtk $31 T.F.S.,.E. Turkey,
Malatya Province, Hekimhan District, Akpinar locality,
attatude +1000) m:.> Cas Arnaz, $501. PS... S. Es: Turkey,
Diyarbakir Province, Lice District, Taykanus ruins,
altitude 850 m.; A. Gdéksin #12, S. Turkey, Adana
Province, Osmaniye District, altitude 800-900 m.; A.
Ipiciirtik #25 T.F.S., E. Turkey, Malatya Province, Pdétirge
District, Bozkir Village, altitude 1000 m; A. Ipiguritk
#19 T.F.S., E. Turkey, Malatya Province, Doganyol
District, G6k¢e Village, altitude 900 m.; C. Araz TiF.a6
#18, E. Turkey, Bingé6l Province, Uzundere Village,
Beluzar Forest, altitude 1350 m.; A. Ipigtirtik #35 T.F.S.,
E. Turkey, Adiyanam Province, Tasliyazi Village,
Haramidere Forest , altitude 850 m.;'C. Araz #1..T.F.S,%

1981 Kasapligil, Oaks of Turkey 5 I

E. Turkey, Elazig Maden, altitude 900 m.; C. Araz #2
Ti .S.40- Eb. Purkey, Siare’ Province, Sirnak: District;
Senola’ Village, Sivridag, altitude: 1000 m:; CC. Araz™ #5
T.F.S:) S.Be* furkey, Batlis: Province, ‘Narlidere’ District,
Bocan Village, Dedetan Series, altitude 900 m.; H.
Pesmen- A. Giiner #1924, S. Turkey, Isparta Province,
Eqridir District,. altitude: 650-1200 'm.\57 Kh. Alpaca: #24
fT PeS.5' 6. Wi. Turkey ~Mugta: Prevince, Koyeeqiz District,
Cayhisar: Village; altitude S00 m.; K.Alpacar #23) T.F.S.,
S. Turkey, Antalya Province, Finike District, Aykirig¢ay,
altitude -700.m.>.K.° Alpacar’ 421," T.FS.7°Ss Turkey,
Antalya Province, Finike District, Belen Village,
altitude. 600 ms;>K. Alpacar #422 T.F. Si) SW. Turkey,
Mugla Province, Kdycegiz District, Gdkbel Forest,
altitude 270;m.: CC.) Arazeelo Tre. Si4) E.aTurkey, Bingol
Province, Yumakia Village, Geng Series,” altitude. £230
my 706. Araz#13°T. PiS., Bes Turkey," Hakkari) Province,
Beytis sebap District, Besagag Village, altitude 1650 m.;
GC) Araz’$3°0.F.S:.,. Bf Turkey, Diyarbakix Province,
Kokulu Pinar locality, Hani District, altitude 900 m.;
he. WpLleurdk #2 TOF.S:,: EB. Turkey, Erzincan., Prevince,
Ekrek Village, altitude 1400 m.; Pesmen #3354, E. Turkey,
Bitla's* Province,. Tatvah. District. KK. Alpacar gos Ti. Ss,
S.W. Turkey, Finike District, Belen Village, altitude
S00. ny. A: Goksin ¢1lGa T.P.S.7°S. Turkey, Adana Province,
Osmaniye District, Nurdag, altitude 830-1000 m.;
K.© Alpacar’ #33 T:F.S.,°S. Turkey; Antalya Province;
Akseki-Murt, Kabiz Road, altitude 450 m.; K. Alpacar
#32. T.F.S., S. Turkey, Antalya Province, Akseki District,
Kuyucak Subdistrict, altitude 1150 m:> K.-Alpacar gil
T2F.S., SiWe Turkey; Antalya’ Province; Finike:- Disteict,
Aykirigay subdistrict, altitude 700 m.; C. Araz #3
TFS. , E.y Turkey, Elazigqg Province,’ Maden, Districe,
alei tuderL020> mi}. Ci Araz- #4 T.FUS.), Bev Purkey aoatece
Province, Sirnak District, Kizilsu Series, altitude
750 m.7) C.. Araz 46 TiP.S.5- Bt! Turkey) Bitlis: Peevince,
Narlidere District, Bocan Village, altitude 910 m.; A.
Ipictirtik #20 T.F.S., E. Turkey, Malatya Province,
Doganyol District, G6k¢ce Village, Memnundere Forest,
abtitude* 900m.) ‘A. Tpicuruk #1). T.F.s-, B- Turkey,
Malatya Province, Ovledik Gegidi District, on the slopes
of Keban G6lii, altitude 1450 c.; B. Kasapligil #5091
S. Turkey, Adana Province, Kozan District, 18 km. from
Buruk Village, altitude 300 m.; B. Kasapligil #5099,
S. Turkey, Mersin Province, Camilli Village.

Quercus boissieri is related to Q. infectoria
and Q. pubescens. The intermediate forms between these
species are common. In fact, Camus (1937) and Menitski
(1968,1971) consider Q. boissieri as a subspecies of Q.
infectoria. It also hybridizes with Q. brantii
producing fertile hybrids. Its wood is used mainly for
fuel and charcoal production. Unfortunately, most of

112 PY F..0.. Lf Gt TA Vol. 49, No. 2

the coppices are in denuded condition due to overgrazing
and human pressure. It is a hardy oak suitable for re-
forestation. Q. boissieri is a beautiful tree with
grayish-green foliage suitable for landscaping. Although
most of the naturally growing ones are in shrubby

habits, they do have tremendous potentiality to reach
enormous sizes under protection.

Quercus brantii Lindl., Bot. Reg. 26 suppl. 41 (1840).
Syne 105) -persicavaub;t-et.Spach. Pi. Or. Spbentabsr Ss
(1842-1843); Q. brantii ssp. persica Jaub. et Spach.

Oye Sebhwarz,” Notizbl.. Bot-«: Gart.. Berlin, 1375 416: 20, (1936)—
Q. baneica Djavanchir. Les chenes de l1'Iran 123 (1967);
Q. globularis Djav. Les chenes de l1'Iran 127 (1967); Q.
SaaaiDaqy es! Abid l43) (4.967 } po Ou! UNGeray/ Diay. 2pad.25t
(2862)>°0O.. aegilops: ssp. branti1i Lindl.» 1936=38.. | Az
Camus Monogr. du Gen. Quercus 1:544 (1934).

Deciduous tree up to 15 m high. Twigs densely
tomentose; greenish brown. Buds ovoid, 3-8 mm long,
pubescent, scales ovoid and stipules persistent. Leaf
blade ovate or oblong-elliptical, 2.9-11.4 cm in length,
1.6-6.8 cm wide, upper surface dark green, lower surface
gray green, both sides densely tomentose, whole lamina
symmetrical, base symmetrical to slightly asymmetrical;
from-narrow elliptic to mostly wide elliptic; apex acute
to obtuse, mucronate; base cordate to sub-auriculate.
Margins serrate, apical sides of teeth concave; basal
Side widely acuminate, sometimes spiny; without glands
on teeth; sinuses rounded; spacing regular; simple
serration; sometimes margins deeply lobed or entire
towards base; apex sinuate; base rounded or somewhat
auriculate. Petiole yellow, 0.7-2.8 cm long, pubescent
and not grooved. Midrib straight, rarely wavy; number
of lateral corresponds to number of lobes, usually 8-11
pairs; intercalary veins faintly visible. Cupules hemi-
spherical or conical, with triangular or filiform scales
spreading or recurved (Figs. A and C); acorns, 3-5 cm
long, 12-22 mm thick, 1/3 of basal portion enclosed
within cupules. Apex of fruit flat or convex with
mucronate tips.

Venation: craspedodromous; primary vein:
moderate width; straight to curved; secondary veins:
narrow acute to wide acute (moderate acute at middle
area); upper secondaries slightly more acute than lower
secondaries; more or less straight, sometimes uniformly
curved; no loop-forming branches; no intersecondaries
off of the midvein, but rarely branching from a second-
ary to join a tertiary from an opposite secondary;
tertiary veins: if blades narrowly elliptic in shape
lower ones at right angles, if widely elliptic: lower
ones at wider angles; percurrent: mostly forked, some-

1981 Kasapligil, Oaks of Turkey 113

times simple sinuous; oblique, angle of tertiaries
decreases in upper leaf area, but otherwise generally
remains uniform; alternate and opposite; the highest
order of venation,5th degree; no excurrent branching,
intersecondaries from secondaries may join tertiaries;
quarternary veins at moderate width, orthogonal; quin-
ternary veins thin, mostly orthogonal, but often
reticulate; ultimate marginal venation fimbrial, of
tertiary size; areoles: mostly aguadrangular, often
irregular; medium; veinlets mostly simple, often with
more than one veinlet in each areole, often branched
once.

Distribution: Mainly in Southeastern Anatolia
(see figure 4.).

Localities?) .C: vAraz: £9 TP. Bsa seer Turkey;
Diyarbakir Province, Egil Subdistrict, Boyali Village,
abEi tude: 850 «m..¢¢C. Araz #l0eT -F:S:g7S. EB.) Turkey; Mardin
Province, Mazidag District, Dasot Village, altitude
L050em.7 -CscAraz, $43; S.E. | Turkey, Diyarbakir ‘Province,
Dakyanus Hani District, 60 km. north of Diyarbakir,
altitude 950 m.; A. Ipiciiriik #48 T.F.S., S.E. Turkey,
Fistik6ézii Village, Halfeti, Oleturdu, denuded coppice,
attitude .800: ma7.C: Arnaz 43, ‘2PeS2pe8s Turkey ,;.cSi25t
Province, Sirnak District, Kizilsu Forest, altitude
$50) mic C. eAreaz. 4? Tu. Ss, ‘S:Ee furkey, -Diyarbakze
Province, Cinar District, Sipyak Village, Bo6lcinar
Porest » altitude 850i m.; C. ArazZ #4) T.F.S.5¢S.82 Turkey,
Bitlis Province, Narlidere District, Bocal Village
altitude 900 m.> Cs. Araz.$2 TaeFiS. > Bestarkey, Mardin
Province, .Idil; subdistrict, -altitude :/50.m.; C.2Araz
#2 T.F.S., E. Turkey, Mardin Province, Midyot District,
Yolagz1 Forest, altitude 1000 m.; C. Araz #6 T.F.S.,
S.E. Turkey, Diyarbakir Province, Ciingiis subdistrict,
Cermik Forest, altitude 900 m.; C. Araz #4 T.F.S.,

E. Turkey, Diyarbakir Province, Kokulu Pinar Hani
subdistrict, altitude 960 -m.; C..Araz, 48 T-E.Ss, E.
Turkey, Bingd6l Province, Agageli District, Arg Forest,
aber bude: »b250. me:;':

This species is related to Q. macrolepis in
western Anatolia and Q. ithaburensis in Syria, Israel
and Jordan. It hybridizes with several other species
such as Q. libani, Q. boissieri, Q. vulcanica; these
hybrids are described under a separate heading. Q.
brantii extends eastward through the mountain steppes
of eastern Turkey to the Kurdish mountains of Irag to
southwestern Iran forming denuded coppices between the
elevations of 700-1800 m. It is usually mixed with the
typical elements of the Irano-Turanian steppes.
Commonly, it is associated with trees and shrubs such as
Pistacia spp., Olea, Ulmus, Amygdalus, Crataegus, Pyrus,

aU

114 Pe eT Obes Fs Vol. 49, No. 2

spp. The most common oaks associated with Q. brantii
are Q. boissieri and Q. libani forming savannah-1like,
denuded forests with loosely scattered small trees.
Most of my specimens are collected from small trees
4-8m. high and their diameters at breast height ranging
from 6 to 35 cm. This polymorphic species has many
subspecies and varieties which are beyond the scope of
this paper, but the interested readers are referred to
Zohary's (1961,1973) and Djavaros' (1967) publications.
The wood of Q. brantii is widely used for fuel
and charcoal production. Under human and grazing
pressures, most of the mixed-broadleaved deciduous
forests of Q. brantii are highly denuded. Occasionally,
mightly specimens are found in cemeteries by the grave-

yards of the notables and "holy individuals". This is
a hardy and drought tolerant oak with beautiful foliage
and crown on a stout bole. Indeed, it is a valuable

tree which deserves attention in landscaping of arid
and semi-arid lands. As far as I know, there are no
cultivated specimens of this oak in the Mediterranean
region, nor in California. Unfortunately, there is
not enough information in literature regarding its
sylvies, seed germination and ecological requirements.
It certainly deserves botanical as well as horticul-
tural research for furthering our knowledge about this
oak.

Quercus calliprinos Webb. Iter. Hisp.:15 (1838).

Syn. Q. pseudococcifera Labill. Icon. Pl. Syr. Rar.
529/ Pl. 6: QO. fenzlii Ky. Eichen Eur. u. des Oriénts
Bu 173-74, “pli 24 (1862)% ‘Ov palaestina Ky: “ibid; ps "S8=
59, Pl. 19 (1862); Q. coccifera var. calliprinos (Webb)
Menits. Gence Quercus L. Nov. Syst. Akad. Nauk, USSR,
921326 002972).

Small trees up to 8 m. high; some larger speci-
mens reaching 1 m. in diameter. Twigs brown and
puberulent. Terminal buds single or clustered, round-
ovate, brown, 1-4 mm. long, more or less tomentose,
scale shape ovate and stipules deciduous. Leaf blade
1.5-5 cm. long, microphyllous, 0.9-2.5 cm. wide, upper
surface yellowish green, more golden below, both sides
glabrous, outline mostly oblong to lanceolate, rarely
ovate, margins not cartilaginous, dentate with sharp
teeth oriented towards the apex, base frequently
oblique, apex cuspidate; whole lamina, symmetrical, base
cordate and sometimes rounded. Margins-entire or
serrate. If serrate: apical side concave to straight;
basal side concave to straight; glands, rounded, some-
times blunt on basal area; sinuses rounded; spacing
regular; serration simple; on complete margin. Petiole
golden, pubescent, 2-4 mm. long and not grooved. Midrib

1981 Kasapligil, Oaks of Turkey 115

generally straight. Cupules greenish brown, involu-
cral bracts 4-10 mm. long, oblong-linear, appressed

at base only, variably curved, but not spiny at tips;
acorns dark brown, roundish-oblong, rarely pubescent,
1-1.3 cm. long, 1.2-1.6 cm. thick, mucronate at apex.

Venation: if margin‘entire-brochidodromous, if
Margin serrate-semi-craspedodromous, each secondary
which enters a tooth arises independently from the
midvein and is not a result of secondary branching
(compare with Q. coccifera); primary vein: moderate to
stout; straight to curved, sometimes zig-zag; secondary
veins sometimes parallel, often branching further; veins
terminating at teeth; 6-9 pairs; secondary veins oppo-
site or alternate along the midrib; divergence moderate
acute to wide acute or at right angles; upper second-
aries more acute than lower ones; thin to moderately
thick; subsinuous to sometimes recurved at lamina base;
loop-forming branches join mostly at right angles,
sometimes acute; rarely composite intercalary veins
present; tertiary veins: at lower right angles, upper
ones at right angles; percurrent: mostly forked, rarely
simple sinuous; running longitudinally at basal lamina
area or oblique, tertiaries at decreasing angles upward
direction; mostly alternate, often opposite; higher
order of venation: the highest order--5th degree; no
excurrent branching, secondaries branch to form loops;
quarternary veins-thick, orthogonal; quinternary veins-
moderate, random; ultimate marginal venation-fimbrial,
of secondary size; areoles: mostly rounded quadrangular
or triangular shapes; small; veinlets-mostly none or
simple linear or curved, sometimes branched once.

Localities: A. Gd6ksin #11 Turkish Forestry
Service, S. Turkey, Maras Province, Andirin District,
altitude 1075 m.; A. Gdksin #10 T.F.S., Andirin District,
altitude 1075 m.; B. Kasapligil #4864, W. Turkey, Izmir
Province, Alasehir Forest District, in pure stands along
foothills’ K. Alpacar #42 T.F.S., S. Turkey, Antalya
Province, Akseki District, Akseki-Manavgat Road in the
cemetery altitude 940 m.; A. Gdksin #18 T.F.S., S. Turkey,
Adana Province, Osmaniye District, Nurdag, altitude 830-
1000 m.; K. Alpacar #1 T.F.S., S. Turkey, Antalya Province,
Biik Research Forest, altitude 500 m.; K. Alpacar #7
T.F.S., S.W. Turkey, Marmaris, between Mugla and
Marmaris, altitude 130 m.; K. Alpacar #6 T.F.S., S.
Turkey, Antalya Province, Kas District, Kalkan sub-
district, along the roadside between Kalkan and Fethiye,
altitude 162 m.; B. Kasapligil #4870, S.W. Turkey, Aydin
Province, Sdke District, Kizilisik Village.

Quercus calliprinos is the predominant species of
macchie occurring widely in the eastern sector of the
Mediterranean basin. It is closely related to Q.
coccifera. As a matter of fact, Boissier (1879),
Schwarz (1934 & 1964) and Menitsky (1972) consider it

116 PRY T OL OH LT A Vol. 49, No. 2

as a variety or subspecies of Q. coccifera. On the
other hand, DeCandolle (1864), Camus (1936-1938), Wenzig
(1887) and Zohary (1961) recognize specific rank for
it. Intermediate forms between Q. calliprinos and Q.
coccifera are quite common in Asia Minor. However, the
distinction between the two species are thoroughly in-
vestigated by Zohary (1961). Q. coccifera is primarily
a small, shrubby, evergreen oak in the western section
of the Mediterranean basin while Q. calliprinos often
has arborescent habit, predominantly occurring in the
eastern section of the Mediterranean basin. Under pro-
tection, they form sizeable trees 8-10 m. high with a
stem diameter of 40-100 cm. Hybridization and intro-
gression is noticeable especially in western and south-
western Anatolia where the distributional areas of both
species overlap each other. Quercus calliprinos grows
successfully especially in calcareous soils of southern
Anatolia associated with Pinus brutia, Juniperus
oxycedrus, Daphne oleoides, Styrax ofEicinalis;) -Cistas,
Pistacia, Platanus, Sorbus spp. and wild olives. In
western Anatolia, it reaches an elevation of 1400 m.,
while along the Taurus mountain range it grows at the
elevation of 1500 m. or even higher. The range of
distribution in the south of Turkey extends to Lebanon
and Jebel Duruz in Syria (Mouterde 1953, 1966), from
Safad to Hebron in Israel (Zohary 1960), from Irbid to
Salt and from Tafila to Shaubak and north of Wadi Musa
in’ Jordan” (Kasapligil 1956 a,b,c). Q. calliprinos is
the most dominant shrub or small tree of the east
Mediterranean macchie. The readers are referred to the
publications of Zohary (1960, 1973) for the ecological
and geobotanical relations of this species. Q. calli-
prinos is a highly polymorphic species with regard to
its habit, morphology and to the great variety of
edaphic and climatic habitats of its natural range.
Zohary (1961) recognizes seven varieties in the Middle
East which are distinguishable through their leaves,
fruits and cupules.

Q. calliprinos is a hardy species, cultivated in
England. Apparently, the legendary 'Abraham's Oak' in
Hebron belongs to this species (Bean 1976). It is
considered a sacred tree by the followers of all three
religions of the Holy Land, hence no one dares to remove
a twig from the aged specimen. However, the open
forests of scattered trees are denuded throughout the
Middle East since the wood of this species is used
widely for fuel and charcoal production.

Quercus GCerris Le'sSpi Pl. 997 (1753). °Synv-On -austradaca
Willd. Sp. Pl. 4:454 (1805); Q. tournefortii Willd.
Sp. Pl. 4:453 (1805); Q. tukhtensis Czecott, Acta Soc.

1981 Kasapligil, Oaks of Turkey 7

Boty «PB. 9244) (1932) 2"0O.> Lanuginosa: Lam: 7''Os CeEris ‘ssp.
austriaca Willd. Schwarz.

Deciduous tree up to 35 m. high forming pyramidal
or broadly open crowns. Twigs rough, glaborous,
reddish-brown hairy and linear stipules persistent. Buds
ovoid, 1-4 cm. long and bud scales oblong. Leaf blade
oblong to narrow obovate or ovate, 4.2-13.8 cm. long,
3.2-6.8 cm. wide, deeply lobed usually 1/3-1/2 to midrib,
dark green hairy, and rough to touch above, grayish
green and pubescent below; whole lamina symmetrical or
asymmetrical, with alternate 4-7 pairs of entire or
dentate lobes, base slightly asymmetrical; apex obtuse,
apical lobe acute to rounded, sometimes retuse or
pointed; base mostly normal obtuse to subcordate, some-
times one side decurrent. Margin-lobate, with alternate,
often very deep lobes: apices mostly rounded to rarely
bluntly pointed, if pointed possibly with a gland;
sinuses rounded; spacing regular; simple and often
compound series; on entire margins. Petiole yellow-
green, pubescent, 0.8-1.8 cm. long (moderate) and
proximal portion grooved. Midrib generally straight.

Venation: craspedodromous; primary vein:
moderate to stout; straight to curved; secondary veins:
terminating at lobes; rarely branching further; narrow
to moderate acute angle, sometimes wide acute in basal
area; upper secondaries more acute than lower second-
aries; moderate to thick; straight to uniformly curved
upward in apical area, downward in basal area, some-
times branching as a secondary or intersecondary to
form a secondary lobe (branching occurs on the lower
side of a secondary); no loop-forming secondaries,
except at base when joining tertiaries; frequent inter-
secondaries, simple, from midvein in non-lobate areas,
sometimes from lower side of a secondary to form a
secondary lobe; tertiary veins: lower ones at right
angle, upper ones also at right angle, sometimes both
angles acute in lobate areas; percurrent: mostly
forked, rarely simple straight to sinuous; oblique,
the angles of divergence in tertiaries, usually decrease
towards apex, predominantly alternate; higher order of
excurrent branching--2nd degree; quarternary veins-
moderate, orthogonal; likewise quinternary veins
moderate, orthogonal; ultimate marginal veins fimbrial
of tertiary size; areoles: irregularly shaped, mostly
medium, often small; veinlets none or simple linear or
branched once. Fruit maturing in the second year,
acorns (Fig. 8, C) light brown, 2.5-3.6 cm. long,
mucronate, rarely concave at apex; cupules 1.6-2.4 cm.
in diameter, 1.5-1.9 mm. deep, enclosing the nuts half
way, cup scales linear-filiform, appressed at base,
upper scale portions loosely arranged and recurving
often.

118 PAE Gti TL Vol. 49, No. 2

Distribution: Western sector of the Black Sea
coasts of Turkey, throughout Anatolia, except the
eastern part. Not recorded from the Saltlake region of
Central Anatolia. Very common along the Istranja
mountains of eastern Thrace, but absent in the remain-
ing areas of European Turkey possibly due to intensive
agricultural land use (see Fig. 9).

Localities: A. Gdéksin #13 T.F.S., S. Turkey,
Adana Province, Osmaniye District, altitude 800-1000 m.;
A. Aldemir’.#35 T.F-S., oN. Turkey, ‘Bolu- Province,

Mudurnu District, altitude 620 m.,; A. Aldemir #34,

Bolu Province, Mudurnu District, Gdlctik Series, altitude
500 miz Ke Alpacar #36 T.F.S., SS. Turkey, Antalya
Province, between Akseki and Manavgat; A. fpiciirtik #43
T.F.S., E. Turkey, Adiyaman Province, G6lbasi District,
abetiude.850.m.>..K. Alpacar. #i4..T.F.8., S20. Turkey,
Mugla Province, Yaras District, kiyit Series, altitude
1020¢m.; .K.“Alpacar #12 -T. FS.) We Turkey ;fDeniziz
Province, Acipayam Road, altitude 950 m.; K. Alpacar

#35 T.F.S., S. Turkey, Antalya Province, Manavgat
District; saltitude 450 m.:)V..Yonellii , $6. Por 7S. zs
Thrace, Istanbul Province, Catalca District, altitude

e0 mL; °K. -Alpacar $34 T.F.S.,\'S. Turkey; Antalya
Province, Akseki District, Murt igi, Kabiz Road along
the creek, altitude 650 m.; V. YOnelli #5 T.F.S., E.
Thrace, Istanbul Province, Gatalca District, altitude
160 m.; A. Ipigctirik #47 T.F.S., E. Turkey, Adiyaman
Province, G6lbasi Forest, altitude 850 m.; C. Aksoy,

A. Suner, Y. Dugarslan #4 T.F.S., N. Turkey, Tokat
Province, Niksar District, Cami¢i Village, Immidogan
locality, altitude 950 m.; K. Alpacar #46 T.F.S., S.W.
Turkey, Mugla Province, Fethiye District, Cengerk6oy,
Hatice ana‘ hill, Kuz Locality, altitude. 250m. ; Av
Aldemir #32 T.F.S., N.W. Turkey, Bolu Province, Mudurnu
District, Ilica Village, Sarot Forest, altitude 440 m.;
A. Aldemir #33 T.F.S., N. Turkey, Bolu Province, Mudurnu
District, Yiirse Series, altitude 480 m.; M. Posat #2
T.F.S., N.W. Turkey, Kocaeli Province, Adapazari District,
Kemaliye Village’.,M. Posat #23 T.F.S., N.W. Turkey,
Adapazari Province, Akyazi District, Karaptirgek Locality;
M. Sahin #35 T.F.S., N. Turkey, Samsun Province, Kavah
District, Mahmutlu Villages M. Sahin #32 T.F.S., N.
Turkey, Samsun Province, Bafra District M. Sahin #29
T.F.S., N. Turkey, Sinop Province, Gerze District, Hatin
Village, Dihmen Forest; V. Y6nelli #46 T.F.S., N.W.
Turkey, E. Thrace, Kirklareli Province, Vize District,
K6mlurk6y Series, altitude 150 m.; Vedat Y6nelli #57
T.F.S., E. Thrace, Kirklareli Province, Vize District,
Kizilagag Village, Kiremithane Series, altitude 200 m.;
V. Y6nelli #61 T.F.S., E. Thrace, Kirklareli Province,
Vize District, Kocatas Locality, altitude 10 m.; V.
Y6nelli #88 T.F.S., E. Thrace, Kirklareli Province,

1981 Kasapligil, Oaks of Turkey 119

Demirkd6y District, Uzunbacak Locality, Karacadag Series;
V.. Yonelli $85-T.F.S., :E...Thrace, Karklareli: Province;
Demirkd6y District, Istihkamtepe Forest, altitude 350 m.;
VV; VYonelii.#283.T.F.S.,. BE.) Thrace; Karklareli Provinee;
Demirkéy District, Karayokus Locality, altitude 500 m.;
VY. Yonelli. #24 T.F.Si; Es. Thrace, Kiarkiarel1; .Provance;
Demirkéy District, Sarapnel Series, altitude 475 m.;

Ve Yonetii £80 -T.F.S.> EB. Thrace; Kirkiareli Province;
Demirkéy District, Isletme Locality, altitude 425 m.;

Vu: Yonetli 476) T:Fos.;, 2... Thrace, Kirkilarel). Provance;,
Derek6y Region, Karlik Series, altitude 430 m.; M.

Posat #3 T.F.S., N.W. Turkey, Kocaeli Province,
Adapazari District, Sogucak Village; M. Sahin #21 T.F.S.
N. Turkey, Sinop Province, Ayancik District, Bakirli
zaviye Village, A. Ipigtirtik #32 T.F.S., E. Turkey,
Malayta Province, Hekimhan District, Akpinar Forest,
altitude, 1000: m::°0M. Posat #21)T.F.S., _N.W.. Turkey,
Adapazari Province, Akyazi District, Merkez Subdis-
Eeice, -Beldibi,.Locality;.V. ¥6nelli.$73. ToFs5<54Es
Thrace, Kirklareli Province, Merkez Region, DUuzorman
Series, altitude 500 m.; A. Ipigcitiriik #24 T.F.S., E.
Turkey, Malatya Province, Doganyol District, Gdokg¢e
Village, Memonun Forest, altitude 900 m.; M. Posat #8
T.F.S., N.W. Turkey, Kocaeli Province, Adapazari District,
Ugurlu Village, M. Posat #17 T.F.S., N.W. Turkey, Izmit
Province, Adapazari, Sosesi; A. Gdksin #9, T.F.S., se
Turkey, Maras Province, Andirin District, altitude

1075 m.; A. Gdksin #8 T.F.S., S. Turkey, Marag Province,
Andirin District, Baskonus locality, altitude 800-1000
m.; K. Alpacar #39 T.F.S., S. Turkey, Antalya Province,
Akseki District, 3 km. along Ibradi Road, in cemetery,
altitude 940 m.; B. Kasapligil #4620 S. Turkey, Burdur
Province, Bucak District, Gerce Village; M. Posat #13
T.F.S., N.W. Turkey, Kocaeli Province, Liitfiye Locality;
B. Kasapligil #5796, E. Thrace, Igneada, Longozlar
Forest, H. Pesmen-A. Giiner #1486, S. Turkey, Isparta
Province, Egridir, Yaka kdyti, altitude 1500-1700 m.;
M.G. #34 T.F.S., N.W. Turkey, Zonguldak Province, Kozlu
District; A. Aldemir #29 T.F.S., N.W. Turkey, Bolu
Province, Gdyniik District, altitude 1380 m.; M.G. #40
T.F.S., N.W. Turkey, Zonguldak Province, Devrek District,
Dirgine, Kazdere Forest; A. Aldemir #31 T.F.S., N.W.
Turkey, Bolu Province, Mudurnu District, Ilica Village,
Sarot Forest, altitude 460 m.; A. Ipigiirtk #41 T.F.S.,
S.E. Turkey, Adiyaman Province, Gdlbas1 Forest, altitude
850 m.; V. YOnelli #32 T.F.S., E. Thrace, Istanbul
Province, Catalca District, Durusu subdistrict, altitude
160 m.; K. Alpacar #18 T.F.S., S.W. Turkey, Mugla
Province, Yerkesik District, Kocadtiz locality,
Narcgalidag Forest, altitude 550 m.; V. Yonelli #62,
T.F.S., E. Thrace, Kirklareli Province, Vize District,
Yumurtatepe Series, Kocatag Locality; A. G6ksin #15,

120 Peart OL Ove. fA Vol. 49, No. 2

T.F.S., S. Turkey, Adana Province, Osmaniye, Nurdag,
altitude 830-1250 m.; V. Y6nelli #10 T.F.S., E. Thrace,
Istanbul Province, Catalca District, Yalikdéy Bdélgesi,
Kozuldere Forest, altitude 20 m.; M.s Sahir #28, T.F.S.,
N. Turkey, Sinop Province, Cerkez Village, Sinop Forest;
B. Kasapligil #4840, S. Turkey, Antalya Province,
Giingdodmus District, Vicinity of Senir Village; B.
Kasapligil #4841, S. Turkey, Konya Province, Aksehir
Tekke Village, altitude 1300 m.; B. Kasapligil #4859,

N. Turkey, Sinop Province, Ayancik, altitude 600 m.;

B. Kasapligil #4867 E. Central Turkey, Afyon Province,
Sandikli District, Burgaz Mountain; B. Kasapligil #4874
S. Turkey, Konya Province, Kadin Hani District, Yukdur
Subdistrict; B. Kasapligil #4877, S. Turkey, Konya
Province, Ilgin District, ¢igil Subdistrict; B.
Kasapligil #5028 N. Turkey, Izmit Province, between
Akmese and Imaniye; B. Kasapligil #5031 N.E. Turkey,
Izmit Province, Yuvacik; B. Kasapligil #5032, S. Turkey,
Adana Province, Osmaniye District, Gavurdagi, altitude
1000 m.; B. Kasapligil #5035, S. Turkey, Adana Province,
Kozan Forest; B. Kasapligil #5077, S. Turkey, Antalya
Province, Manavgat Forests; B. Kasapligil #5078 N.
Turkey, Corum Province, Hacilar hani region; B.
Kasapligil #5109 E. Turkey, between Malatya and
Hekimhani, Akpinar locality.

; Q. cerris is the most widely distributed oak in
Turkey (Fig. 9). In Europe, it extends from the Balkan
Peninsula through Central Europe to France. In the
Middle East, south of Turkey, Q. cerris grows along the
coastal regions of Syria and Lebanon. In the U.S.A.
and Great Britain, it is cultivated in parks and ar-
boretums. It is a quick growing and hardy species
suitable for streets especially in cold climatic zones.
Several varieties are listed tentatively by Zohary (1961).
The natural hybrids of Q. cerris with Q. libani in
southern Anatolia and with Q. infectoria in northern
Anatolia are recorded. However, Q. x libanerris Boom
(1959) described from a cultivated specimen in
Trompenburg Arboretum in Holland is not represented in
my collections from the natural populations in Turkey.
Karamanoglu (1976) enumerates two subspecies from Turkey.
Q. cerris ssp. austriaca (Willd.) Schwarz reaches the
elevation 1500 m. in northern Anatolia between
Kastamonu and Cankiri (Krause Nos. 2419 & 2424). Q.
cerris ssp. tournefortii (Willd.) Schwarz is reported
occurring between the elevations of 750-900 m. in
Bolkardagi of the Taurus range in Mersin (Kotschy Nos.
386, 405), c& Karamanoglu (1976). Both subspecies
occur in steppic, arid regions as well as in mesic
broadleaved forests of the Black Sea coasts of Asia
Minor and eastern Thrace. Unfortunately, high forests
of pure stands are rare, except in Osmaniye district

1981 Kasapligil, Oaks of Turkey TAE

of Adana Province, I observed in 1944, a seedling forest
of magnificent tall trees mixed with Pinus nigra in
Hagbel Frenk locality at the elevation of 1220 m.
Depending on the vegetation type of its natural distrib-
ution areas in Turkey, Q. cerris associates with a large
variety of trees and shrubs: Pinus brutia, P. sylves-
tris, 0 Pibany;”0.- 1nfectoria, OO. vuilcanica, Castanea
sativa, Fagus orientalis, Laurus nobilis, Carpinus,
Fraxinus, Acer, Arbutus, Platanus, Populus, Phillyrea,

eee ee OO——EEEeE

Cornus, Paliurus, Mespilus, Sorbus, Melia, Styrax,

ee Oe Oe eee

a

Erica; Smilax, etc.

Along the coastal mountain ranges of the Black
Sea coast, Q. cerris is quite an invasive species forming
coppice type of forests following the clear cutting of
broad-leaved high forests. It has a tremendous capacity
of regeneration through suckers and easy establishment
of young seedlings. The wood is not desirable for con-
struction purposes, but it is used mainly as fuel wood.
Under human and grazing pressures, most of the coppices
are in denuded state. Being related to Q. suber = Cork
oak, Q. cerris = Turkey oak is classified under Subgenus
Cerris (Spach) Orsted by Schwarz (1964). In fact,
hybridization between the two has been reported from
western Europe. Like Q. suber, Q. cerris also is
suitable for cork production. During the World War II,
Turkey was unable to import cork from Spain and Portugal.
The Turkish Forestry Service commissioned Dr. Kazim
Mih¢ioglu to investigate and find a substitute for the
COrk.. During early 1940s, Dr. Mihgioglu settled with
Q. cerris after a long survey of cork producing trees
in Turkey. He established several experimental lots in
southeastern Turkey where the primary cork from the barks
of aged trees were ax-peeled, punched and used as cork
stoppers for bottled beverage industries. The primary
cork was too brittle and hard enough to resist the cork
screws. However, a few years later, the secondary oak
developed from the cork cambium was soft, elastic, and
pliable. The foresters of the Osmaniye (Adana Province)
called it "female cork" to distinguish from the brittle,
porous "male cork" obtained from the primary barks of
the trees. The early reports of the cork production
from Q. cerris barks were published by Dr. Mih¢ioglu
in the 1942 volume of "Orman ve Av = Forestry and
Hunting" journal of the Turkish Forestry Association.
At the end of the World War II, Turkey was able to
import cork again and consequently the experiments on
cork production from the barks of Turkey oak were dis-
continued.

No doubt, the best use of this species would be
in landscaping the streets and parks in the urban areas.
In 1959, I planted three seedlings of Q. cerris at the

122 P.8.Y.,T 0.4.0.6 TA Vol. 49, No. 2

International Grove of the University of Washington in
Seattle during the 5th International Forestry Congress.
Twenty years later, when I visited the University of
Washington campus again, I located the two survivors
which were about 15 m. high with vigorous branching from
Single boles (Fig. 10). According to Bean (1976), it
has been in cultivation in Great Britain since 1735.

Quercus: Cocc i fera) is, Sp. -P1.2995 (1753) 2. “Syne 0.
pseudococcifera Desf., Q. coccifera var. pseudo-
coccifera (Desf.) A. DC., Ilex coccigera Clus.

Small evergreen shrubs or trees 203 m. high.
Twigs grayish brown; pubescent. Terminal buds more or
less glabrous, ovoid dark brown, bud scales ovoid and
stipules persistent. Leaf blade ovate to wide ovate,
rarely oblong, 1.7-5 cm long (usually about 2 cm.),
microphyllous, 1.2-2.3 cm. wide, dark green and shiny
above, pale green and glabrous below; whole lamina
symmetrical, base symmetrically cordate or rounded;
apex acute, mucronate. Margins--in ours serrate, with
abruptly divarcating spines, more or less cartilaginous
with spiny teeth not directed towards the tips of
leaves, often radiating outward directions, apex
cuspidate, base rarely oblique; if serrate: apical
side concave, basal side straight, no glands on teeth;
sinuses rounded; with regular spacing; simple serration;
Or occasionally complete margin. Petiole yellowish
brown, pubescentat young stage, soon becoming glabrous,
1-6 mm. long. Midrib straight; rarely wavy. Fruits
matruing in two years, concave and mucronate at tip,
half way or 2/3 of acorn enclosed within cupule. The
scales of the cup radiating in all directions, quite
spiny. Fruits develop singly on short stalks, usually
abundant.

Venation: semicraspedodromous, where secondary
branching occurs almost midway between the margin and
midvein, each of the 2 branches enter adjacent teeth;
primary vein: stout; straight to subsinuous; secondary
veins: mostly medium acute to wide acute angle, tending
to right angle and recurved toward base of lamina;
upper secondaries at a more acute angle; moderate width;
straight and sometimes branching, lower secondaries re-
curved; loop-forming secondary branches at acute angle
or tight angles; seldom composite intersecondaries;

4-7 pairs of secondary veins terminate at teeth;
secondary veins either opposite or alternating along

the midrib; tertiary veins: lower ones at wide or right
angles, upper ones at right angle; percurrent: mostly
forked, rarely simple, sinuous; oblique, tertiary angle
decreases upward, lower tertiaries sometimes parallel

to midvein; alternate and opposite; higher order of

1981 Kasapligil, Oaks of Turkey 123

venation: the highest order--5th degree; highest order
of excurrent branching--2nd degree; quarternary veins--
thick, orthogonal; quinternary veins--moderate, random;
ultimate marginal venation fimbrial, of secondary size;
areoles: mostly rounded quadrangular or triangular
shapes; small; veinlets mostly none or simple linear,
sometimes curved.

Distribution: Mainly southeastern Europe and
North Africa, Mediterranean regions of Greece and Turkey.
Localities: B. Kasapligil #4682, W. Turkey,
Izmir Province, Menemen District, Emiralem, foothills
of Dumanli Dag, altitude 50 m.; B. Kasapligil #4625
S. Turkey, Konya Province, Ermenek District, Merkez
subdistrict, B. Kasapligil #5105 W. Turkey, Manisa
Province, Aksehir Village, altitude: 1050 mo;5' Kx Alpacar
#26 T.F.S., S.W. Turkey, Mugla Province, Kdycegiz
District, Yarg1 Village, altitude 1300 m.; K. Alpacar
#19 T.F.S., S.W. Turkey, Mugla Province, Yerkesik
District, Mar¢galidag Forest, Karahayit locality,
altitude’ 600 m7 K.« Alpacar: #20) T. FeS.y 'S.W.! Turkey,
Mugla Province, Gdkbel Village, Sarisu Forest,
atta pudec2 75) m. 3K. Alpacar:q4) T.F.S.)) Se urkey,
Antalya Province, Akseki District, Akseki-Manavgat
Road near the cemetery, altitude 940 m.; K. Alpacar
#10 T.F.S., S.W. Turkey, Mugla Province, Kavaklidere
Village, Road between Mugla-Aydin, altitude 350 m.;
Mehmet Posat #10 T.F.S., N.W. Turkey, Kocaeli Province,
tzmit District; B. Kasapligil #5070, W. Turkey, between
Izmir and Manisa; B. Kasapligil #5082, S.W. Turkey,
Mugla Province, Fethiye District, Karadere Village;
B. Kasapligil #5102 S.W. Turkey, Antalya Province,
Bukbeli Districé,' altitude: 650) mis, 8.) hasapligil
#5103, S.W. Turkey, Antalya Province, Finike District,
altitude 500-600 m.; B. Kasapligil #5104 N.W. Turkey,
Canakkale Province, Eceabat District, Sarikiz Forest,
altitude 850 m.; B. Kasapligil $5106, S. Turkey; deel
Province, Mut District, KarabagG subdistrict;),.>) ku.
north of Mut, altitude 1120 m.; B.. Kasapligil $5107,
S. Turkey, Mersin Province, south of Kuzucubelen,
altitude 625 m.; B. Kasapligil #4835, S.W. Turkey,
Denizli Province, Bozdogan #4839, S. Turkey, Antalya
Province, Gtindogmus District; B. Kasapligil #4841
Konya Province, Aksehir Forest District; B. Kasapligil
#4856, N.W. Turkey, Bursa Province, Gemlik District;
B. Kasapligil #4862 N.W. Turkey, Edirne Province,
Kesan District; B. Kasapligil #5048, S. Turkey, Adana
Province, Kozan Forest; B. Kasapligil #5050 W. Turkey,
Manisa Province, Spil Mountain; B. Kasapligil #5051,
S. Turkey, Mersin Forest.

Q. coccifera commonly known as Kermes oak is a
predominant elemngt of the macchie in western and
southern Turkey (Kasapligil 1952). The vertical range

124 Py Eit.T @.LcO06 2h Vol. 49, No. 2

of the Kermes oak reaches an altitude of 1400 m. in
western Anatolia (Karamanoglu 1976). It is often
associated with Pinus nigra, P. brutia, Quercus cerris,
Arbutus unedo, A. andrachne, Juniperus oxycedrus, Olea
europaea, Phillyrea media, Paliurus spinachristi and
many other Mediterranean elements. Under grazing
pressure it becomes a creeping shrub which is typical
in Phrygana vegetation in Kocaeli peninsula of north-
western Anatolia (Kasapligil 1947). Under protection
or in cultivation, they become sizable trees about
6-7 m. high. The largest specimen in Turkey measures
8 m. in height with a stem diameter of 35 cm.

Kermes oak is used primarily for fuel and char~
coal production. The wood is dense and very hard;
that is why it is called Steineiche in German and Tag
mesesi (Stone oak) in Turkish. The name Kermes oak
is derived from the Kermes insect (Chermes ilicis)
which lives on the barks of this oak. The female in-
sects were the source of "grana chermes", an important
dye material during the Middle Ages (Baytop 1963). The
leaves and fruits are very variable since it hybrid-
izes freely with Q. calliprinos and Q. ilex in nature.
It is a beautiful oak with shiny green, spiny foliage
resembling Ilex aquifolium. Although several forms of
Kermes oak are cultivated in Western Europe, I have
never seen them under cultivation in Turkey and in the
United States. For the infraspecific classification of
Q. coccifera the readers are referred to the monograph
Gf Vaeroso’ (1950):.

1981 Kasapligil, Oaks of Turkey 125

A provisional list of Miocene oaks of Turkey and their
possible relationships to the extant species.

Quercus boissieri Reut. ssp. pesmenii, ssp. nov.

Very close to modern Q.boissieri, leaves
obovate, serration sharply pointed towards
apex.

Quercus drymeia Unger
Related to Q.sartorii Liebm. from Mexico.
Lanceolate blades with sparsely serrate
Margins, lamina base often rounded. Midrib
and petiole strongly developed.

Quercus furcinervis (Rossm.) Heer
It may belong to Castanopsis. lanceolate
leaves with craspedodromous venation;
lamina tips often curved.

Quercus cf. hartwissiana
Cupule 2 = 2.5 cm. wide with tuberculate
scales.

Ouercus’ heidingeri Ettingh.
Be has dite. to Q.ilex but leaves are much
larger.

Quercus kodorica Kol.
Blade margins shallowly lobed. Midrib and
petiole strongly developed.

Suercus Kubiny. “UKRov.) Cz.
Affinities to Q.macrolepis, O-Libanx,
Q.trojana; most common oak of Tertiary
period found in many fossil deposits of
central Europe. See the distrubution map
by H. PTraiau.;

Quercus lonchitis Ung.
Lanceolate or obovate leaves with entire

Margins.

Quercus mediterranea Ung.
Aff. with Q.ilex. Base of lamina entire,
upper 2/3 of lamina coarsely serrate.

Quercus mioaxelrodii sp. nov.
Aff. with Q.castaneifolia + lanceolate

leaves. Midrib zig-zag, blade margins
serrate.

126

Quercus

Quercus

Quercus

Quercus

Quercus

Quercus

Quercus

Quercus

Pe ET Oe Oe Vol. 49, No. 2

miopontica sp. nov.
Aff. to Q.pontica from N. E. Turkey and
AbDkihasia. U.S .5.8.., ana to Q.sadleriana
R. Br. Campst. from N. California and
Southern Oregon.

miovariabilis Hu et Chaney
Roughly resembles chestnut leaves, it is
identical with the Chines Miocene species
described from Shantung Prov. of China.
Palaeont. Sinica, ser. 412, mew Ser. 2,
Dat 36-3715. Dhevsl Dien tee Be Bel

neriifolia A. Braun
It may be related to Q.imbricaria Michx.
from East. U.S.A. . Narrowly lanceolate
leaves with entire margins, petiole well
developed.

paulmouterdei sp. nov.
Aff. to Q.-libani and Q.regia. Cupules are

3-3.5 cm. wide, scales triangular, pro-
minently overlapping.

sclerophyllina Heer.

Aff. with Q.coccifera =Kermes oak. Petiole
short and thick, midrib strongly developed,
leaf margins sharply dentate, pointed out-

wards.

semecarpifolia fossils

Aff. with Q.sosnovskyi. Leaves 6-7cm. long,
2.5-3 cm.wide, midrib and petiole strongly
developed. Secondaries diverging at broad

angles. 2utt of hairs. at angles.

seviraedii A. Br.
Aff. with Q.phelios of Eastern U.S..

sosnovskyi Kol. f.angustifolia Kol.

It has aff. to Q.glauca Thunb. from Japan.
Blader 12-14 cm. long, lanceolate with en-
tire margins. The leaves are quite vari-

able.

1981 Kasapligil, Oaks of Turkey 127

Pitre Wms Bay D S.vF Ors DL bb, Bis eA Tot. ees
rigure. 1s

Quercus aucherii Jaub et Spach A) A flowering branch
with staminate catkins x 1/2 drawing by Anne Crocker;
B) Contact prints of two cleared leaves, to the left:
with entire margins, to the right: partly dentate
blade showing the brochidodromous venation in the
lower portion of the lamina and semicraspedodromous
condition in the upper portion of the leaf. C) Detail
of the minor venation and ultimate areoles, x 30, Cl -
areole, C2 - ultimate linear veinlet, C3 - fimbrial
vein along the leaf margin; Kasapligil #5075. D)
Fruit variation x 1/2, drawings by Elizabeth Fall.

Dl - Bell-shaped cupule with imbricate bracts enclosing
1/4 of the slender acorn; D2 - A hemispherical cupule
with adpressed scales enclosing 1/3 of a thick acorn.
Note the mucronate tips of the acorns. Specimens
provided by Kenan Alpacar #54 from Belen village of
Antalya Province.

Figure 2:

Quercus boissieri Reut. A) A fruiting branch showing the
variation of leaf outlines. Although most of the leaves
are obovate, some of them are oblong-elliptical.

Drawing by Anne Crocker. B) Acorns with mucronate
apices. Note the prominently tuberculate cupule with
short pedicel. C) Detail of two leaves with obtuse
apices and cuneate bases. D) Contact prints of two
cleared leaves showing the craspedodromous venation in
the upper dentate-lobate portions of the leaves and
brochidodromous venation in the lower blade portions
with entire margins. All illustrations x 1/2.

Figure 3:

Distributions of Q. boissieri Reut. (circles), Q. libani
Oliv. (solid circles) and Q. trojana Webb. (triangles).

Figure 4:

Distributions of Q. brantii Lindl. (circles) and Q.
macrolepis Kotschy (solid circles).

Figure 5:

Quercus brantii Lindl. A) Fruiting branch showing the
leaf variation and the young stage of fruit development.
Specimen from Silvan south of Mus, T.F.S. #14. Note
that the cupular bracts are more or less filiform at an
early stage. Drawing by Anne Crocker. B) Contact
print of a cleared leaf showing the detail of major

128 PB Yor Oa, Oreo a, A Vol. 49, No. 2

venation, B. Kasapligil #4626. Note that the upper
secondaries diverging at narrow angles and terminating
at marginal teeth while the lower secondaries near the
base of lamina diverging at wider angles and reticulo-
dromous where blade margins entire. The tertiaries are
oblique in relation to the primary vein. Slide and
print by Margret Mukai. C) Variation of mature fruits
and cupules. Basal involucral bracts are triangular,
imbricately appressed, but their tips are free. The
upper scales near the cupular margins are recurved.

The specimen to the left: C. Araz #6 T.F.S., from
Diyarbakir Province, Fermik Forest; The specimen to

Ene Metes* As ipaguruk Fol TlE.S., ee. Luckey,
Fistik6zii Village, Halfeti loc. Both drawings by Elizab.
Part. “Alt ~LTigqures x 1/2.

Figure 6:

Ay) Fruveing branch of*Q. calliprinos KRasapligitt

#4864. Note that the teeth of the serrate leaves are
pointed towards the leaf apex. The involucral bracts
of this particular specimen are not recurved and that
the cylindrical acorns are halfway enclosed within the
eupules. “Drawing by Anne "Crocker, x 1/2. “B)_..Contace
print of a cleared leaf with entire margins showing the
brochidodromous venation pattern in which the bifurca-
ting secondaries forming loops. C) Another leaf
clearing from the same specimen with dentate margins
and semi-craspedodromous venation. Note that the teeth
in the lower half of the leaf are pointed ourwardly and
that the secondary veins diverge at narrow, right and
broad angles. The intercalary veins are evident in
both leaf samples (Kasapligil #4864). The slides and
prints of B & C by Margret Mukai, X 2.

Figure 7:

Quercus calliprinos, details of leaf vasculation,
Kasapligil #4625. A) Contact print of a serrate leaf
with strongly developed primary vein and semi-craspedo-
dromous secondaries alternating with intercalary veins.
The teeth are spinescent and pointed toward the leaf
apex, x 2-1/2. B) A secondary vein terminating at the
ie Of a marginal tooth. \ 2°, 3° 49% 5°; veine-or the
2nd-5th degrees respectively, mf: marginal fimbrial
vein. C) Bifurcation of a secondary vein near the
blade margin giving rise to semicraspedodromous condition
(bs), (ar): Areoles with compactly arranged palisade
parenchyma cells. Photomicrographs by Margret Mukai,
Kies

1981 Kasapligil, Oaks of Turkey 129
Figure 8:

Quercus cerris = Turkey oak. A) A fruiting branch with
persistent stipules and an immature fruit enclosed with-
in the cupule covered by long, filiform bracts.
Kasapligil #4620, drawing by Anne Crocker, x 1/2; _ B)
Contact print of a cleared leaf showing the detail of
major venation and the frequency of intercalary veins.
Note that some of the lobes are coarsely dentate.
Kasapligil #4620. Slide of leaf whole mount by Joan
Amoroso, contact print by Margret Mukai, x 1/2; C)
Variation of the mature fruits showing the detail of the
cupular bracts. A. Aldemix #32, “F-E2S., Bolu’ province,
Mudurnu (left), A. Ipigtruk #46, T.F.S., Adiyaman
province Gdlbas1 (right). Both drawings by Elizabeth
Baeble sel fz

Figure 9:

Distribution of Quercus cerris (Turkey oak) in Anatolia
and Eastern Thrace.

Figure 10:

Quercus cerris (Turkey oak): A) Contact print of a
cleared leaf showing a deeply dissected blade with
simple lobes almost reaching the midrib region. Note
the lobes and the secondary veins are alternating.
Again the intercalary veins are common. See the text
for the detailed description of the major and minor
venation. Hasan Pesmen & Giiner #1486, slide and the
contact print by Margret Mukai x 1/2; B and C)
cultivated specimens of Turkey oak in the International
Grove of the University of Washington, Seattle. The
trees were planted by this author in 1959 for commem-
orating the 5th International Congress of Forestry on
behalf of the Turkish Government. The photographs
were taken twenty years later on August 10, 1979 by
B. Kasapligil.

Figure ll:

Quercus coccifera L. Kermes oak. A) A fruiting branch
showing the variation of the leaves and acorns in the
same individual plant. Kasapligil #4625, drawing by
Anne Crocker, x 1/2; B-C) Contact prints of cleared
leaves showing the detail of the major and minor vena-
tions. Note the remarkable resemblance of the foliar
vasculature in Q. coccifera and Q. calliprinos (Figs.

6 and 7). Kasapligil #4625, both slides and prints by
Margret Mukai, x 2.

130 Pe TT Oi Ore TB Vol. 49, No. 2
Figure 12:
Distributions of Q. aucherii Jaub et Spach (circles),

Q. calliprinos Webb. (triangles), Q. coccifera L.
ilex L. (solid circles).

(solid Triangles), and Q.

1981 Kasapligil, Oaks of Turkey 132

SEED a gts a Bee
PY ery Via
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Fig.4. Quercus aucheri

Vol. 49, No. 2

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Kasapligil, Oaks of Turkey 133

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Kasapligil, Oaks of Turkey

1981

136 Peet OL Oe: LA Vol. 49, No. 2

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1981 Kasapligil, Oaks of Turkey 137

Vol. 49, No. 2

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Kasapligil, Oaks of Turkey 139

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Kasapligil, Oaks of Turkey 141

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1981 Kasapligil, Oaks of Turkey 143
REFERENCES

Batat,,, EL... 2978
Analysis of morphological variation in Quercus
pubescens Willd. ssp. anatolica Schwarz. Ph.D.
Dissert.; UC’ Davis, Davis, California

Baytop,, T..1963
Tiirkiyenin tibbi ve zehirlibitkileri; Istanbul
Univ. Publ. No. 59. Istanbul.

Bean, W.J. 1976

Trees and shrubs hardy in the British Isles.
8th ed. revised by D.L. Clarke, vol. 3, Quercus
p. 454-521, illustr. John Murray Publ. Ltd.,
London.

Boom 1959
Q. x libanerris Boom 1959 in Jaarb. Nederl.

pendrol., Ver: , No: 21,: py 1SE,s, fig. 4d.; Pp:
P59.

Camus, A. 1936-1954
Les Chenes: Monographie du genre Quercus,
1-3, atlas and text, Paris.

Engler, A. (Revised by H. Melchior 1964)
Syllabus der Pflanzenfamilien. 12th. ed. vol.
2, Quercus:50-51. Gebriider Borntraeger, Berlin.

Ettinghausen, C.F. 1895
Uber die Nervation der Blatter bei der Gattung
Quercus mit besonderer Beriick-sichtigung ihrer
vorweltlichen Arten. Denkschr. Akad. Wiss.,
Wien, math. natw. 63:117-180.

Hutchinson, J. 1967
The genera of flowering plants. Vol. 2:Dicotyl-
edons. Clarendon Press, Oxford.

KaramanoGlu, K. 1976
Tiirkiye Bitkileri. Vol. 1, Quercus p:57-70.
Ankara tiniv., Eczacilik Fak. No. 32, Ankara.

144 PE rh Oe. OC Tak Vol. 49, No. 2

Kasapligil, B, 1947
Kuzey Anadoluda Botanik Gezileri (Botanical
excursions in northern Anatolia). Orman Genel
Mudurlugui Yayinlarindan No. 32, Istanbul.

1952
The forest vegetation in the Mediterranean
regions of Turkey. Istanbul Univ., Orman

Fakiiltesi Dergisi 2 (2):47-65. Istanbul

1956a
A report on the vegetation profiles of the
forest and grazing Lands in’ Jordan. 24 pp:
+ 1 map,” +’ 6 profiles: Ministry ‘of Economy,

Amman.

1956b
Plants of Jordan with notes on their ecology
and economic uses. 130 pp. + a veget. map

1750; 000.7 Dept.) of “Forests, Ministry of
Agric., Amman, Jordan.

L956¢
Report to the Government of the Hashemite
kingdom of Jordan on an ecological survey of the
vegetation in relation to forestry and grazing.
Bolt slg. OLE A-O. Report. Now 549), 39° pp. +
2 maps, + i prorate.” FJA.0O. of the’ U.N. 7° Rone:

Komarov, V.L. (edit.) 1936
Elera Of the U-s.5.k.;, Vol. 5, Quercus i. ,
pp. 322-354, treated by V.P. Maleev., Akad.
Nauk SSSR, Moskva-Leningrad.

Kotschy, Theodor 1862
Die Eichen Europas und des Orients. Eduard
H6lzel's Verlag, Wien and Olmuz.

Krussmann, G. 1962
Handbuch der Laubgehédlze. vol. 2:2, p. 295-
322, Verlag Paul Parey, Berlin & Hamburg.

Menitsky 1968
Conspectus speciorum Caucasicarum subsectionum
robur (Reichb.) Maleev et Galliferae (Spach)
Maleev generis Quercus. Nauka, Moscow.

1972
Generis Quercus L. species Asiae Austro-
Occidentalis Novosti sistematiki vysshikh
rastenii. Akad. Nauk SSSR, vol. 9:105-140,
Leningrad.

1981 Kasapligil, Oaks of Turkey 145

Mihcioglu, K. 1942a
Tiirkiyede Sacgli meseden mantar istihsaline
daily ‘bir arastirma. “Orman ve Av? vod. 25: 151,

Ankara.

1942b
Gavurdaginin Sa¢gli mese ormanlarinda yapilan
yerli mantar istihsali. Orman ve Av, vol.

L5e187-. Ankara:

Mouterde, P. 1953
La Flore du Djebel Druze. Universite Saint
Joseph, Beyrouth.

1966
Nouvelle Flore du Liban et de la Syrie. Vol. l,
Editions de L'Imprimerie Catholique, Beyrouth.

Rechinger, Karl Heinz (filius) 1938
Enumeratio Florae Constantinopolitanae.
Fedde's Repert. spec. nov. regni veget. Beih.
Vol. 98:1-73. Dahlem, Berlin.

Schwarz, O. 1934
in K. Krause's Beitraege zur Flora Kleinasiens,
G. Die in der Tiirkei vorkommenden Baeume und
Straeucher: Quercus. Fedde's Repert. spec.
nov. reg. veget. 33:321-338, Dahlem, Berlin.

936-37
Monographie der Eichen Europas und des
Mittelmeergebietes, 2 vols. Fedde's Repert.
spec. nov. reg. veget. Sonderbeiheft D, text:
200 pp., atlas :64 pls., Dahlem, Berlin.

1964
in T.G. Tutin and W.H. Heywood (editors)
Flora Europaea, vol. 1:61-64.

Stojanoff, N. & B. Stefanoff 1929
Beitrage zur Kenntniss der Pliozanflora der
Ebene von Sofia.
Zeitschrift Bulg. Geol. Geselsch. 2,3

Vicioso, Carlos 1950
Revision del Genero Quercus en Espana. Minist.
de Agric., Direccion Gen. de Montes, inst.
Forestal de Investig. y Experiencias. 21 (51):
1-194, Madrid.

146 PEt) EO le OG 2. A Vol. 49, No. 2

Zohary, M. 1960
The Maquis of Quercus calliprinos in Isreal

and Jordan. Bull. of Research Council of
Israel, ad. (2) iSi-72;

1961
On the Oak Species of the Middle East. Bull.
Res. Council of Israel 9D (4) :161-186.

1966
Flora Palaestina. The Israel Acad. Sci. and
Humanities, Jerusalem. Text vol. 1, p. 30

Quercus L. p:30-36.

L973
Geobotanical Foundations of the Middle East.
2 vols., Gustav Fischer Verlag,: Stuttgart.

ADDITIONS TO THE LITERATURE CITED

Boissier, E. 1879
Flora Orientalis, Vol. 4, H. Georg, Genevae et
Basileae.

De Candolle, A.P. 1864
Prodromus systematis naturalis regni vegetabilis.
16-2, Paris.

Bitener, David LL. 1974
Approaches to the identification of angiosperm
leaf remains. Bot. Rev. 40(1):1-157.

Hickey). i002) (4973
Classification of the architecture of dicotyledo-
nous leaves. Amer. J. Bot. 60(1):17-33.

Zohary, M. 1962
The Plant Life of Palestine (Israel and Jordan)
Chroenica Bot., NcS,.P1i. Sei. Books 33(6):262pp.,
Ronald Press Co., New York.

CERTAMEN MELASTOMATACEIS XXXIII.

John J. Wurdack
U. S. National Herbarium, Smithsonian Institution

With this installment of novelties, all of the easy
taxonomic decisions have been completed in the outpouring of
recent collections from Bahia, Brazil. However, much more work
is needed, particularly in Microlicia, to present a reasonable
treatment of the Melastomataceae for this still poorly known
region.

LAVOISIERA HARLEYI Wurdack, sp. nov.

L. nervulosae Naud. affinis, foliis 3-5-nervatis non
reticulato-venosis stylo glabro differt.

Ramuli primum tetragoni demum teretes primum sicut folia
sparsiuscule glanduloso-setosi (pilis 1-2 mm longis) demm
glabrati; nodi dense glanduloso-setulosi (0.2-0.3 mm). Folia
sessilia imbricata; lamina 20-35 X 10-20 mm elliptico-oblonga
apice rotundato basi late acuta vel obtusa, firmula et essentia-
liter integra, ad margines cartilaginea et glanduloso-ciliata,
3-5-nervata nervis secundariis nervulisque invisis. Flores 6-
meri; hypanthium (ad torum) 10 m longum usque ad medium densi-
uscule glanduloso-setosum (pilis ca 1.5 m longis); calycis tubus
ca 1 mm longus, lobis ca 3 mm longis basaliter expansis extus
sparse glanduloso-setosis glanduloso-ciliolatis intus glabris.
Petala 34-35 X 16-19.5 mm ad margines apicem versus modice gland-
ulosa (0.1 mm) extus ad basim sparsissime glandulosa alioqui
glabra. Stamina dimorphica glabra; filamenta 13-13.5 mm vel 11.5-
12 mm longa; connectivum 10 mm vel 3 m prolongatum ventraliter
ad basim in appendice 3 X 1 X 1m vel 2 X O.8 X 0.8 mm hebeti
productum. Stylus 12.5 X 1.3-0.4 mm glaber; ovarium 6(?)-locu-
lare apice glabro leviter intruso.

Type Collection: R. M. Harley 19690 (holotype CEPEC 19692;
isotypes K, US), collected on northeast face of summit ridge,
Pico das Almas ca 25 km WNW of the Vila do Rio de Contas, Bahia,
Brazil, elev. 1600-1800 m, 19 March 1977. "Bushy shrub to 2 n.
Leaves viscid, rather bright green. Petals bright pinkish
magenta. Anthers golden-yellow.”

Paratype (topotypical, fruiting): Mori 12455 (CEPEC, US).

The suggested relative (isotype US) has leaves 7-9-nerved
(at the base) and markedly reticulate-venose beneath, as well as
somewhat longer calyx lobes and basally glandular-setulose style.
Of the other species in this affinity with moderate-sized leaves,
L. selloana Cogn. differs (ex descr.) at least in the much
smaller flowers and glabrous hypanthia, while L. cogniauxiana
Mello Barreto (ex descr. and photo) has larger 9-nerved leaves
and 8-merous flowers. To L. nervulosa, I have referred (with

some reservations) two collections (Mori 12551, Harley 18788)
147

148 Fr Rat 204.0 6:2 A Vol. 49, No. 2

from just south of Mucuje; both have sparser pubescence than
Blanchet 3333 and petals yellow with the exposed part externally
red in bud (rather than wholly pink). All of the examinable
flowers and fruit of L. harleyi were 6-merous; the anther thecae
were all insect-destroyed.

MICROLICIA HATSCHBACHII Wurdack, sp. nov.

M. balsamiferae (DC.) Mart. affinis, ramulis setulosis
staminum maiorum connectivis plus prolongatis differt.

Ramuli primum obscure tetragoni mox teretes modice vel
densiuscule et subpersistenter glanduloso-setulosi pilis 0.2-0.4
mm longis. Petioli 0.5-1 mm longi subtus sparsiuscule glandu-
loso-setulosi; lamina (0.7-)1-1.5 X (0.4-)0.6-0.8 cm elliptica
apice hebeti-acuto basi acuta, rigida et integra, ubique dense
glanduloso-punctata, supra glabra, subtus glabra vel secus costam
basim versus decidue glanduloso-setulosa, eciliata, 3-nervata.
Flores in ramulis terminales 1-3, pedicellis ca 2 mm longis sicut
hypanthiis basim versus sparse glanduloso-setulosis. Hypanthium
(ad torum) 4.5-5.5 mm longum teres glutinosum; calycis tubus 1.3-
1.5 mm longus, lobis 1.8-2 X 2.4-2.8 mm deltoideis hebeti-acutis.
Petala glabra 16-17 X 9-13 mm asymmetrice obovata apice late
obtuso. Stamina paulo dimorphica, filamentis 6-7.3 mm vel 6-6.5
mn longis, antherarum rostris obscure (0.5 mm) evolutis. Stamina
maiora: thecae 4.4-6 X 0.8 X 1.1 mm; connectivum 1.8-2.5 m pro-
longatum, appendice ventrali O.6-0.7 mm longa hebeti. Stamina
minora: thecae 4-5.3 X 0.7 X 1 mm; connectivum 0.9-1.5 m pro-
longatum, appendice ventrali O.4-0.5 mm longa hebeti. Stigma
non expansum; stylus 8.5-12 X 0.5-0.2 mm in ovarii apicem O.4-
O.5 mm immersus; ovarium 3-loculare glabrum apice rotundato-
lobulato. Semina 0.7 X 0.4 mm areolata.

Type Collection: G. Hatschbach 39696 (holotype MBM 48802;
isotype US), collected at Rio Agreste, Mun. Morro do Chapéu,
Bahia, Brazil, 17 Jan. 1977. "Arbusto muito ramoso, 1 m 70, flor
rosada. Solo arenoso, junto a afloramentos de arenito."

Paratypes (all Morro do Chapéeu region, Bahia, Brazil):

Hatschbach & 0. Guimaraes 42423 (MBM, US), Rodovia BA-052 8 km
natscnbach wuUTmMaAracs
from Morro do Chapeu, elev. 900 m ("Arbusto ramoso, copado,
70 em, flor rosada"); Hatschbach 39637 (MBM, US) ("Arbusto muito
ramoso, 2m, encosta de morro. Flor rosada"); Harley 19185
(CEPEC, K, US), Rio do Ferro Doido on BA-052 highway to Mundo
Novo, 19.5 km SE of Morro do Chapéeu, elev. 900 m ("Bushy shrub
to 2 m with stems bare below. Leaves rigid, spreading, mid-
green, slightly paler below. Sepals yellow green. Petals
bright magenta. Anthers dark golden yellow. Stigma yellow with
pinkish style"); Irwin, Harley, & Smith 32246 (US), Serra do
Tombador 18 km E of Morro do Chapeu, elev. 1100 m ("Brittle
ramose shrub ca 1.5 m tall. Corolla magenta; filaments red-
violet; anthers yellow"); E. Pereira 2131 (RB, US), elev. 1000 m
("Arbusto de fl. roxas"); Davidse, Ramamoorthy, & Vital 11896
(MO), 19 km E of Morro do Chapeu, elev. 900 m ("Shrub 70 cm
tall; petals reddish purple; leaves sticky").

Microlicia balsamifera is glabrous, with costulate

1981 Wurdack, Certamen Melastomataceis 149

(especially basally) hypanthia, longer (ca 1.5 m) anther rostra,
and large stamen connectives prolonged only 1-1.5 mm; recent
collections include Harley 15127, 15138, and 15423, all from the
region north of Rio de Contas, Bahia. Other glabrous relatives
include M. sincorensis (DC.) Mart., as well as perhaps M.
blanchetiana (Naud.) Cogn. and M. crebropunctata Pilger. Among
the pubescent species in Cogniaux’ monograph, M. hatschbachii
would perhaps key to near M. baccharoides Naud., which however
has denser eglandular stem pubescence, leaf blades obtuse to
rounded at the base, calyx lobes remote and subulate, and pro-
longation of the large stamen connectives 8-9 mm; recent
materials referred by me to M. baccharoides include Harley 15901
and three Mori collections (12581, 12589, 12631), all from south
of Mucuje, Bahia.

MICROLICIA HARLEYI Wurdack, sp. nov.

M. pallidae Cogn. et M. insigni Cham. affinis, foliis lati-
oribus calycis lobis glanduloso-ciliatis petalis amplioribus
differt.

Ramuli quadrangulati subtetrapteri glabri. Folia ascen-
dentia sessilia rigida essentialiter epunctata ciliolato-sub-
serrulata (ciliis 0.5-0.8 mm longis subappressis in foliis
floralibus glanduliferis in foliis inferioribus eglandulosis) ad
apicem 0.5-0.8 mm pungentia alioqui glabra apice acuto base late
acuta; folia inferiora 15-20 X 5-8 mm oblongo-elliptica 3-5-
nervata; folia superiora 8-10 X 3-4 mm lanceata 3-nervata.
Flores in ramulis foliosis terminales solitarii vel pauci; pedi-
celli ca 1 mm longi. Hypanthium (ad torum) 3 mm longum glabrum;
ecalycis tubus ca 0.3 mm longus extus in torii zona modice
glanduloso-setosus (setis ca 1.5 mm longis), lobis 6 X 1.8-2 m
oblongo-lanceatis ad basim contiguis ciliolato-serrulatis (ciliis
glanduliferis ca 1 mm longis) ad apicem 1-1.2 mm pungentibus
alioqui glabris. Petala 16-17 X 8-9 mm oblongo-obovata apice
late acuto et setula caduce glandulifera terminato alioqui
glabra. Stamina dimorphica glabra, filamentis 5 mm longis,
antheris oblongis 0.2-0.3 mm rostratis poro 0.2 mm diam. ventra-
liter inclinato. Stamina maiora: thecae 3.3 X 0.7-0.8 m:;
connectivum 7-7.5 mm prolongatum et ultra filamenti insertionem
1.5 mm porrectum incrassatum hebes. Stamina minora: thecae 2.3
mm longae; connectivum 2 mm prolongatum et ultra filamenti
insertionem 0.3 mm porrectum non incrassatum hebes. Stigma non
expansum; stylus 9 X O.4-0.1 mm glaber; ovarium ca 2.5 mm longum
glabrum 3-loculare.

Type Collection: R. M. Harley 19665 (holotype CEPEC 19690;
isotypes K, US), collected in a marsh in the first caldera above
the first escarpment, middle and upper northeast slopes of Pico
das Almas ca 25 km WNW of the Vila do Rio de Contas, Bahia,
Brazil, elev. 1600-1800 m, 19 March 1977. "Subshrub to 1 nm.
Petals magenta; anthers red.”

Both suggested relatives have leaves up to 11 X 3.5 m,
eciliate calyx lobes, and petals up to 11 m long.

150 PF Bak 2:0«L-01\6 © & Vol. 49 ,No. 2

PYRAMIA CAMBESSEDESIOIDES Wurdack, sp. nov.

A congeneribus foliis parvis fasciculatis hypanthiis glabris
differt.

Ramuli obscure quadrangulati sicut petioli modice pilis
pinoideis 0.1-0.2 mm longis setulosi. Folia plerumque ad nodos
fasciculati; petioli 0.1-0.2 mm longi; lamina 0.5-1 X 0.1-0.3 cm
oblonga apice hebeti-acuto vel obtuso vel rotundato basi late
acuta vel obtusa, rigida et integra, supra glabra, subtus sparse
pilis pinoideis 0.1 mm longis deciduis setulosa, 1-3-nervata ner-
vis secundariis nervulisque invisis. Panicula 2-5 cm longa
foliosa submultiflora; flores 5-6-meri, pedicellis 5-7 mm longis,
bracteolis 0.8-1 mm longis oblongo-ovatis persistentibus ca 1-
1.5 mm infra hypanthium insertis. Hypanthium (ad torum) 5-5.5 um
longum glabrum; calycis tubus 0.4-0.8 mm longus, lobis 0.6-0.8 mm
longis ovatis extus valde carinatis; torus intus sparse glandu-
losus. Petala glabra eciliata 7.5-10.5 X 4.4-5.3 mm elliptico-
oblonga apice acuto et apiculato. Stamina in dimensionibus paulo
dimorphica; filamenta 5.8-7.4 mm vel 4.8-6 mm longa apicem versus
glandulosa; antherae 5.7-6 X 0.8-1 mm vel 4.8-5.2 X 0.8-0.9 mm
paulo subulatae, thecis ad basim ventraliter 0.2-0.3 mm pro-
longatis. Stylus 16-16.5 X 0.4 mm basim versus sparse glandulo-
sus; ovarium 5(-6)-loculare apice modice glanduloso-setuloso
(0.1-0.3 mm).

Type Collection: S. A. Mori, R. M. King, T. S. dos Santos,
& J. L. Hage 12576 (holotype CEPEC 17835; isotype US), collected
3 km south of Mucujé, Municipio Mucujé, Bahia, Brazil, elev.

1000 m, 26 July 1979. "“Arbusto 1,5 m. Calice verde, corola
vermelha alaranjada, com uma mancha amarela nas bases das
pétalas, estames completamente amarelos."

Paratypes (all Bahia, Brazil): Edmundo Pereira 2181 (US,
RB), from between Palmeiras and Lengois ("Arbusto, petalas com
a metade superior cor de abobora e a base e os estamas amarelo");
A. P. Duarte 9358 (Pereira 10071) (RB, US), from Lengois
("arbusto 1 m, £1 amarela e vermelha"); Mori & Benton 13195
(CEPEC, US), from between Andarai and Mucuje 2 km south of
Igatu, elev. 800 m.

The three previously known species of Pyramia have non-
fasciculate leaves with blades 0.7-3 cm wide, as well as densely
puberulent hypanthia. The Bahia endemic suggests the evolution-
ary end of Pyramia towards Cambessedesia, the 5-6-celled ovaries
being the technical character of the former genus; the pubes-
cence qualitatively is like that in the Bahia variants of C.
membranacea Gardn. and C. harleyi Wurdack (vide infra).

CAMBESSEDESIA HARLEYI Wurdack, sp. nov.

C. membranaceae Gardn. affinis, petalis glanduloso-ciliola-
tis differt.

Ramuli primum rotundato-quadrangulati demum teretes sicut
inflorescentia hypanthiaque modice glanduloso-setuloso (pilis
plerumque 0.2-0.5 mm longis ad basim imperspicue papillosis) et
sicut laminarum subtus venae primariae petiolique modice vel
densiuscule setulosi (pilis asperis eglandulosis ca 0.2-0.3 m

1981 Wurdack, Certamen Melastomataceis 151

longis). Petioli 0.2-0.5 cm longi; lamina (1-)1.5-2.5(-3) x
1-2(-3.5) cm ovato-suborbicularis apice rotundato basi paulo
(usque ad 0.5 cm) cordata, subrigida et distincte irregulari-
terque crenato-serrulata, supra glabra.et dense bullata, subtus
foveolata et in venulis superficieque modice pilis asperis 0.2-
0.3 mm longis setulosa, 5(-7)-nervata nervis exterioribus usque
ad basim liberis vel paulo (ad 2 mm) coalitis. Panicula 3-7 cm
longa foliolosa submultiflora; flores 5-meri, pedicellis plerum-
que ca 2 mm longis, bracteolis ca 1-1.5 X 1-1.5 mm suborbicu-
laribus vel ovatis persistentibus. Hypanthium (ad torum) 6 m
longum paullulo costulatum; calycis tubus 0.5-0.6 m longus,
lobis 0.5-0.6 mm longis late ovatis paullulo hebeti-apiculatis;
torus intus inconspicue glanduloso-puberulus (0.1 m). Petala
5.7-6.5 X 367-45 mn elliptico-ovata apice hebeti-acuto apicem
versus crenulata et glanduloso-ciliolata (0.3-0.4 mm) alioqui
glabra. Stamina in dimensionibus paulo dimorphica; filamenta
10-10.5 mm vel 7.5-8 mm longa inconspicue glanduloso-puberula;
antherae 6.5 X 1 mm vel 4.6 X 0.7 m oblongo-subulatae, thecis
ad basim ventraliter 0.2-0.3 mm prolongatis, poro 0.15-0.2 mm
diam. ventraliter inclinato. Stigma punctiforme; stylus 20 X
O.4 mm basim versus glanduloso-puberulus; ovarium 3-loculare
inconspicue glanduloso-puberulun.

Type Collection: R. M. Harley 19189 (holotype CEPEC
19687; isotypes K, US), collected at Rio do Ferro Doido south-
east of Morro do Chapeu on BA 052 highway to Mundo Novo, Bahia,
Brazil, elev. ca 900 m, 1 March 1977. "Subshrub to 1.5 m.
Leaves strongly and finely bullate, mid-green above, pale
beneath. Calyx red-tinged. Petals bright vermilion with small
yellow basal zone. Stamens golden-yellow."

Paratypes (all Bahia, Brazil): Harley 16692 (CEPEC, K,
US), from Serra do Curral Feio 16 km northwest of Lagoinha (5.5
km southwest of Delfino) on road to Minas do Mimoso, alt. 950-
1000 m ("Shrub to 1m. Petals vermilion with yellow base.
Anthers yellow"); Irwin, Harley, & Smith 32431 (NY, US), topo-
typical ("Slender brittle shrub to ca 1.5 m. Corolla red,
yellow at base; stamens yellow"); Hatschbach 39609 (US), from
Morro do Chapéu ("Subarbusto 60 em, flor vermelha, estames
centraes amarelos"); Davidse, Ramamoorthy, & Vital 11864 (Mo),
from Chapada da Diamantina 34 km east of Morro do Chapeu, elev.
910 m ("Shrub to 1 m; petals red; stamens and style yellow,
turning orange in older flowers").

Cambessedesia membranacea has eciliate petals and (usually)
glabrous stamen filaments and style; the type collection (NY)
and Hatschbach 34127 (Serra Petrolina, M. Grosso) have leaf
blades beneath glabrous and hypanthia nearly or quite glabrous.
In Goias and Bahia, two variants of C. membranacea have been
collected, both with the branchlet internodes and leaf blades
beneath sparsely to moderately setulose with roughened hairs
like those in C. harleyi. One of these variants, centered on
Pico das Almas (Harley 19508 and 19609) and Serra do Rio de
Contas (Harley 15161, Mori 12343), has glabrous hypanthia; the
other variant, with glandular-setulose hypanthia, occurs in both

152 POHPY (TP Oeb OC: Len Vol. 49, No. 2

Goias (BR 354, Mineiros, Hatschbach 34255) and Bahia (Serra Agua
de Rega, Irwin 30899; Serra Sincora, Froes 20154, Harley 15585
and 15852). No material of C. eichleri Cogn. (Serra Caraga,
Minas, fide Glaziou) has been seen, but from the description the
much larger ovate acute leaves and eciliate petals are differen-
ces from C. harleyi. However, the puberulous variants above
referred to C. membranacea should be compared with C. eichleri.
The glandular-ciliolate corolla and glandular-puberulous fila-
ments and style of C. harleyi are like those in C. weddellii
Naud.; that Minas Gerais species, however, lacks roughened tri-
chomes, having only smooth (and glandular) hairs.

CAMBESSEDESIA GRACILIS Wurdack, sp. nov.

C. tenui Markgraf affinis, foliis angustis esetulosis
staminibus minoribus differt.

Ramuli primum obscure quadrangulati demum teretes glabri
(in nodis obscure caduceque puberuli pilis ca 0.1 mm longis).
Petioli 0.1-0.4 cm longi glabri; lamina (1-)2-3(-3.8) X (0.1)0.2-
0.3 cm lineari-oblonga apice basique acuto, firme chartacea et
integra vel apicem versus obscure distanterque serrulata, utrin-
que (glandulis paucis praecipue subtus exceptis) glabra, 1-3-
nervata nervis secundariis nervulisque invisis. Flores 5-meri
solitarii terminales, pedicellis ca 2 mm longis sparse glandu-
losis. Hypanthium (ad torum) ca 4.5 mm longum modice glanduloso-
setulosum (setulis 0.6-0.8 mm longis); calycis tubus ca 0.3 m
longus, lobis ca 1.2 mm longis ovatis extus carinatis. Petala
sparse glanduloso-ciliolata (ciliis 0.1-0.2 mm longis) ca 6 X
wm elliptica apice obtuso et ca 0.3 mm apiculato. Filamenta
2.5-3 mm longa sicut stylus sparse glandulosa; antherae 4 vel
2.5 X 0.5 mm oblongae, thecis ventraliter ad basim O.4-0.5 mm
vel 0.1-0.2 mm protractis, poro minuto ventraliter inclinato.
Ovarium 3-loculare apice modice glanduloso-setuloso (0.2-0.4 mm).

Type Collection: S. A. Mori, R. M. King, T. S. dos Santos,
& J. L. Hage 12533 (holotype CEPEC 16969; isotype US), collected
on Pico das Almas 18 km from Rio de Contas, Bahia, Brazil, elev.
1300-1400 m, 24 July 1979. "Subarbusto. Calice verde, o
restante da flor amarela."

Paratype: Mori & Benton 13574 (CEPEC, NY), near-topotypi-
cal, elev. 1200 m. "Subarbusto 50 cm de altura Calice verde con
pelos avermelhados, corola e estames amarelos."

The suggested relative has sparsely glandular-setulose
branchlets, lower leaf surfaces, and leaf margins, much wider
oval leaves (1-2.5 cm wide) and large anthers 6-6.5 mm long.
Cambessedesia hilariana (Kunth) DC. var. longifolia Cogn. has
leaves rather like those of C. gracilis and glandular-ciliolate
petals, but the leaves are fascicled, the branchlet nodes
distinctly villosulous, and the large anthers much longer; the
variety is perhaps specifically distinct from typical C.
hilariana and has both filaments and style sparsely glandular -
puberulous basally (a feature also true to some extent in other
varieties). .

1981 Wurdack, Certamen Melastomataceis 153

MARCETIA SINCORENSIS Wurdack, sp. nov.

M. nervulosae Markgraf affinis, ramulis (nodis exceptis)
esetulosis, foliis ad apicem acutis nervulis subtus ob indumen-
tum subamorphum occultis differt.

Ramuli quadrangulati modice resinoso-granulosi esetosi
(nodis exceptis); nodi obscure caduceque glanduloso-setulosi
pilis 0.1-0.2 mm longis. Folia essentialiter sessilia, petiolis
vix 0.5 mm longis; lamina (8-)10-18 X (5-)7-9(-13) mm ovata vel
oblongo-ovata apice acuto basi rotundato-truncata, rigida, supra
primum sparse resinoso-glandulosa mox glabrata, subtus dense
resinoso-granulosa, (7-)9-nervata nervulis subtus paulo denseque
elevato-reticulatis ob indumentum obscuris. Flores in foliorum
superiorum axillis oppositis solitarii (in cyma pauciflora foli-
osa 2-3 cm longa aggregati); pedicelli 1-2 mm longi, bracteolis
ca 3 X 0.7 mm persistentibus. Hypanthium (ad SSS) 6.5 mm
longum sparsiuscule resinoso-granulosum esetulosum; calycis tubus
O.4 mm longus, lobis 3-3.5 X ca O.7 mm subulatis remotis. Petala
7-5-8 X 5 mm elliptica (apice acuto) minute (0.05 mm) glanduloso-
ciliolata alioqui glabra. Stamina isomorphica glabra; filamenta
6.5 mm longa; thecae 6.5-6.6 X 1 X 1 mm oblongo-subulatae, poro
0.15 mm diam. ventraliter inclinato, connectivo ad basim paulo
incrassato. Stigma punctiforme; stylus 12.5 X 0.5-O.1 mm glaber;
ovarium 4-loculare glabrum.

Type Collection: R. M. Harley 18855 (holotype CEPEC 19249;
isotypes K, US), collected mee -15 km north of Mucujé on road to
Andaratf, Serra do Sincora, Bahia, Brazil, elev. ca 1100 n,

18 Feb. 1977. "Shrub to 1.5 m. Leaves grey, paler beneath.
Petals white. Stamens with showy yellow anthers and white
filaments."

Paratypes: Mori, King, dos Santos, & Hage 12657 (CEPEC, US;
"Arbusto 1,5 m de altura. Corola branca, filetes e estilete
brancos.' ry and 12666 (CEPEC, US; fruiting), both from 10-12 km
northwest of Mucuje on road to Andaratf, Bahia, Brazil, elev.

1000 m.

Marcetia nervulosa has branchlet internodes densely setulose
with rather robust caducously gland-tipped hairs, leaves with
obtuse to rounded apices and with the nervules beneath obvious
and moderately setulose with gland-tipped hairs O.1-0.3 mm long,
and hypanthia definitely (but rather sparsely) setulose with
gland-tipped hairs O.1-0.3 mm long. While the petals of M.
nervulosa were thought by Markgraf to be yellow, this may well
have been a drying artifact. ‘The only recent collection seen of
M. nervulosa is Mori 12915 (Pai Inacio ca. 15 km NE of Palmeiras,
Mun. Palmeiras, Bahia, elev. 1000-1200 m), fruiting.

MARCETIA MACROPHYLLA Wurdack, sp. nov.
M. grandiflorae Markgraf affinis, ramulorum pubescentia
longiore petalis angustioribus antheris paulo subulatis differt.
Ramuli quadrangulati dense setosi pilis gracilibus ca 3-4
mm longis eglandulosis pilis glanduliferis ca 1 mm longis modice
intermixtis. Folia essentialiter sessilia, petiolis ca 1-1.5 mm
longis; lamina plerumque 2.5-4 X 1.5-2(-2.5) em oblongo-elliptica

154 Rhye OH O'S TA Vol. 49, No. 2

apice obtuse rotundato basi paullulo (1-2 mm ) cordata, rigida,
supra dense subsericeo-strigosa pilis gracilibus eglandulosis

ca 2 mm longis, subtus dense appresso-setosa pilis gracilibus

ca 2 mm longis pilis glanduliferis ca 0.1 mm longis inconspicue
intermixtis, (7-)9(-11)-nervata nervulis subtus crebro elevato-
reticulatis ob pilos occultis. Panicula foliosa 2-4 cm longa
pauciflora; pedicelli plerumque 4-6 mm longi sicut hypanthia
(extus) et sepala (ubique) densiuscule setosi pilis gracilibus
1-2 mm longis pro parte glanduliferis. Hypanthium (ad torum) 6-
7-5 mm longum; calycis tubus ca 0.2 mm longus, lobis 5.2-7 X l-
1.2 mm lanceatis. Petala 6.5-8 X 3.7-5 mm anguste ovata apice
acuto et setula 0.5-0.7 mm longa terminato glanduloso-ciliolata
et extus circum margines glanduloso-puberula. Stamina isomorph-
ica glabra; filamenta 7.2-8 mm longa; antherae 7-7.5 X 0.9-1 mm
paulo subulatae poro 0.15-0.2 mm diam. ventraliter inclinato;
connectivum ad basim vix incrassatum. Stigma punctiforme; stylus
19-20 X 0.4-0.5 mm glaber; ovarium 4-loculare glabrum.

Type Collection: R. M. Harley 15986 (holotype CEPEC; iso-
types K, US), collected near Rio Cumbuca ca 3 km north of Mucujé
on Andaraf road, Serra do Sincora, Bahia, Brazil, elev. ca 850 n,
5 Feb. 1974. "Shrub ca 1.5 m. Leaves dark green with impressed
veins above, paler beneath. Calyx red. Petals magenta; stamens
with white filaments and deep yellow anthers."

Paratypes (both Serra do Sincora, Bahia): R. L. Froes
20236 (NY, US), from Mucujé; Harley 18667 (CEPEC, K, US), from
south of Andaraf 16 km along Mucuje road near Xique-Xique, elev.
700-900 m. "Shrub to 3 m, with bright red-brown bark. Leaves
spreading, rigid, dull green above, paler beneath. Calyx red;
petals pale magenta. Stamens with golden-yellow anthers and pink
filaments; style deep pink.”

The suggested relative has stem pubescence 1-1.5 mm long,
leaf blades 14-16 X 10-11 mm with 13 primary nerves, calyx lobes
10 mm long, petals 18 X 18 mm, and oblong anthers with a broad
(0.3 mm) but slightly ventral pore. Another relative, M.
velutina Markgraf, has short cauline pubescence like that of M.

randiflora, subrotund leaf blades 10-20 X 12-18 mm, calyx lobes
only ca 3 mm long, and anther pore dorsally inclined (and anthers
shaped as in M. macrophylla.) ; two recent collections of this
Bahia endemic are Harley 15164 (Serra do Rio de Contas) and Mori
et al 12560 (Mucuje).

MARCETIA LANUGINOSA Wurdack, sp. nov.

M. nummulariae Markgraf affinis, foliorum subtus ramulorum-
que pubescentia arachnoideo-lanuginosa differt.

Ramuli quadrangulati sicut foliorum laminae subtus densis-
Sime arachnoideo-lanuginosi pilis eglandulosis. Folia sessilia;
lamina (0.7-)1-1.3 X (0.5-)0.9-1.3 cm ovato-suborbicularis apice
late obtuso vel rotundato basi paulo (1 mm) cordato-amplexicauli,
rigida, supra modice strigulosa pilis eglandulosis robustis
plerumque 0.5-0.7 mm longis, 5-7-nervata. Flores ad ramorum
apices solitarii vel in inflorescentia foliosa usque ad 3 cm
longa aggregati; pedicelli 1-2 mm longi sicut hypanthia dense

1981 Wurdack, Certamen Melastomataceis 155

pilis gracillimis laxis et paulo crispulis minute glanduliferis
1.5-2 mm longis induti. Hypanthium (ad torum) 3.5-4 mm longum;
calycis tubus 0.2 mm longus, lobis 3.5 X 0.8 mm anguste oblongis
remotis ubique dense glanduloso-setulosis. Petala 8.5-11 X 7-
7-7 mm oblongo-elliptica apice setula glandulifera 0.6-0.7 mm
longa terminato glanduloso-ciliolata alioqui glabra. Stamina
isomorphica glabra; filamenta 5.1-5.4 mm longa; antherae 5-5-5 X
0.8-0.9 mm, poro 0.15 mm diam. paulo ventraliter inclinato,
connectivo ad basim paullulo incrassato. Stigma punctiforme;
stylus 21.5 X 0.4 mm glaber; ovarium 4-loculare glabrum.

Type Collection: R. M. Harley 15828 (holotype CEPEC; iso-
types K, US), collected ca 10 km north of Barra da Estiva by Rio
Preto on Ibicoara road, Serra do Sincora, Bahia, Brazil, ieee
1100 m, 2 Feb. 1974. ‘Eahohenn 20 em high. Flowers magenta.’

Paratypes (all Bahia): Harley 15095 (CEPEC, K, US), from
ca 6 km north of town of Rio de Contas on Abaira road, Serra do
Rio de Contas, elev. mae m ("Brittle-stemmed decumbent shrub to
ca 20 cm"); Harley 15714 (CEPEC , K, US), from north face of Serra
de Ouro, 7 km south of Barra da Estiva on Ituacu road, Serra do
Sincora, elev. ca 1150 m ("Subshrub with magenta flowers");
Harley 20007 (CEPEC, K, US), from ca 5 km east of Vila do Rio de
Contas on Marcolina Moura road, Serra do Rio de Contas, elev.
1000 m ("Erect shrublet with single stems to 10 cm high. Stem
and underside of leaves white-woolly. Leaves above dark green,
with white appressed hairs. Calyx lobes maroon; petals bril-
liant mauve; filaments white; anthers lemon yellow; style
magenta"); King & Bishop 8613 (UB, US), from 16 km north of
Livramento do Brumado along road to Arapiranga, elev. 900 m
("Corolla magenta").

Marcetia nummularia has leaves of similar shape but stems
and primary leaf veins beneath with discrete erect setulae.
Evidently Markgraf’s floral dimensions were taken from a bud
(packet on Luetzelburg 175, M) and there are now only fruit
visible on the specimen branches; the petals are probably larger
than 2 mm.

MARCETIA VISCIDA Wurdack, sp. nov.

M. Sincorensi Wurdack affinis, foliis distincte petiolatis
laminis ad basim acutis calycis lobis pedicellisque longioribus
differt.

Ramuli obscure quadrangulati sicut pedicelli calycis lobi
hypanthiaque glabri. Petioli 1.5-2.5 mm longi; lamina 12-24 x
4-6 mm lanceata vel oblongo-lanceata apice hebeti-acuto basi
late acuta, subrigida, supra glabra, subtus densiuscule arach-
noideo-furfuracea, (5-)7-nervata nervulis subtus ob indumentum
occultis. Flores in foliorum superiorem axillis oppositis sin-
guli, pedicellis ca 5 mm longis, bracteolis 3-5 X 0.6-1 m
persistentibus. Hypanthium (ad torum) 4.5 mm longum; calycis
tubus 0.3 mm longus, lobis 6.3-6.6 X 1.1-1.2 mm oblongis remotis
apice rotundato; torus intus sparse glanduloso-setulosus. Petala
12 X 8-8.2 mm elliptica (apice late acuto vel obtuso) minute
glanduloso-ciliolata. Stamina isomorphica glabra; filamenta 4 m

156 2 ey ro'h od TA Vol. 49, No. 2

longa; antherae 4.5-4.8 X 0.7-0.8 X 0.7 mm oblongae poro 0.2 mm
diam. ventraliter inclinato, connectivo ad basim ventraliter
bilobulato-incrassato. Stigma punctiforme; stylus 6.6 X 0.4 mm
sparse glandulosus; ovarium semper 4-loculare glabrum.

Type Collection: R. M. Harley 15162 (holotype CEPEC; iso-
types K, US), collected 12-14 km north of town of Rio de Contas
on road to Mato Grosso, Serra do Rio de Contas, Bahia, Brazil,
elev. ca 1200 m, 17 Jan. 1974. "Viscid wiry subshrub to ca 35 cm.
Petals white or cream.”

Paratype: Harley 20149 (K), collected 2.5-5 km south of
Vila do Rio de Contas, Bahia, Brazil, elev. ca 980m. "Spindly
shrub to ca 75 cm with bare stems and leaves mainly fallen.
Leaves rigid, dull dark green, paler beneath. Calyx red-tinged,
very viscid. Petals white; anthers yellow; filaments white.”

Marcetia sincorensis has essentially sessile and thicker
leaves with rounded-truncate base, pedicels 1-2 mm long, sub-
ulate calyx lobes 3-3.5 mm long, and petals 7.5-8 X 5 m, as
well as more subulate anthers with less distinct ventro-basal
connective thickening.

MARCETIA FORMOSA Wurdack, sp. nov.

M. viscidae Wurdack affinis, foliis 9-nervatis floribus
maioribus petalis magentis differt.

Ramuli rotundato-quadrangulati sicut foliorum venae pri-
mariae subtus hypanthiaque densiuscule caduceque resinoso-
granulosi esetosi. Petioli 2-2.5 mm longi crassi; lamina 20-30 X
6-12 mm oblongo-elliptica apice hebeti-obtuso basi late acuta,
rigida, supra primum modice resinoso-granulosa glabrata, subtus
in superficie dense arachnoideo-furfuracea, 9-nervata. Flores in
foliorum superiorem axillis oppositis solitarii, pedicellis 10-le
mm longis, bracteolis non visis. Hypanthium (ad torum) 6.5 mm
longum paulo obtuseque 8-costatum; calycis tubus 0.5 mm longus,
lobis 10.5-11 X 2 mm (apice rotundato); torus intus dense glan-
duloso-ciliatus pilis gracillimis 2.5-2.7 mm longis. Petala
24-25 X 20-21 mm minute (0.05 mm) glanduloso-ciliolata alioqui
glabra. Stamina isomorphica glabra; filamenta 6 mm longa;
antherae 5-5.5 X 1.6-2 X 1.5 mm oblongae, poro O.3 mm diam.
ventraliter inclinato, connectivo ad basim dorsaliter incrassato.
Stigma punctiforme; stylus 10 X 0.35 mm glaber; ovarium }+-locu-
lare glabrun.

Type Collecticn: R. M. Harley 15452 (holotype CEPEC; iso-
types K, US), collected on slopes of Pico das Almas ca 25 km WNW
of Rio de Contas, Bahia, Brazil, elev. ca 1600 m, 23 Jan. 1974.
"Wiry shrub to ca 1 m with leaves very dark green above, greyish
beneath. Calyx viscid. Petals deep very bright magenta."

Both species share the feature of the torus being glandular-
puberulous within. Certainly M. formosa has the largest flowers
now known in the genus.

MARCETIA HARLEYI Wurdack, sp. nov.
M. sincorensi Wurdack affinis, foliis floribusque minoribus
ovario 3-loculari differt.

1981 Wurdack, Certamen Melastomataceis 157

Ramuli obscure tetragoni sicut folia subtus calycis lobi
(intus et extus) hypanthiaque dense furfuracei indumento ut
videtur subamorpho sed magnificatione maxima glanduloso. Folia
essentialiter sessilia (petiolis crassis obscuris ca 0.3 m
longis) 5-8(-11) X 2-5 mm oblongo-elliptica apice hebeti-acuto
vel obtuso basi late acuta, rigida, supra glabra, 5-7(-9)-
nervata nervulis on indumentum occultis. Flores in foliorum
superiorem axillis oppositis solitarii, pedicellis ca 2 m
longis, bracteolis 2-2.5 X 0.8-1 mm persistentibus. Hypanthium
(ad torum) 3 mm longum obscure 8-costatum; calycis tubus 1.2-
1.5 mm longus, lobis 2.4-2.5 X 1.5 mm ovato-oblongis obtusis
remotis, appendicibus intercalycinis non evolutis. Petala 8-
9.5 X 7-5-9.5 mm late obovata (apice rotundato) minutissime
glanduloso-ciliolata alioqui glabra. Stamina isomorphica glabra;
filamenta 4 mm longa; thecae 2.7-2.9 X 0.9-1 X 0.8-1 mm lance-
atae, poro 0.15 m diam. ventraliter inclinato, connectivo ad
basim paulo incrassato. Stigma punctiforme; stylus ca 5 X 0.35-
O.4 mm glaber; ovarium semper 3-loculare glabrun.

Type Collection: R. M. Harley 15698 (holotype CEPEC; iso-
types K, US), collected on north slope of Serra de Ouro 7 km
south of Barra da Estiva on Ituacu road, Serra do Sincora, Bahia,
Brazil, elev. ca 1150 m, 30 Jan. 1974. "Subshrub 20 cm high.
Flowers magenta.”

Paratypes (all Bahia, Brazil): Harley 15518 (CEPEC, K,
US), from 6 km north of Barra da Estiva on Ibicoara road, Serra
do Sincora, elev. 1100 m ("Wiry shrub to ca 50cm. Leaves
glossy, dull green above, grey beneath. Calyx red; petals
magenta; filaments white; anthers bright yellow"); Harley 15841
(CEPEC, K, US), from 14 km north of Barra da Estiva near Ibicoara
road, Serra do Sincora, elev. 1100 m ("Wiry subshrub to ca 20 cm.
Leaves dark green above, grey beneath. Petals magenta; anthers
yellow"); Harley 15094 (CEPEC, K, US), from 6 km north of Rio de
Contas on Abaira road, Serra do Rio de Contas ("Wiry subshrub to
ca 20 cm. Leaves dull green. Petals bright magenta; stamens
orange-yellow"); Irwin, Harley, & Smith 31075 (NY, US), from 24
km north of Seabra on road to Agua de Rega, Serra da Agua de
Rega, elev. ca 1000 m ("Subshrub to ca 40 cm tall. Corolla
rose-pink, aging to deeper pink"); Irwin, Harley, & Smith 32413
(NY, US), from Morro do Chapéeu, Serra do Tombador, elev. 1125 m
("Subshrub ca 35 cm tall, from thick rootstalk. Corolla red-
violet; filaments white; anthers yellow"); Hatschbach 39599
(US), from Morro do Chapéu ("Ereta 35 cm, flor purpurea, anteras
e estames amarelos"); King & Bishop 8612 (UB, US), from 16 km
north of Livramento do Brumado on road to Arapiranga, elev.

900 m ("Corolla magenta").

In qualitative foliar features, M. harleyi and M. sincoren-
sis are quite compatible; however M. sincorensis has much larger
flowers with acute petals. Technically the 3-celled ovary would
remove M. harleyi from Sect. Marcetia, but the other species
features (and lack of intersepalar appendages ) do not suggest any
close affinity with M. gracillima Cogn. and M. luetzelburgii
Markgraf of Sect. Pseudomarcetia.

158 rH YT O40 OTA Vol. 49, No. 2

PTEROLEPIS HATSCHBACHII Wurdack, sp. nov.

P. weddellianae (Naud.) Triana affinis, floribus minoribus
trimeris differt.

Suffrutex ut videtur ca O.4 m superne ramosus; ramuli
subalato-quadrangulati (alis ca O.1 mm altis modice strigulosis
pilis ad basim non productis) inter alas glabri. Petioli ca 0.5
mm longi crassi; lamina 1-2.5 X O.4-0.8 cm lanceata apice acuto
basi obtusa, chartacea et essentialiter integra appresso-cilio-
lata, supra sparsiuscule strigosa (pilis gracilibus ca 0.8-1.3
mn longis ad basim ca 0.2 mm adnatis), subtus in superficie
sparse strigulosa pilis 0.3-0.5 mm longis, trinervata nervis
primariis lateralibus supra invisis. Flores fere semper 3-meri
in axillis foliosis superioribus plerumque solitarii, pedicellis
1-1.5 mm longis. Hypanthium (ad torum) 4 mm longum sparse stri-
gosum pilis omnibus simplicibus gracilibus eglandulosis 1-2 mm
longis; calycis tubus 0.2 mm longus appendicum axibus 0.7-0.8 mm
longis, lobis 5-5.2 X 2.2-2.4 mm oblongo-lanceatis ciliolatis
seta terminali ca 2 mm longa alioqui glabris. Petala 10 X 10-11
mm late obovata apiculata seta terminali ca 2 mm longa excepta
glabra. Stamina in dimensionibus paullulo dimorphica glabra;
filamenta 5.5 mm vel 4.8 mm longa; thecae 3.9-4 X 0.6 mm vel
3.4 X 0.5 mm subulatae, poro 0.15 mm diam. ventraliter incli-
nato; connectivum non prolongatum, lobis ventralibus incrass-
atis ca 0.4-0.5 X 0.5-0.6 mm. Stigma non expansum; stylus 11 X
O.4-0.15 mm glaber; ovarium 3-loculare pilis apicalibus 0.2-

O.7 mm longis pro parte glanduliferis.

Type Collection: G. Hatschbach & 0. Guimaraes 42363 (holo-
type MBM 62898; isotype us), collected in moist sandy soil at
Serra do Tombador, Mun. Morro do Chapéu, Bahia, Brazil, elev.
1050 m, 15 July 1979. "Flor lilas."”

Pterolepis weddelliana has 4-merous flowers with the hy-
panthium plus calyx tube 6-7 mm long and equaled by the calyx
lobes, as well as anthers ca 6 mm long and connective prolonged
ca 0.5 mm; to P. weddelliana I have referred Woolston 906 (US,
from Primavira, Alto Paraguay, Paraguay) and Steinbach 5151
(US, from Buena Vista, Prov. Sara, Depto. Santa Cruz, Bolivia,
elev. 500 m). The other species placed by Cogniaux in this
complex, P. repanda (DC.) Triana, has cauline pubescence on all
sides (rather than confined to the angles), as well as larger
Plowers with longer stalks on the intersepalar hairs. I do not
believe that the other trimerous species, P. trimera Ule, is
closely related to P. hatschbachii (nor is the Sect. Trimero-
calyx Ule natural). To P. trimera has been referred Hatschbach
& Guimaraes 42353, from Serra do Tombador, Bahia, elev. 1000 n,
agreeing well with Ule's description (but with lilac petals); of
the examinable flowers in this recent collection, 26 had 3 calyx
lobes and 20 had 4 calyx lobes. All except 2 of the 42 examin-
able flowers or fruits in Hatschbach 42363 showed 3 sepals.

ADDITIONAL NOTES ON THE GENUS PRIVA. X

Harold N. Moldenke

PRIVA Adans.

Additional & emended bibliography: Brongn. , Enum. Gen. P1l.,
ed. 1, 65. 18435 Walp., Repert. Bot. Syst. 6: 687. 1847; Brongn.
Enum. Gen. Pl., ed. ‘2 5:119. 1850; Briq. in Engl. & Pranti, Nat.
Pflanzenfam., ed. 1, 4 (3a): 133, 137, 139, 142--144, 153 ,& 155,
fig. 59 E. 1895; Mold. , Phytologia 49: 58--64. 1981.

>

PRIVA ASPERA H.B.K.
Additional bibliography: Walp., Repert. Bot. Syst. 6: 687.
1847; Mold., Phytologia 49: 61. 1981.

PRIVA GRANDIFLORA (Ort.) Mold.
Additional bibliography: Mold., Phytologia 49: 63. 1981.
Miranda encountered this plant in pine woods and pedregal.
Additional citations: MEXICO: Distrito Federal: Miranda 712
(Me--74094). México: Miranda 449 (Me--94095).

PRIVA LAPPULACEA (L.) Pers.

Additional bibliography: Briq. in Engl. & Prantl, Nat. Pflan-
zenfam., ed. 1, 4 (3a): 143. 1895; Mold., Phytologia 49: 61 & 63.
1981.

The Cuatrecasas & Castaneda 25521 and Proctor 3341, distribu-
ted as typical P. lappulacea, actually represent f. albiflora
Mold., while Daniel 5639 (at least insofar as the United States
National Herbarium specimen is concerned) is not verbenaceous.

Additional citations: NICARAGUA: Estelf: w. D. Stevens 2603
(Ld), 9082 (Ld). Leén: W. D. Stevens 4693 (Ld). Managua: W. D.
Stevens 2650 (Ld), 2895 (ld), 3433 (Ld). Masaya: Araquistain 234
(Ld); Vincelli 772 (Ld). Matagalpa: W. D. Stevens 9380 (Ld).
Rivas: W. D. Stevens 3777 (Ld). PANAMA: Panama: Croat 34678 (W--
2846392). COLOMBIA: Choco: Forero, Jaramillo, Ledén, & Forero P.
1902 (N), VENEZUELA: Lara: Steyermark & Espinoza 108774 (N).
GALAPAGOS ISLANDS: Santa Cruz: Fournier 207 (W--2853561). PERU:
Madre de Dfos: Foster, Foster, Brokaw, & Brokaw 3307 (W--2888882).
BRAZIL: AmazOnas: Lasseigne P.21182 (N).

PRIVA LAPPULACEA f£. ALBIFLORA Mold.

Additional bibliography: Lopez-—Palacios, Revist. Fac. Farm.
Univ. Andes 20: 30. 1979; Mold., Phytologia 44: 98, 102, & 104--
105. 1979; Mold., Phytol. Mem. 2: 50, 66, 74, 78, 79, 84, 101,
2245 118, 123,0429, 2315 & 573201980;

Recent collectors refer to this plant as an herb with weak
stems to 1 m. long, yellow anthers, and the fruit green with a
fruiting-calyx easily attaching itself to the passerby, and have
encountered it at the upper edge of a potrero as well as on wood-
ed islets on slightly elevated ground in savannas "visited in
part by cattle", in shade along roadsides, in coffee plantations,

159

160 FHI TOL@eezT A Vol. 49, No. 2

on steep uncultivated slopes of soft volcanic rock, in weedy
Panicum maximum pastures on neutral to slightly calcareous soil
"of coluvial origin", and "occasional along stony paths", at 80--
1050 m. altitude, flowering and fruiting in April and from Sep-
tember to November. The corollas are said to have been "white
with purple lines" on Hinton 17664 and "white, pink-veiny" on
Chrostowski 69-132. The vernacular name, "yerba amarosa"”, is re-
ported.

Material of this form has been misidentified and distributed
in some herbaria as the typical form and also as P. mexicana (L.)
Pers. Calderdén reports the vernacular name, "cadillo de bolita”.

Additional citations: MEXICO: Nuevo Ledn: Hinton 17664 (Au).
Oaxaca: Calderdn 24 (Au, N). NICARAGUA: Estel{: Stevens & Ara-
quistain 14942 (Ld). Managua: W. D. Stevens 3922 (Ld). NORTHERN
SOUTH AMERICAN ISLANDS: San Andres: Proctor 3341 (W--1979220).
COLOMBIA: Guajira: Cuatrecasas & Romero Castaneda 25521 (W-2342058).
Magdalena: Kirkbride 2528 (W--2835048). GUYANA: Maas & Westra
4109 (Ld). PERU: San Martin: Chrostowski 69-132 (Ws).

PRIVA MEXICANA (L.) Pers.

Additional & emended bibliography: J. F. Gmel. in L., Syst.
Nat.5 ed. 13, Ampw. 42 2241. 9791:. Poise. in| Lams, Tabi .¥ucyel.
Méth. Eots: [IlLlustr, Gen.] 1: 59 & 60. 1791; Loud.,. Hort. Brit.,
ed. 1, 246. 1830: Schiecht. & Cham., Linnaea.5:. 98-—99= 1830;
foud., Hort. Brito, ed.(23 246.1832; -G. Don din-Loud.,.Hort:Brit.,
ede. 132 247) 28395 (Gs Donwin-Sweets Hort... BEit:, edi 3°4.5529c1839;
Steud... Nom.) Bot. Phan., eds 2 .,:25..397 & 750.1841; D.. Dictrs;

Syn. Pl. 3: 606. 1843; H. N. & A. L. Mold., P1. Life 2: 33. 1948;
Mold., Phytologia 44: 102 & 105--108. 1979, Mold., Phytol. Mem.
224394, 665735, 165.94, ° 1025" 359594515462, & 57321980.

Arguelles reports finding this plant growing along with Alnus,
Fraxinus, Litsea, Taxodium , Quercus, Salvia, and various composites.
Ventura reports the stamens white, the corollas lilac, and the fruit
green, and the species scarce at 2400 m. altitude.

Material of this species has been misidentified and distributed
in some herbaria as Verbena sp. On the other hand, the Hinton
17664, distributed as P. mexicana, actually is P. lappulacea f.
albiflora Mold.

Additional citations: MEXICO: Distrito Federal: Miranda 311
(Me--74098); Ventura A. 3031 (N). Hidalgo: Ventura A. 1651 (Me--
275718). Mexico: Hinton 18010 (Au). Querétaro: Arguelles 1299
(Me--275938).

PRIVA MEYERI Jaub. & Spach

Additional bibliography: C. Muell. in Walp., Ann. Bot. Syst. 5:
705. 1860; Vierh., K. Akad. Wiss. Wien Denkschr. 71: 114 [434].
19072 H.-N; 06 ,A. L.-Mold. ,.Pisibife 27,72.°1948s Mold., Phytotogia
44: 92 & 108--109. 1979; Mold., Phytologia Mem. 2: 201, 224, 228,
238, 241, 242, 244, 246, & 573. 1980; Mold., Phytologia 49: 62.
1981.

Recent collectors have found this plant at 1600 m. altitude, in

1981 Moldenke, Notes on Priva 161

flower and fruit in February.

The Dahlstrand 1638, distributed as P. meyeri, actually is P.
cordifolia var. australis Mold.

Additional citations: SOUTH AFRICA: Cape Province: Drége a
[Mo. Bot. Gard. photo 866 in part] (Z--photo of cotype). Natal:
Collector undetermined 2202 [Mo. Bot. Gard. photo A.866 in part]
(Z--photo). Transvaal: Dahlstrand 1505 (Go).

PRIVA MEYERI var. MADAGASCARIENSIS Mold.
Additional bibliography: Mold., Phytologia 44: 109. 1979;
Mold., Phytol. Mem. 2: 251 & 573. 1980.

PRIVA PEDICELLATA Mold.
Additional bibliography: Mold., Phytologia 44: 109--110. 1979;
Mold., Phytol. Mem. 2: 273 & 573. 1980.

PRIVA PERUVIANA Mold.

Additional bibliography: Mold., Phytologia 44: 110. 1979;
Mold. , Phytol. Mem. 2:° 135 & 573°. 1980.

Recent collectors have encountered this plant on alluvial soil
in clearings in mature forests, flowering and fruiting in October,
describing it as an herb with green fruit. The corollas are said
to have been “white” on the collection cited below.

Additional citations: PERU: Madre de Dios: Gentry, Aronson, &
Ramirez 26928 (Z).

PRIVA PORTORICENSIS Urb.
Additional bibliography: Mold., Phytologia 44: 110. 1979;
Mold., Phytol. Mem. 2: 98 & 573. 1980.

PRIVA SOCOTRANA Mold.

Additional & emended bibliography: Balf. f., Trans. Roy. Soc.
Edinb. 31: [Bot. Socotra] 232--233. 1888; Vierh., K. Akad. Wiss.
Wien Denkschr. 71: 114 [434]. 1907; Mold., Phytologia 44: 110.
1979; Mold., Phytol. Mem. 2: 253 & 573. 1980.

ADDITIONAL NOTES ON THE GENUS VITEX. XX

Harold N. Moldenke

VITEX Tourn.

Additional & emended bibliography: P. Herm., Mus. Zeyl., ed.
47. 717; Le, Fl. Zeyl., imp. 2, 194—195. 1748; PS Browne aa
Sloane, Civil Nat. Hist. Jamaic., ed. 1, 267. 1756; Kwa-wi
[transl. Savatier], Arbor. 4: pl. 1. 1789; P. Browne in Sloane,
Civil Nat. Hist. Jamaic., ed. 2, 267. 1789; Wall. in Roxb., Fl.
Ind., ed. 1, imp. 1, 1: 481--482. 1820; Roxb., Fl. Ind., ed. 1,
imp. 1, 3: 70 & ‘71. 1824: Chan. , Linnsea 7: 107—-109 5371-375,

162 PRY 28 6.6 sak Vol. 49, No. 2

& 400. 1832; Wight, Icon. Pl. Ind... Oriente 2, (332 1,01 451%.
1842; Brongn., Enum. Gen. Pl., ed. 1, 65. 1843; D. Dietr., Syn.
Pl. 3: 371 & 610--612. 1843; Walp., Repert. Bot. Syst. 6: 690.
1847; Walp., Ann. Bot. Syst. 1: 542 & 545. 1849; Brongn., Enum.
Gen. Ply, Cie) 25.119: & .120...3850:> Hook, £5, Fie SM. Zeal, ie):
203. 1852; Miq., Fl. Ind. Bat. 1 (1): 858--865. 1856; C. Muell.

in Walp., Ann. Bot. Syst. 5: 712. 1860; A. Wood, Class-book, [ed.
42], imp. 3, 539. 1864; Pfeiffer, Nom. Bot. 2 (2): 858 & 896.
1874; Mercado in Blanco, Fl. Filip., ed. 3, 4 Lib. Med. 36. 1880;
Hemsl. in Thomson & Murray, Rep. Scient. Res. Voy. Challenger 3,
Bot. 1: 110 & 177—178.. 18855; Vidal: y, Soler, Phans Cusine > Paii—
ip. 15, 39, 44, 64, 72,. 134, &.135;. 1885: F. Meet. See. Svat.
Census Austr. Pl. 1: 173. 1887; Kuntze, Rev. Gen. Pl. 2: 510--

S11 & 513. 1891; Bocq., Fl. Febrig. Colon, Frane. 68. 1895;

Briq. in Engl. & Prantl, Nat. Pflanzenfam., ed. 1, 4 (3a): 132--
144 & 169--172. 1895; Woodrow, Journ. Bomb. Nat. Hist. Soc. 5:

12 & 359. 1899; Diels, Fl. Cent.-China 549. 1902; Post & Kuntze,
Lexicon 103, 134, 589, & 688. 1904; Laing & Blackwell, Pl. N.
Zeal., ed« 1, 210, 349--351, 98 456,: fie. 114.: 1906s Koordy & Valu,
Atlas Baumart. Java 2: 6 & 201, pl. 292--299. 1914; Masuda, Bot.
Mag. Tokyo 28: [418]. 1914; Chiov., Result. Scient. Miss. Stef.

3 144. & 218. 1916; Basu, : Indian; Med. Pl. ,, ed:.3, 32.3) & 1936——
1940, pl. 740--742. 1918; Parker, Forest Fl. Punj., ed. 1, 391 &
394. 1918; H. J. Lam in Lam & Bakh., Bull. Jard. Bot. Buitenz.,
sec. 3, 3:,4/-—50. 1921; Bodding,.Mem. Asiat. Soc., Bene. 40:. 3;
[9 12. BO, Obs, Ids Gls Bogeot, 965 .100,) 103, S299. 1925;
Cheeseman, Man. N. Zeal. Fl., ed. 2, 763--765 & 1163. 1925; J.
Havchine., Fam.: Flow... Pi.,.eda 1, 309 &,.327.,,£46.¢ 2631, 4.9265
Heyne, Nuke. Pi. Ned. Ind. , eds) 2,22 1313)& 4317--1515., 1927;
Laing & Blackwell, Pl. N. Zeal., ed. 3, 354--356 & 468, fig. 127.
1929; Osmaston, Forest Fl. Kumaon 405--406. 1927; Fedde & Schust.,
Justs Bot. Jahresber. 47 (2): 423 & 426. 1929; W. Trelease, Wint.
Bot., Jed. 3,, imp. 1, 323, 325, & 335. 1931; Madrid Moreno, Deciar.
Virt. Arb. Pl. 110 & 173. 1936; Makins, Ident. Trees Shrubs 259.
£930; Breyn, Frod. Fase. Rars PL. ed., 2, 2+. 106..1939; Lam &
Meeuse, Blumea 3: [248]--254. 1939; Laing & Blackwell, Pl. N.
Zeal., ed. 4, 321, 371--373, & 499, fig. 139. 1940; Lall, Indian
Forest. 48: 181--185. 1942; Van Melle, Shrubs Trees Small Place
48, 54, 55, & 177. 1943; Mitra, Chandran, & Rao, Science Cult. 14:
315--317. 1949; S. C. & D. Datta, Indian Pharm. 6. 1950; Masamune,
Sci. Rep. Kanazawa Univ. 4: [Enum. Trach. 7]: 48--49. 1955; Patt-
naik, Journ. Bomb. Nat. Hist. Soc. 54: 149. 1956; J. Hutchins.,
Fam. Flow. Pl., ed. 2, 1: 395. 1959; Jacks. in Hook. £. & Jacks.,
ind, Kew., imp. 3, 2: 1214. 1959: Puri, Indian Forest Ecol. 1: 31,
a73,, 183, & 229, pl. 50 (1960) and 2: 535. & 657. 1960: Li... Woedy
Pl. Taiwan 16, 816, 832--834, & 973, fig. 334. 1963; Sarma, Drag.
Vier. 1..2963; laine & Blackwell, Pi. N.. Zeal... ed. 7, 42%, 371-—
373, & 499, fig. 139. 1964; Poole & Adams, Trees Shrubs N. Zeal.
230, 196h- Reard,..Desert pz ‘Gat. West. Austz..; Pl...) ed... 93. 1965;
Mallik & Chaudhuri, Bull. Bot. Soc. Beng. 22: 105--108, pl. 1.
1968; Guhabakshi & Naskar, Bull. Bot. Soc. Beng. 23: 175. 1969;
Hiremath & al., Journ. Karnatak Univ. [14]: 30--48. 1969; Nisa &

1981 Moldenke, Notes on Vitex 163

Qadir, Pakist. Journ. Forest. 19: 205, 208--216, & 218. 1969;
Beard, Descrip. Cat. West Austr. Pl., ed. 2, 113. 1970; Rao & Na-
rayana, Riech. Arom. Korperpflirg. 20: 215--216, 218, 220, & 222.
1970; Shrivastava & Sisodia, Indian Vet. Journ. 47: 170--175.
1970; M. L. & M. M. Dhar, Dhawan, Gupta, & Srimal, Indian Journ.
Exp. Biol. 9: 101. 1971; Farnsworth, Pharmacog. Titles 6 (6): xv
& title 10764. 1971; Patel, Forest Fl. Gujarat 25, 228, & 230--231.
197 Ts 04S. G Ss", "Biol. Abstr. 52° 36592: 1971; Molds; ‘Fitth Sum:
Hast, 54, 6b> 98, ‘107, Sid, 12, 1285 2384 179, +207 2085. 239%
2A GALS, 204',°2665° 2695 :270; 279, 2815 2905, 291); «2935, $2949 298.
363,” 300,, SLE, 313% 31859319. 328, 330) 3745°385;2 386)" S396
(1971) and 2: 534, 573, 660, 684, 710--716, 718--721, 723--726,
¥z6-—730, 7/81, 785, & 92735 1971; Anon. ¥ Commonw. Myc. “Inst. lmdex
Fungi, 3: 824. 1972; Fletcher in Hillier, Man. Trees Shrubs, ed. 2,
imp. ed., 416. 1972; Fong, Farnsworth, Henry, Svoboda, & Yates,
Lloydia 35: 35 & 46. 1972; Mahli & Trivedi, Quart. Journ. Crude
Drug Res. 12: [1927]. 1972; Mold., Phytologia 23° 211—-212, 3i5—
316, 413--416, 418--421, 423, 424, 427, 430, 434, 438, 503, 506,
& 512. 1972; Farnsworth, Pharmacog. Titles 9 (6): xii. 1973;
Vohora, Khan, & Afaq, Indian Journ. Pharm. 35: 100--102. 1973;
Vohora, Khan, & Afaq, Biol. Abstr. 57: 6733. 1974; Tsagarelli,
Bull. Acad.'‘Sci. ‘Georgian SSSR 78: 383 & 384.'°1975; Fo GoMey. 5
Journ. Arnold Arb. 57: 120 & 130. 1976; Babu, Herb. Fl. Dehra

Dun 14 & 18. 1977; Fosberg, Falanruw, & Sachet, Micronesica 13:
30. 1977; Kodanda Rao & E. & D. Venkata Rao, Biol. Abstr. 64:
6284. 1977; Fosberg, Sachet, & Oliver, Micronesica 15: 239. 1979;
Hocking, Excerpt. Bot. A.33: 86. 1979; Subramanian & Misra, Biol.
Beser. G7: 23358. 1979: Mold., Phytol... Mem.) 25°25 6,794.) beet oe
2, 24-—-28,' 445435 475'.54, 555" 573} 'S95 69, 139 d= 1 7G te ee
34,°96,5°98; 99, 101=-104)5 112,121, 123-126 ).830; 1356) 17a,
176, 179--182, 193, 195--202, 204--207, 209--218, 211--224, 228--
232, 234--236, 238, 239, 241--244, 246--248, 251--258, 265--267,
269, 271, 274, 275, 280, 282, 283, 287--290, 294, 297-=299, 302--
305, 309--311, 318, 319, 321, 323--325, 327--334, 339--343, 365--
369) 372, ..3795 395,°400,. 405, 413, 422, 423, -430, ‘43190435, “aaa,
442, 446, 456--460, 462, & 588--596. 1980; Mold., Phytologia 45:
485 & 491 (1980) and 48: 326, 413--419, 441, 442, 452--500, 505-
a105° 6° 5524 1988:

VITEX ACUMINATA R. Br.

Additional & emended bibliography: F. Muell., Sec. Syst. Cen-
sus Austr. PL. °22°173.~°18893.F. M. Bailey, »Cat? ‘Indie, Heres
Queensl. 35. 1890; Beard, Descrip. Cat. West Austr. Pl., ed. l,
93° (1965) and ed. 2, 113. 19703 Mold., Phytologia 45:°479. 1980;
Mold., Phytol. Mem. 2: 339, 458, & 588. 1980.

Additional citations: MOUNTED ILLUSTRATIONS: F. M. Bailey, Com-
preh. Cat. Queensl. Pl. fig. 362. 1909 (Z).

VITEX AGELAEIFOLIA Mildbr.
Additional bibliography: Mold., Phytologia 44: 224--225. 1979;
Mold., Phytol. Mem. 2: 215, 221, & 588. 1980

164 P HSE T:0 hb) O44, A Vol. 49, No. 2

VITEX AGELAEIFOLIA var. RUFULA Mold.
Additional bibliography: Mold., Phytologia 44: 225. 1979;
Mold., Phytol. Mem. 2: 221 & 588. 1980

VITEX AGNUS-CASTUS L.

Additional bibliography: Briq. in Engl. & Prantl, Nat. Pflan-
zentam., eds 1,24 (a)*«.133—-135,, 143, & 172s 18955). Butchins..,
Fame Ftows (Pic ted; 1, De 309;..fig. 263 19263, Savage, Cak>. Lind.
Heth 3 Lomd. 110. 1945+ Mold, : ‘Phytol. Mem. (22 137 Bap: 175-195" 22,
25S 26 OAL 4S; 4s 54 55 57, 59) 695 965 SERS, MOL, 905. 925,
171, 193, 195--200, 205, 241, 246, 254--256, 304, 318, 365, 366,
369, 372, 400, 436, 456--460, & 588. 1980; Mold., Phytologia 48:
415--417, 476, 477, 485, & 487--489. 1981.

Additional a itustrations: J. Hutchins., Fam. ‘Flow. Pi., eds,
1263095 £405 263242926:

VITEX AGNUS-CASTUS £. LACINIOSA (Ces.) Mold.
Additional bibliography: Mold., Phytologia 44: 342. 1979;
Mold., Phytol. Mem. 2: 197, 456--458, & 588. 1980.

VITEX AGNUS-CASTUS var. PSEUDO-NEGUNDO £. ALBIFLORA Mold.
Additional bibliography: Mold., Phytologia 44: 347. 1979;
Mold., Phytol. Mem. 2: 254 & 588. 1980.

VITEX AGNUS-CASTUS f£. ROSEA Rehd.
Additional bibliography: Mold., Phytologia 44: 347. 1979; Mold.,
Phytol. Mem. 2: 43, 197, 198, 366, 456, & 588. 1980.

VITEX AGNUS-CASTUS f£. VARIEGATA Mold.
Additional bibliography: Mold., Phytologia 44: 347--348. 1979;
Mold., Phytol. Mem. 2: 366, 456, & 588. 1980.

VITEX AJUGAEFLORA Dop
Additional bibliography: Mold., Phytologia 44: 348. 1979; Mold.,
Phytol. Mem. 2: 294, 366, & 588. 1980.

VITEX ALTISSIMA Let.

Additional bibliography: Savage, Cat. Linn. Herb. Lond. 110.
1945: Mold., Phytologia 48: 417--418 & 476. 1981.

Recent collectors describe this plant as a tree, 10--25 m. tall,
the leaves deciduous, the "petiole with a pulvinus at the base",
the young ones prominently winged, the leaf-blades "tomentose be-
neath, the calyx pinkish, the anthers "dark" or "black", the fila-
ments white, the fruit round, smooth, green, drupaceous, "turning
purple in age", and have encountered it in wet deciduous, semi-
evergreen, and dry deciduous forests, along streams, and on grassy
slopes on hillocks, at 820 m. altitude, in anthesis from March to
July, and in fruit in April and July. Ramamoorthy and Saldanha
both report it "common" in Mysore, India. The corollas are de-
scribed as having been "white but with the petals [=lobes?] and
lower lip purplish" on Saldanha 16875, "purple" on Saldanha 13425,
"Ghite" on Saldanha 13974, "white with a prominent purple lip" on

1981 Moldenke, Notes on Vitex 165

Saldanha 16938, "white with a blue lower lip"on Saldanha 13162,
"corolla lip purple" on Nicolson & al. HFP.158, and "lower lip of
corolla purple, side lobes and upper 2 lobes purplish-white" on
Saldanha 16553. Ramamoorthy, on his no. HFP.1917, comments
"petioles winged toward base", but the wings are not obvious on
the specimen of this collection so far seen by me (flowering).

Additional citations: INDIA: Karnataka: Jarrett, Saldanha, &
Ramamoorthy HFP.595 (W--2794856); Nicolson, Saldanha, & Rama-
moorthy HFP.158 (W--2794857), HFP.206 (W--2794858); Ramamoorthy
HFP.1917 (W--2794849); Saldanha 13162 (W--2794859), 13425 (W--
2794854), 13707 (W--2794850), 13974 (W--2794851), 14093 (W--
2794848), 16553 (W--2794852), 16875 (W--2794853), 16938 (W--
2794855).

VITEX ALTISSIMA f£. SUBGLABRA Thwaites
Additional bibliography: Mold., Phytologia 48: 418. 1981.
Kostermans refers to this plant as "very common" in dry val-
leys.
Additional citations: SRI LANKA: Kostermans 26727 (Lc).

VITEX ALTMANNI Mold.
Additional bibliography: Mold., Phytologia 44: 360. 1979; Mold.,
Phytol. Mem. 2: 319 & 588. 1980.

VITEX AMANTENSIS Pieper
Additional bibliography: Mold., Phytologia 44: 360--361, 386,
& 480. 1979; Mold., Phytol. Mem. 2: 228 & 588. 1980.

VITEX BREVILABIATA Ducke

Additional bibliography: Mold., Phytologia 45: 481. 1980; Mold.,
Phytol. Men. 25° E72. 5S6i271980'.

Additional citations: BRAZIL: Amazonas: Prance, Berg, Bisby,
Steward, Monteiro, & Ramos 18027 (Mu).

VITEX BREVIPETIOLATA Mold.
Additional bibliography: Mold., Phytologia 44: 392. 1979; Mold.,
Phytol. Mem. 2: 171 & 589. 1980.

VITEX BUCHANANIT J. G. Baker

Additional bibliography: Mold., Phytologia 44: 392--393. 1979;
Mold., Phytol. Mem. 2: 228, 236, °238; 239, 241, 457, & 539. 1980;
Mold., Phytologia 48: 466. 1981.

VITEX BUCHANANII vat. QUADRANGULA (GUrke) Pieper
Additional bibliography: Mold., Phytologia 44: 393. 1979; Mold.,
Phytol. Mem. 2: 228, 239, & 589. 1980.

VITEX BUCHNERI Glrke
Additional bibliography: Mold., Phytologia 44: 393 (1979) and
46: 30. 1980; Mold., Phytol. Mem. 2: 221, 224, & 589. 1980.

VITEX BUDDINGITI Mold.

166 P eee Loe Vol. 49, No. 2

Additional bibliography: Mold., Phytologia 44: 393. 1979; Mold.,
Phytol. Mem. 2: 319 & 589. 1980.

VITEX BUNGUENSIS Mold.
Additional bibliography: Mold., Phytologia 44: 393--394. 1979;
Mold., Phytol. Mem. 2: 228 & 589. 1980.

VITEX BURMENSIS Mold.
Additional bibliography: Mold., Phytologia 44: 394. 1979; Mold.,
Phytol. Mem. 2: 274 & 589. 1980.

VITEX CALOTHYRSA Sandw.
Additional bibliography: Mold., Phytologia 48: 419. 1981.
Additional citations: COLOMBIA: Vaupés: Schultes, Baker, & Cab-
rera 16171 \(Lc).

VITEX COFASSUS Reinw.

Additional bibliography: Mold., Phytologia 48: 454. 1981.

Craven & Schodde describe this species as a buttressed tree, 12--
35 m. tall, the trunk with a diameter of 14 cm. at breast height,
the bark stringy , flaky, pale-gray ,with a pale-brown blaze, or
striate , pale-green outside, pale-cream inside, the wool deep-
cream , the sapwood yellow "to gr ay-brown in heartwood", the leaves
dull mid-green or rather dull dark-green above, paler beneath, the
young fruit dull-green or dull mid-green, and have found it growing
in alluvial-freshwater tidal forests at sealevel and in primary
rainforests at 125 m. altitude, flowering and fruiting in Februazy.
The corollas are said to have been "mauve-purple" on their no. 704
(of which a wood sample was also collected) and "mauve-blue" on
their no. 4497.

Additional citations: NEW GUINEA: Papua: Craven & Schodde 704
(W--2896262); Schodde & Craven 4497 (W--2896062).

VITEX CYMOSA Bert.
Additional bibliography: Briq. in Engl. & Prantl, Nat. Pflanzen-
fam., ed. 1, 4 (3a): 143. 1895; Mold., Phytologia 48: 456. 1981.
Additional citations: BRAZIL: Amazonas: Byron 145 (W--2920718).

VITEX GLABRATA R. Br.

Additional & emended bibliography: F. Muell., Sec. Syst. Census
Austr. Pl. 1: 173. 1889; F. M. Bailey, Cat. Indig. Nat. Pl. Queensl.
35. 1890; Beard, Descrip. Cat. West Austr. Pl., ed. 1, 93 (1965)
and ed. 2, 113. 1970; Mold., Phytologia 48: 458. 1981.

VITEX LEUCOXYLON L. f.
Additional bibliography: Savage, Cat. Linn. Herb. Lond. 110.
1945; Mold. , Phytologia 48: 458, 460, 483 ,& 487. 1981.

VITEX LEUCOXYLON £. SALIGNA (Roxb.) Mold.
Additional bibliography: Briq. in Engl. & Prantl, Nat. Pflanzen-
fam., ed. 1, 4 (3a): 134. 1895; Mold., Phytologia 48: 460. 1981.

1981 Moldenke, Notes on Vitex 167

VITEX LOBKOWITZII Ettingsh.

Additional bibliography: Briq. in Engl. & Prantl, Nat. Pflanzen-
fam: , ed.. 1, 4 Ga)> 143% 1895; Mold. ; Phytologia 46: 2121980;
Mold., Phytol. Mem. 2: 369 & 591. 1980.

VITEX LUCENS T. Kirk

Additional bibliography: Poole & Adams, Trees Shrubs N. Zeal.
230 & 233. 1964; Mold., Phytologia 48: 461. 1981.

Additional illustrations: Poole & Adams, Trees Shrubs N. Zeal.
233. 1964.

Poole & Adams (1964) point out that the specific epithet awarded
this species refers to the glossy or lustrous upper leaf-surface.
They describe the species as a tree, to 20 m. tall, with plainly
tetragonal branchlets, the leaflets 3--5 in number, each 5--12 cm.
long, elliptic-oblong or obovate in shape, the flowers 2.5 cm.
long , the corollas red, and the drupes also red, 2 cm. in diameter.
They describe its distribution in New Zealand forests from North
Cape to the Mahia Peninsula and Cape Egmont.

Additional citations: MOUNTED ILLUSTRATIONS: Poole & Adams,
Trees. Shrubs: Ns. Zeal.) 233. 1964.(Z, Z)’.

VITEX MOLELTS H.B.K.

Additional bibliography: Walp., Ann. Bot. Syst. 1: 545. 1849;
Mold., Phytologia 48: 462. 1981.

Additional citations: MEXICO: Guerrero: Lopez Forment 928 (Me--
284806).

VITEX NEGUNDO L.

Additional bibliography: Savage, Cat. Linn. Herb. Lond. 110.
1945; Mold., Phytologia 48: 466--500. 1981.

The Ramamoorthy HFP.374 and Saldanha 12470, distributed as
typical V. negundo, actually represent its var. trifoliolata Mold.

VITEX NEGUNDO var. CANNABIFOLIA (Sieb. & Zucc.) Hand.-Mazz.

Additional bibliography: Walp., Ann. Bot. Syst. 1: 542. 1849;
Mold., Phytologia 48: 487, 489, & 492--500, fig. 1--5. 1981.

In the previous supplement to my Additional Notes on this genus
I began a quotation from the late 0. F. Cook's description of
anisophylly in what was taken to represent this variety [but actu-
ally is var. intermedia (P'ei) Mold.]. His description continues
as follows: "Even when the leaf that should be underneath in the
normal position is brought to the upper side by the twisting of
the branch the growth of the larger leaf is not inhibited by its
more exposed position, nor does the smaller leaf take advantage of
its more protected position to increase in size. In any particu-
lar case it is impossible, of course, to say that the leaves of a
pair are more or less unequal than they might have been if grown
in some different position. But it is evident that there is no
general loss or even an apparent reduction of the normal aniso-
phylly as a result of modification of position and exposure.

"A further possibility of securing evidence of this kind by
observing the behavior of leaves of equal pairs has been consider-

168 FAR TO MOoema Vol. 49, No. 2

ed, but without securing any very definite results. In some
cases there seemed to be quite a tendency to inequality between
the leaves of the equal pairs when the exposure was abnormal, but
in other cases there was no apparent difference. Equal pairs
standing in vertical positions seemed to be as nearly equal as
where they had lateral positions on adjoining branches. It was
thought at first that the tendency to bilateral asymmetry disap-
peared when leaves of equal pairs developed in vertical positions,
but this was not always the case. When branches were turned
over by twisting the larger divisions were developed on the side
of the leaf that had the superior position, even in pairs of

Z
=
'
Zz
a
Z
Z

}

Fig. 6. Paired leaves showing difference in form as well as size

1981 Moldenke, Notes on Vitex 169

leaves that showed an unusual tendency to inequality of size.
Thus in figure 8, representing a pair of leaves grown in reversed
position on a twisted branch, the larger divisions were produced
on the sides of the leaves that has a superior or more exposed
position. That greater exposure has no pronounced effect of in-
hibiting the development of leaves in more exposed positions was
also shown in many cases where the lower leaf was smaller in a
normally equal pair, developed in a vertical position.

Fig. 7. Paired leaves showing differences in form as well as size
in case of shade leaves

"Instead of ascribing the anisophylly of vitex to direct ef-
fects of gravitation or light upon the development of the parts
of the plant it seems to have more connection with the structure
and habits of growth of the plant. In view of the normally as-
cending position of the branches it is easy to understand the ad-

170 P oY? OL 08S A Vol. 49, No. 2

aptive advantage of larger development on the outer or more ex-
posed sides of the branches and of smaller development of leaves
and branches in the more unfavorable axillary position, between
the lateral branch and the upright parent shoot. Whether the in-

Fig. 8. An asymmetric pair of equal-sized. leaves

1981 Moldenke, Notes on Vitex 171

feriority of the leaves and buds of the upper or inner face of

the branch is due directly to the less favorable position of
their primordia on the axillary side of the branch or represents

a specialized character in heredity may be difficult to determine,
but it is evident that the differences depend upon the peculiari-
ties of the plant rather than upon the influences of the external
environment. The fact that the inequality diminishes or disappears
in shoots that start in an upright position is not a reason for
looking upon the specializations of oblique and horizontal

branches as direct results of environmental influence. On the
contrary, it may serve as a further indication that the inequality
of the sides of the lateral branches represents a specialized cm-
dition of heredity, but capable of being reversed like other states
of expression of characters. That the plants are capable of pro-
ducing symmetrical upright shoots only shows the more clearly that
the peculiarities of the lateral shoots are in the nature of spec-
jializations. That different kinds of shoots can be produced in
different positions does not prove that the changes are caused by
the external conditions, but is an evidence of the adaptive abil-
ity of the plant.

"One more possibility is worthy of consideration, that the very
pronounced anisophylly of Vitex is a product of two factors. The
inequality of the two sides of the branches, as considered above,
may be supplemented or intensified by an inequality of internodes.
If we consider that the opposite positions of the leaves repre-
sents a suppression of alternate joints or internodes of the stem
it becomes possible to understand that the large joints might natur-
ally tend to produce larger leaves and branches than the alterna-
ting small joints.

"Even when branches are brought into positions where the larger
leaves of the unequal pairs should be uppermost, as when shoots are
given off near the tops of large branches that are bent over, the
usual position of the leaves is often regained through the twisting
of the branches by the greater weight of the leaves on one side,
tending to this side underneath.

"The idea that too much light may inhibit the growth of the
leaves on the upper sides of the branches may find some support in
the fact that anisophylly is often unusually pronounced on shoots
that arise on very exposed parts of the plants, though all the
leaves tend to be small on such branches, in comparison with the
size attained in lower or more sheltered places. But if this factor
were significant the inequality should disappear on branches that
are heavily shaded (Fig. 7), or where the lower side receives more
light than the upper, but even in such cases the inequality often
remains very great."

All things considered , it seems to me that the plants with which
Cook was experimenting here are neither typical V. negundo L., nor
its var. cannabifolia (Sieb. & Zucc.) Hand.-Mazz. nor var. hetero-
phylla (Franch.) Rehd., but represent the var. intermedia (P'ei)
Mold.

Material of v. negundo var. cannabifolia has been misidentified
and distributed in some herbaria as V. heterophylla Roxb., V. incisa

172 PHT TCLoOCs & Vol. 49, No. 2

Lam. ,V. negundo L., V. negundo var. heterophylla (Franch.) Rehd.,
V. negundo var. incisa (Lam.) C. B. Clarke, and V. quinata (Lour.)
F. N. Will. On the other hand, the Chow 7, distributed as var.
cannabifolia, probably is better regarded as var. intermedia
(P'ei) Mold.

Additional citations: INDIA: Punjab: T. Thomson s.n. [Punjab,
1--4000 ped.] (Mu--655). Tamil Nadu: G. Thomson s.n. [Maisor &
Carnatic] (Mu--656). CHINA: Fukien: Cheng 3366 (Mu); En 2810
(Mu); Ging 5143 (Ws),6651 (Ws). Kwangsi: Wan & Chow 79016 (N).
Kweichow: Tsiang 8518 (Mu). Shantung: Zimmermann 442 (Mu--3958).
CHINESE COASTAL ISLANDS: Lantau: Hu 10244 (W--2731441). THAILAND:
Zimmermann 2 (Mu--3964). MALAYA: Johore: Poore 296 (K1--296).
Perak: Chin 843 (K1--15297). MALAYAN ISLANDS: Langkawi: Abd 15
(K1--2015). JAPAN: Honshu: Maximowicz s.n. [Yokohama 1862] (Mu--
1518); Siebold s.n. [in Japonia legit] (Mu--626--isotype, Mu--
627--isotype). CULTIVATED: India: Herb. Hort. Bot. Calcut. s.n.
(Pd). New York: Moldenke & Moldenke 11865 (N). MOUNTED CLIPPINGS:
Walker, Pl. Okin. South. Ryuk. 894. 1976 (W).

VITEX NEGUNDO var. DENSIFLORA Haines, Bot. Bihar Orissa, ed. 1, 4:
1126. 1922.

Additional & emended bibliography: Haines, Bot. Bihar Orissa,
edijids es 72 (1922), and ed. 2,2: 746. 1961; Mold., Phytotogia.15;
308--309. 1967; Mold., Fifth Summ.: 2 279 (1971) and 2: 927. 1971;
Mold., Phytol. Mem: 2: 256, 266, 459, & 592.1980.

VITEX NEGUNDO var. HETEROPHYLLA (Franch.) Rehd.

Additional & emended synonymy: Vitex sinuata Raeusch., Non.
Bot., ed. 3, 182, nom. nud. 1797; Steud., Nom. Bot., ed. 1, 888,
nom. nud. 1821. Vitex negundo Bot. Mag. ex Sweet, Hort. Brit.,
eds 13085323, invusyn. 1826-([not V. negundo 1.,.1753,. nor :Lour, 5
1934, nor Noronha, 1790, nor Royle, 1919, nor Willd., 1918]. Vitex
iveisa Fat ).Sns, Contrib. Mat, Med, China: 227, sphalm, 1871.. Vi-
tex incisa var. heterophylla Franch., Nouv. Arch. Mus. Hist. Nat.
Paris, ser. 2, 6: 112. 1883. Vitex negundo sensu Curtis apud
Rehd., Journ. Arnold Arb. 28: 258, in syn. 1947. Vitex negundo
heterophylla Blackburn, Trees Shrubs East. N. Am. 303. 1952. Vi-
tex negundo var. incisa Clarke ex Roberty, Pet. Fl. Ouest-Afr. 178.
1954. Vitex negundo incisa (Bunge) Clarke ex Enari, Ornament.
Shrubs Calif. 170, in syn. 1962. Vitex negundo cv.'Incisa' Enari,
Ornament. Shrubs Calif. 170. 1962. Vitex negundo 'Heterophylla'
Sherk & Buckley, Ornament. Shrubs Canada 164. 1968. Vitex negundo
incisa (Lam.) Clarke ex Mold., Fifth Summ. 2: 724, in syn. 1971.
Vitex negundo var. heterophylla Rehd. apud R. G. & M. L. Br.,

Woody Pl. Md. 288 & 289. 1972. Vitex laciniata "Hort. ex Schauer"
ex L. H. & E. Z. Bailey, Hortus Third 1162, in syn. 1976. - Vitex
negundo var. incisa "(Lam.) C. B. Clarke in Hook. f.'' ex Mold., Phy-
tol. Mem. 2: 459, in syn. 1980.

Additional & emended bibliography: Lam., Encycl. Meth. Bot. 2:
612. 1788; Raeusch., Nom. Bot., ed. 3, 182. £797 3° Desf.% Tape. Ecol.
Pet., eds hl; 53. 18045" Walid., Enum. Pils Hott, Borel: 2: 660. 1809;

1981 Moldenke, Notes on Vitex 133

Balbis, Cat. Stirp. Hort. Acad. Taur. 81. 1813; Roxb., Hort.
Bene. , imp. 1, 46...1814: Desf., Tabl. Ecol. Bot.,fed.+ 2564. S685?
Sweet, Hort. Brit .cwed. 1s \322..18265) Wall .> Nemese. List (424.

no. 1746. 1829; Loud., Hort. Brit., ed. 1, 246. 1830; Sweet, Hort.
Brit... edx,2,. 416.1630; Wall.,;. Numer... List; 86, .no-..1746D., 1831:
fang... Hor’, Brit.) eda2, 246..1832:, Roxb. Fie Ind. jveds.2 4.4mp-
1, 3: .72--73. 1832; Bunge, Enum. Pl. Chin.-Bor. 52. 18335 Bunge,
Mém. Div. Sav. Acad. Sci. St. Pétersb. 2: 216. 1835; G. Don in
Loud.,- Hort. Brit.,..ed. 3».246. 1839; .G. Don dn-Sweet,,, Hort.
Beth. sed. 5. 551... 1839: Spach, Hist... Nak. Veg. Phan..9> 232.
1840;, Hassk., Cat. Pl. Hort. Bot. Bogor. Cult.Alt.. 134., 1844;
Voigt, Hort. Suburb. Calc. 469. 1845; Buek, Gen. Spec. Syn. Can-
doit. 3:, 502...1858; Dupuis, Nouv. Fi. Usuel.. Med. 2: 298. 1860: A.
Gray..Man.. Bot....North. .U.\ 5... ed«3, ixviis(1862),..ed. 4A,, amp. 8,
Ixvii (1863), and ed. 4, imp. 2, lxvii. 1864; A. Gray, Field For.
Gard. EBot.>s ed. 1, imp. .1, 243, (1868). and. ed.. 1, imp.. 2,243;
£369> A. Gray; Man. Bot. North. U. S., ed. 4) 4mp.. 3, Jxvii. 1970;
A. Wood, Am. Bot. Flor., ed. 1, imp. 1, 237 (1870) and ed. l,

Pipi 2e de tore F..P.. Sma,,Contrib.. Mats,.Med. Cirina, 277. 187i;
A. Wood, Am. Bot. Flor., ed. 1, imp. .3,. 237 (1872). ed... Lt, imp. 4,
23h A813), and ed..1, imp..5, 23/.. 18/4;> Boxbh.4 El... ind. ied. 2,
rap. 2, 482.1874; A. Wood, .Am. Bot. Flor:,..ed. 2, imp...6, 23/7 -
$75 Ae Cray. Field For...Gard.. Bot. , eded, -imps,,35 243. 1680;
Franch., Nouv. Arch. Mus. Hist. Nat. Paris, ser. 2, 6: 112. 1883;
Franek. , PL. David... imp. 1, 12 232..1884¢ Of R...Willts.4n A.
Wood., Am. Bot. Flor., ed. 2, 237. 1889; Forbes & Hemsl., Journ.
Sinn= »See>. Lond: -Bot...26,:[ind. Fl. Sin... 2) 2457... 1890; Vesa an
Vilm., Blumengurt. 1: 829. 1895; K. Schum. & Lauterb., Fl. Deutsch.
Schutzgeb. Slldsee 524. 1900; Diels, Fl. Cent.-China 549. 1902;
Stuart, Chin. Mat. Med. 1911; Dunn & Tutcher, Kew Bull. Misc.

inf. Addit: Ser...10: 204. 1912: Rehd. in, Sare., PL. Wite.«3: 43,
373, & 374. 1916; H. Hallier, Meded. Rijks Herb. Leid. 37: 44.
1918; Lazaro e Ibiza, Compl. Fl. Espafi., ed. 3, 3: 298. 1921;
Haines, Bot. Bihar Orissa, ed. 1, 4: 712. 1922; Nakai, Fl. Sylv.
Kor. 14: 38, pl. 12. 1923; Wangerin, Justs Bot. Jahresber. 51 (1):
554. 1923; Haines, Bot. Bihar Orissa, ed. 1, 6: 712. 1924; Parker,
Forest Fl. Punj., ed. 2, 395. 1924; Janssonius, Mikrogr. Holz.
812. 1926; E. D. Merr., Lingnan Sci. Journ. 5: 158. 1927; Vansell
& Eckert, Univ. Calif. Agr. Exp. Sta. Bull. 517, imp. 1, 52 & [60].
1931; Fedde, Justs Bot. Jahresber. 51 (2): 385. 1933; L. H. & E.
Z. Bailey, Hortus 639. 1935; Makins, Ident. Trees Shrubs 259.
1936; Parks, Texas Agr. Exp. Sta. Bull. 155: 113. 1937; W. Trelease,
Pl. Mat. Decorat. Gard. Woody Pl., ed. 5, imp. 1, 146. 1940;

Fedde & Schust., Justs Bot. Jahresber. 60 (2): 576. 1941; Vansell
& Eckert, Univ. Calif. Agr. Exp. Sta. Bull. 517, imp. 2, 76.

1941; Everett, Cat. Hardy Trees Shrubs 120. 1942; E. L. D. Sey-
mour, New Gard. Encycl., ed. 3, 1292. 1944; Savage, Cat. Linn.
Herb. Lond. 110. 1945; E. L. D. Seymour, New Gard. Encycl., ed. 4,
1292 (1946) and ed. 5, 1292. 1951; Blackburn, Trees Shrubs East. N.
Am. 303. 1952; Pételot, Pl. Méd. Camb. Laos Vietn. 2; 248 (1954)
and 4: 171. 1954; Roberty, Pet. Fl. Ouest-Afr. 178. 1954; Bean in
Chittenden, Dict. Gard., imp. 1, 4: 2249 & 2250. 1956; Wyman, Shrubs

174 PH? TOL OL Ts Vol. 49, No. 2

Vines Am. Gard. 351 & 352. 1956; Viertel, Trees Shrubs Vines,
imp. 1, no. 406. 1959; Haines, Bot. Bihar Orissa, ed. 2, 2: 746.
1961; Enari, Ornament. Shrubs Calif. 170. 1962; E. L. D. Seymour,
New Gard. Encycl., ed. 6, 1292 (1963) and ed. 7, 1292. 1964; Bean
in Chittenden, Dict. Gard., imp. 2, 4: 2249 & 2250. 1965; Everett,
Reader's Digest Compl. Book Gard. 447 & 661. 1966; Mold., Phyto-
logia 17: 15, 17--19, 22, 23, 28, & 29. 1968; Mold., Résumé Sup-
pl. 16: 29 (1968) and 17: 8 & 12. 1968;Sherk & Buckley, Ornament.
Shrubs Canada 164. 1968; W. Trelease, Pl. Mat. Decorat. Gard.
Woody Pl., ed. 5, imp. 2, 146. 1968; G. W. Thomas, Tex. Pl. Eco-
log; sma, 76, 1969" “Yranch;, Pi, David., imp. °22 24 2322 1970-
McGourty [editor], 1200 Trees [Plants Gard. 26 (2):] 53. 1970;
Mold. in Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex.
Res. Found. 6:] 1340 & 1878. 1970; E. L. D. Seymour, New Gard. En-
eyel., ed. @, 1292, "1970; Viertel, Trees ‘Shrass Vines, tie. 2, 02
406. 1970; Hartwell, Lloydia 34: 388. 1971; Mold., Fifth Sum. 1:
au, OF, 1265290. 316, 3742 585, & 306 (1971) “am 2. 554, 710;
TiZD Iie JLo s, ILI; 1 20, F255 1245728, & O27 Tete Priactcer,
Detect. Sem. Spor. P1, Hort. Bot. Univ. Hung. 59. 1971; Roxb.';
Pi, “sid, Gd. 2, imp, 3, 462.°4971: Wyman, Gard, “ineye.., imp. 1;
eft 4 0972) ane tmp. 2, ‘1171. (29722 ‘Re 6s 8 Ms ok. “er. s Woaay Pr.
Md. 288 & 289. 1972; Encke & Buchheim in Zander, HandwUrterb.
Pflanzennam., ed. 10, 525. 1972; Farnsworth, Pharmacog. Titles 7
(4)? axvi & 222. 19723 Moid.., Phytologia 23: 427 & 438.1972: "R.
R. Stewart, Annot. Cat. in Nasir & Ali, Fl. West. Pakist. 608.
1972; Gibbs, Chemotax. Flow. Pl. 3: 1754. 1974; Howes, Dict. Use-
ful Pl. 52. 1974; Whitney in Foley, Herbs Use Delight [204].
1974; [Farnsworth], Pharmacog. Titles 7, Cum. Gen. Ind. [118].
1975; Kooiman, Act. Bot. Neerl. 24: 462. 1975; Lépez-Palacios,
Revist. Fac. Farm. Univ. Andes 15: 101. 1975; Mold., Phytologia
387 S00. 19752 “Wyman, Gard. ‘Sourn. 25: [45})- 6.46. 1975) LR: &
E. Z. Bailey, Hortus Third 1161 & 1162. 1976; Mold., Phytologia
94-°279; 1976; C.-W. ‘Ei, China Reeonstr.''27 (2): 4. 1978: Wang,
Act. Entomol. Sin. 21: 343--344. 1978; Mold., Phytologia 44: 225.
1979; Wang, Biol. Abstr. 68: 4667. 1979; Mold., Phytol. Mem. 2:
Ze, 4/4 35. 96, 260, °309, 367, 458, 4593" & S922 19603 Roxb,
Hort. Beng., imp. 2, 46. 1980; Mold., Phytologia 48: 489. 1981,

Additional illustrations: Bean in Chittenden, Dict. Gard.,
imp. 2, 42° 2250 (7956) and imp: 2,4: °22507 1965; RK. G. °& MeL.
Br., Woody Pl. Md. 289. 1972.

This is a natural variety, native to northern China and Mon-
golia (Baileys, 1976) or only northern China (Bean, 1976, Spach,
1840). Desfontaines (1804), Sweet (1826), and Ldzaro e Ibiza
(1921) give its native distribution merely as "China". There is
no justification whatever in changing its status to that of a
cultivar as some recent writers have proposed! The same is true
of the other infraspecific taxa in V. negundo, V. agnus-castus,
etc. Modern writers are all too prone to regard all taxa grow-
ing in cultivation as ipso facto cultivars -- in many cases this
is not at all the case!

Wyman (1956) states that the present variety was first in-
troduced into cultivation in 1750 from "China or Korea", but

1981 Moldenke, Notes on Vitex 175

Sweet (1826) and Don (1830) give the date as 1758 from "China".
Voigt (1845) lists it as in cultivation in Calcutta in 1845,
while Lazaro e Ibiza (1921) record it from Spanish gardens. Ro-
berty (1954) tells us that this is the "form" of Vitex negundo
["& folioles profondément dentées"] generally cultivated in wes-
tern Africa. Enari (1962) reports that in California it is usu-
ally cultivated as "Vitex lacimiata", having "deeply toothed, in-
cised or cut leaflets". Sherk & Buckley (1968) claim that it is
hardier than V. agnus-castus and is even root-hardy as far north
as Ottawa, Canada, where it blooms in September and October.
Priszter (1971) offered its seeds to the horticultural trade from
cultivated plants in Hungary. Everett (1942) found the "castane-
aefolia" form cultivated in the Lu Shan Arboretum. He claims
that var. heterophylla will grow in the Rocky Mountain region of
western North America. The Meebold collection from Vienna, Aus-
tria, is said to have come from seeds imported from "tropical
Asia".

Spach (1840) says "Cette espéce, originaire du nord de la
Chine, se cultivé fréquemment comme arbrisseau d'ornement; elle
est tres-rustique; sa floraison a lieu en aout et septembre."
Clarke (1885) says of it that it occurs "Throughout India. Dis-
trib. E. Asia. -- The extreme pinnatifid form of this var. is
Chinese; the wild Indian examples are crenate-serrate, i.e. in-
termediate.

Voss (1895) says of it: “blassblau, die Bl¥ttchen sind fieder-
lappig. Blltezeit: Juli, August. -- Verwendung in Sommer als an-
genehme, im Freien aufzustellende KibelstrYucher. tberwinterung
im Kalthaus oder hellen Keller. Vermehrung durch Stecklinge,
Grundisprosse und Wurzelschnittlinge. Ansucht aus Samen, die
lauwarm zu halten. -- Keimkraft der Samen 1--2 Jahre; Keimung
frischer Samen erst nach 1 Jahr; Yltere liegen noch 14nger."

Smith (1871) records the name , "man-king", for this plant in
China ,where, he says, the fruits are imported from Honan, Shen-
si, and Pehchihli. He describes these fruits as "berries globu-
lar , black, nucumentaceous, usually covered with remains of the
calyx or mixed with its dried leaves. Interior is white, ligne-
ous, and made of 4 carpels in a state of adhesion". He says
that they have little taste or smell and are “inert as sold in
Hankow". The material is prescribed in the treatment of head-
aches, catarrh, watery eyes. It is supposed to promote the
growth of the beard - "that great object of the middle life of
every Chinaman". Li (1978) reports that in animal tests and
in clinical use the same results obtain as from the use of var.
Cannabifolia fruit. Gibbs (1974) asserts that syringin is ab-
sent from the stems and that negative results are obtained with
the HCl/methanol test.

Parks (1937) calls the variety the "Japanese vitex" and says
that "It grows more as a bush than the other species and has
light green leaves with no odor. The plant bears throughout the
summer large numbers of light blue flowers. These have the habit
of opening about ten o'clock A. M. and remaining open until after
dark. It is a fine honey plant and it is not uncommon to see

176 Puy Trot oe sz Vol. 49, No. 2

bees working it until the darkness causes them to leave or to
remain on the plant. This is a species that should be very large-
ly procured from nurseries [in Texas] as it is a most valuable
ornamental." Vansell & Eckert (1931) also describe the corollas
as blue, adding that the color of the honey is greenish-white,

but that the value of the plant as a source of nectar and pollen
for bees is minor.

Stewart (1972) regards this variety as identical with typical
Vitex negundo L., but, of course, it is very distinct. Blackburn
(1952) describes typical V. negundo as having its leaflets rath-
er shallowly toothed "or occasionally entire". while var. hetero-
phylla has the leaflets "deeply toothed or cut" and f. multifida
has them "cut into narrow segments reaching almost to the midrib"

The Baileys (1976) describe the leaflets of var, heterophylla
as "smaller, deeply toothed or cut". Dunn & Tutcher (1912) say
for the typical V. negundo "Leaflets large, entire or coarsely
toothed" and for our variety "Leaflets small, deeply cut". Par-
ker (1924) avers that "The form with more ovate-lanceolate coarse-
ly serrated leaves is var. incisa (sp. Lamk.) [now known as var.
heterophylla]; it passes into the typical form and both kinds of
foliage may sometimes be found on the same plant" [the presentday
var. intermedia]. Atkins (1936) calls it "A smaller shrub, with
smaller fl.-panicles and berries". The fruits, of course are
drupes, not berries.

Hallier (1918) describes the present variety as "Ein junger,
noch nicht bllhenden BY¥umchen einer wahrscheinlich neuen dem V.
incisa verwandten Art mit fllnfgliedrigen BlY¥ttern und gestielten,
nach Art der Quercus palustris scharf fiederlappigen Bl¥ttchen
fand ich am 21.11.1904 auf den Boden des gelichteten Hochwaldes
hinter San Ramon auf S.W.-Mindanao (no. 4712, Hb. Hamb., Manill.,
E7-83),

Van Melle (1943) regards the "deeply toothed or incised" form
as typical V. negundo and the "pinnately divided" form as var.
incisa [=heterophylla]". Actually, the former is var. hetero-
Phylla, while the latter is f. multifida.

Recent collectors describe Vitex neguz:do var. heterophylla as
a shrub, 1.5--2.5 m. tall, with diffuse branches and aromatic
leaves and have found it growing in rocky clay limestone soil, at
200 m. altitude, in flower in May and July. Liogier encountered
it "en manigua a orilla de la carretera". The corollas are said
to have been "lavender" on Fogg s.n. and "R[oyal] Hf[orticultural]
S[ociety] Aster violet, the lip 38, the lateral lobes 38/1, the
upper lobes 38/3" on Huttleston 1709.

Common and vernacular names recorded for this taxon include
"Chinese chaste tree", "cutleaf chaste-tree", "cutleaf chaste
tree", "cut-leaved chaste tree", 'cut-leaved chaste-tree",
"“cut-leaved chastetree", "gattilier incisé", "Japanese vitex",
"man-king", "negundo", "pimentillo", and "vitex incisé".

The Schumann & Lauterbach (1900) reference in the bibliography
(above) is usually cited by its titlepage date, "1901", but the
New York Botanical Garden Library received its copy of the work
on December 7, 1900.

1981 Moldenke, Notes on Vitex pI /

The plant described and illustrated by Wyman (1975) as this
variety actually is Vitex agnus-castus f. latifolia (Mill.)

Rehd. On the other hand, the plant described and illustrated by
Viertel (1959, 1970) as "V. negundo incisa" actually seems to be
V. negundo var. intermedia (P'ei) Mold.

he Ve .al ba (HOT... *Ws,talba ‘Lam. ; Vs alba vars incisa shorts,
and V. incisa var. alba Hort., listed as synonyms of V. negundo
var. heterophylla by me previously (1957) are now regarded by me
as belonging to the synonymy of V. negundo var. heterophylla
f. alba (Garr.) Mold:

Vitex chinensis Mill., generally accepted as a synonym of V.
negundo var. heterophylla, is based on an unnumbered P. Miller
specimen in the British Museum herbarium from the Chelsea Physic
Garden, determined by R. Brown as "Vitex incisa'". The specimen
is in young fruit and on the reverse side of the sheet Solander
has written "Miller, ex Hort."

Wang (1978) reports that the bee, Scolia clypeata Sickman,
secures nectar from the plant here under discussion, as well as
from Tamarix chinensis, Mentha arvensis, M. spicata, Melilotus
suaveolens, Medicago sativa, Solidago sp., Rudbeckia laciniata
var. hortensis, and Salix matsudana, in the area of Peking, China.
Schumann & Lauterbach (1900) cites Lesson s.n. from Ceram.

Material of this taxon has been widely misidentified and dis-
tributed in herbaria as typical V. negundo L. On the other hand,
the Chang 8253, Ging 5143 & 6651, and Zimmermann 442, distributed
as var. heterophylla, actually represent var. cannabifolia
(Sieb. & Zucc.) Hand.-Mazz., while F. Murray 1148 and H. Rhodes
47-64-122 SM.84A are V. negundo var. heterophylla f£. multifida
(Carr.) Rehd. and E. M. Alexander 35, Allard 11390, Cutler 5043,
and Ging 5388, 5956, & 6740 are var. intermedia (P'ei) Mold.

Additional citations: OKLAHOMA: Payne Co.: G. E. Hall 22 (Au--
122931). Tulsa Co.: Hays 91 (Au--122930). HISPANIOLA: Haiti: A.
H. Liogier 21397 (N). CHINA: Hopeh: Herb. Inst. Bot. Acad. Sin.
75084 (Ac, Ac, N); Tatarinow s.n. [Fl. Pekin.] (W--2525123, W--
2560205). PHILIPPINE ISLANDS: Luzon: Merritt & Darling s.n.
[Herb. Philip. Forest. Bur. 14051] (W--711498). CULTIVATED: Ari-
zona: Thornber 7369 (Au--245703). Austria: Meebold 17404 (Mu).
California: Jerabek s.n. [Balboa Park, June 1945] (Sd--36461),
s.n. [Balboa Park, Sept. 1945] (Sd--37000). Egypt: Mahdi s.n.
[16/7/1963] (Gz, Gz), s.n. [10/11/1963] (Gz, Gz), s.n. [25/6/
1965] (Gz, Gz), s.n. [4/6/1967] (Gz, Gz). England: P. Miller s-n.
[Chelsea Physic Gard.; Bailey Hort. neg. 5055] (Ba--photo). Flo-
rida: Godfrey 55555 (Ld). New Zealand: Sykes 88/62 (Nz--125876).
Pennsylvania: Fogg s.n. [August 7, 1969] (Ba); Huttleston 1709
[Longw. Gard. 571138] (Ba). Texas: Rowell 5801 (Au--187069); R.
Runyon 65 (Au--270235), 3552 (Au--270201). Virginia: Allard
11390 (Ws).

VITEX NEGUNDO var. HETEROPHYLLA f. ALBA (Carr. ) Mold.
Additional & emended synonymy: Vitex incisa alba Desf., Tabl.

Ecol. Bot., ed. 1, 53. 1804. Vitex alba Hort. ex Mold., Prelim.

Alph. List Inv. Names 49, in syn. 1940. Vitex incisa var. alba

178 PH Yi 0 4.0uG ek Vol. 49, No. 2

Hort. ex Mold., Prelim. Alph. List Inv. Names 51, in syn. 1940.
Vitex alba Lam. ex Mold., Phytologia 5: 502, in syn. 1957. Vitex
alba var. incisa Hort. ex Mold., Phytologia 5: 502, in syn. 1957.
Vitex negundo alba Hort. ex Mold., Phytologia 5: 502, in syn. 1957.

Additional bibliography: Desf. , Tabl. fcol. Bot., ed. 1, 63
(1804) and ed. 2, 64. 1815; Mold., Phytologia 17: 19. 1968; Mold.,
Fifth Summ. 1: 374 & 385 (1971) and 2: 927. 1971; Mold., Phytol.
Mem. 2: 367 & 592. 1980.

VITEX NEGUNDO var. HETEROPHYLLA £. MULTIFIDA (Carr.) Rehd.

Additional synonymy: Vitex negundo f. multifida Rehd. ex L. H.
Bailey, Stand. Cycl. Hort. 6: 3574 [as "Vv. Ww. £. mitifida"]. 1917.
Vitex negundo heterophylla f. multifida Blackburn, Trees Shrubs
East. N. Am. 303. 1952.

Additional bibliography: C. B. Clarke in Hook. f., Fl. Brit.
India 4: 584. 1885; Dunn & Tutcher, Kew Bull. Misc. Inf. Addit.
Ser. 10: 204. 1912; Lazaro e Ibiza, Compl. FiJ Espan., ed. 3, 3:
298. 1921; Van Melle, Shrubs Trees Small Place 48, 55, & 177. 1943;
Blackburn, Trees Shrubs East. N. Am. 303. 19523; Mold., Phytologia
17: 18 & 19. 1968; Mold., Résume Suppl. 16: 29 (1968) and 17: 12.
1968; Mold., Fifth Summ. 1: 291, 374, 385, & 386 (1971) and 2:
116, J19, 723,°924,, /81,. 785, & 927. IS7LS Motdey Paycol. Mem, 2:
280, 367, & 592. 1980.

This form is a naturally occurring one from the Peking area of
Hopeh, China, now widely cultivated (and even escaped), with the
leaflets small and pinnatifidly cut or divided into narrow and
distant. segments reaching almost to the midrib. It appears to be
the form regarded as "var. incisa" by Van Melle (1943) and by Dunn
& Tutcher (1912). It is included in "V. incisa" by Lazaro e Ibiza
(1921). Clarke (1885) regarded it as "the extreme pinnatifid.....
Chinese" form of V. negundo var. incisa as distinguished from
"the wild Indian......crenate-serrate" form.

Murray describes the plant as 2 m. tall and found it in fruit
in September.

Additional citations: CULTIVATED: Pennsylvania: FE. Murray 1148
(Ba).

VITEX NEGUNDO var. INTERMEDIA (P'ei) Mold.

Additional synonymy: Vitex negundo f. intermedia (Pei) Mold.
ex Venkatareddi, Bull. Bot. Surv.India 11: 258. 1969. Vitex
negundo var. intermedia (P'ie) Mold. ex G. W. Thomas, Tex. Pl.
Ecolog. Summ. 78, sphalm. 1969. Vitex negundo intermedia [(P'ei)
Mold.] ex Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex.
Res. Found. 6:] 1878. 1970. Vitex negundo intermedia (P'ei) Mold.,
Fifth Summ. 2: 724, in syn. 1971.

Additional bibliography: C. B. Clarke in Hook. f., Fl. Brit.
India 4: 584. 1885; Liogier, Rhodora 67: 350. 1965; Mold., Phyto-
logia 17: 17--20. 1968; G. W. Thomas, Tex. Pl. Ecolog. Sum. 78.
1969; Venkatareddi, Bull. Bot. Surv. India 11: 258. 1969; Mold.
in Correll & Johnston, Man. Vasc. Pl. Tex. [Contrib. Tex. Res.
Found. 6:] 1340 & 1878. 1970; Viertel, Trees Shrubs Vines no. 406.
1970; Mold., Pitth Summ. 1: 31, 61,’ 96, 107, 112; 207, 266, 279,

1981 Moldenke, Notes on Vitex 179

291, 293, 294, 298, 313, 328, 331, & 374 (1971) and 2: 711, 719,
#23, (724, 6 927. 1971; Mold... Phytologia 23: 414 (1972), 25: 244
(1973), and 28: 446 & 452. 1974; Alain in Leon & Alain, Fl. Cuba
imp. 2, 2: 318. 1974; Fosberg, Rhodora 78: 113. 1976; Mold.,
Phytologia 34: 280. 1976; Hsiao, Fl. Taiwan 4: 434. 1978; Mold.,
Phytol. Mem. 2: 25, 54, 91, 99, 104, 197, 198, 254, 266, 280,
Poe Zoos, foe, 302, 304, 309, 319, 321, "36/7, 459;5°R 592. 1908;
Mold., Phytologia 48: 489 & 494--500, fig. 1--5. 1981.

Illustrations: Viertel, Trees Shrubs Vines no. 406 [as V.
negundo incisa]. 1970; Mold., Phytologia 48: 495 & 497--500, fig.
1--5. 1981.

Venkatareddi (1969), citing his nos. 97634 & 97860, asserts
that this variety "Grows often in association with the typical
variety", flowering and fruiting "All the year". Clarke (1885),
in his discussion of what he calls "var. incisa'", comments that
"The extreme pinnatifid form of this var. is Chinese; the wild
Indian examples are crenate-serrate, i.e. intermediate." It is
not clear if he is here referring to var. intermedia or, more
likely, to var. cannabifolia (Sieb. & Zucc.) Hand.-Mazz. The il-
lustration given by Viertel (1970) is labeled "Vitex negundo
incisa", but plainly depicts, not that variety, but var. inter-
media.

Recent collectors describe V. negundo var. intermedia as a
spreading deciduous shrub, 1--4 m. tall, with fragrant blossoms.
They have found it growing on rocky ground, in dry land on wooded
hillsides, along roadsides, "between houses", in mixed woods,
and (in Texas) in sandy soil in oak or Populus-Vitex communi-
ties. They have encountered it at 1000 m. altitude, flowering
in May and from July to December. Hu refers to it as "common"
in Hong Kong. The Wroten collection, cited below, does not have
any statement on its accompanying label that it represents culti-
vated material, but I am assuming, from the locality of collection,
that it does.

The corollas are said to have been "pink" on Tsang 27843,
"purple" on Chevalier 12, and "blue & blue-violet, with a yellow
spot in the mouth" on Cutler 5043.

Material of this variety has been widely misidentified and
distributed in herbaria as V. agnus-castus L., typical V. negundo
L., V. negundo var. cannabifolia (Sieb. & Zucc.) Hand.-Mazz., and
V. negundo var. incisa (Lam.) C. B. Clarke.

Additional citations: TEXAS: Collingsworth Co.: L. C. Higgins
6263 (N). Donley Co.: L. C. Higgins 3910 (N). Harris Co.: Tra-
verse 1322 (Au--179257). INDIA: East Punjab: Koelz 8278 (N).
Karnataka: G. Thomson s.n. [Maisor] (Pd). Uttar Pradesh: Kapoor
& Thamman 27194 (Mu). CHINA: Anhwei: Chow 7 (Ac). Fukien: Ging
5388 (Ws), 5956 (Ws). Kiangsu+: Chiao 22343 (Ws). Kwangsi: Tsang
27843 (Ca--1286197): wan « Chow 79016 (Ld). CHINESE COASTAL IS-
LANDS: Hainan: piang 63036 (Mu). HONG KONG: yu 6858 (W--2711970),
9243 (W--2711719). JAPAN: Honshu: Maximowicz s.n. [Yokohama,
1862] (Pd). TAIWAN: Oldham 383 (Pd). PHILIPPINE ISLANDS: Luzon:
Rothdauscher s.n. [Manilla, 1879] (Mu--1523). CULTIVATED: Cayman
Islands: N. Chevalier 12 (N). Cuba: Cutler 5043 (Ba). Louisiana:

3

180 POE XY yl Ody Onl DE A Vol. 49, No. 2

Wroten C.423 (Ne--33949). Oklahoma: FE. M. Alexander 35 (Tu--
129545). Pakistan: Iqbal s.n. [7-9-1957] (Kh). Virginia: #H. A.
Allard 11390 (Se--134450).

VITEX NEGUNDO var. LAXIPANICULATA P'ei

Additional bibliography: Mold., Phytologia 8: 67. 1961;
Mold., Fifth Summ. 1: 291 (1971) and 2: 927. 1971; Mold., Phytol.
Mem. 2: 280 & 592. 1980.

VITEX NEGUNDO var. MICROPHYLLA Hand.-Mazz.

Additional bibliography: Mold., Phytologia 15: 311. 1967;
Mold... bat th,Summ. 12 291 (1971) ‘and..2:.. 927 .-497ts Mold... Phytol.
Mem. 2: 280 & 592. 1980.

VITEX NEGUNDO var. PHILIPPINENSIS Mold., Phytologia 38: 308. 1978.

Synonymy: Vitex negundo var. philippensis Mold., Biol. Abstr.
65% 6/769; 1978:

Bibliography: Mold., Biol. Abstr. 65: 6769. 1978; Mold., Phy-
tolapia 3382 308. 1978; Hocking ,) Excerpt. Bot....4.33:, 66. 1979;
Mold... Phytol. Mem. +25. 309%, 3193367, 459,, & 592. (1980: Mold...
Phytologia 48: 490. 1981.

Collectors have found this plant in flower in January, March,
May, July, and November and in fruit in March. The Loher 4433
specimen in the Munich herbarium is a mixture with something non-
verbenaceous.

Most of the material cited below was originally distributed
and previously cited by me as typical V. negundo L.

Citations: PHILIPPINE ISLANDS: Luzon: Elmer 8125 (N--type);
Loher 4433 in part (Mu--3966, Z); E. D. Merrill 1503 (N), 1636
(N), 2320 (N), 3429 (N), Sp. Blanc. 440 (Gg--31493 ,N); Roth-
dauscher s.n. [Manilla, 1879] (Mu--1522). GREATER SUNDA ISLANDS:
Java: Teijsmann 16728 (N). CULTIVATED: Java: Herb. Hort. Bogor.
XV.J.A.XXXIV.6 (Bz--26411, Bz--26412, Bz, Bz, Bz, Bz, N); Koorders
42133b (Bz--24462, Bz--25663, N).

VITEX NEGUNDO f£. PURPURASCENS Sivarajan & Mold. in Mold., Phyto-
logia 28: 404. 1974.

Bibliography: Mold., Phytologia 28: 404 & 445. 1974; Hocking,
PExcerpe,. DOL. A. 25: 379. 1975; Mold... Phytol. Mem. 2; 266, 367, &
592. 1980; Mold., Phytologia 48: 490. 1981.

Collectors describe this plant as a shrub or small tree, the
leaves "densely deep purplish tomentose beneath when alive" (Si-
varajan 1327), "purplish beneath" (Fortune 25), or with "underside
of leaves violet" (Corner s.n.). Corner also describes the "in-
florescence and flowers violet, corolla with a white spot in the
throat". The leaflets are all 3 per leaf on some sheets of For-
tune 25 and either 3 or 5 on others, in all cases remarkably small
in size. Corner reports the vernacular name,. "kemuning hitam",
in Malaya, and reports that the form is cultivated "elsewhere in
Malaya", too.

Citations: INDIA: Kerala: shivarajan s-n- [Calicut] (Ld); Si-
varajan 1327 (Ld,Uc),; CU.1849 (Z--type). HONG KONG: Fortune 25

1981 Moldenke, Notes on Vitex 181

(E--2168600, Mu--650). CULTIVATED: Malaya: Corner s.n. [Jan.
97a] (Ld).

VITEX NEGUNDO var. SESSILIS Mold.

Additional bibliography: Mold., Phytologia 15: 311. 1967;
Mold... Fifth Summ. 1: 374 (1971) and 2: 927. 1971; Mold., Phytol.
Mem. 2: 367 & 592. 1980.

VITEX NEGUNDO var. TRIFOLIOLATA Mold., Phytologia 25: 432. 1973.

Bibliography: Mold., Biol. Abstr. 56: 3000. 1973; Mold., Phy-
Edlogia 25% 432. 1973.

Recent collectors refer to this plant as a shrub or small tree
"common" in open places especially along streams, the leaves green
above and white beneath, and have found it in flower in January
and July. The corollas are said to have been "purple" on the
Ramamoorthy collection and "blue" on that of Saldanha, cited below.

Material of this variety has in the past been distributed in
herbaria as typical V. negundo L.

Citations: PAKISTAN: Baluchistan: K. H. Rechinger 29984 (W--
2637733--type). INDIA: Karnataka: Ramamoorthy HFP.374 (W--
2794868); Saldanha 12470 (W--2794869).

VITEX NEGUNDO var. VARIEGATA L. H. & E. Z. Bailey, Hortus Third
1162 [as "Cv. 'Variegata']. 1976; Mold., Phytol. Mem. 2:
459. 41930.

Synonymy:Vitex negundo cv. 'Variegata' L. H. & E. Z. Bailey,
Hortus Third 1162. 1976.

Unfortunately the Baileys fail to provide a description for
this variety, but obviously the name is intended to apply to the
form of the species with its leaflets variegated with white or
yellow blotches [foliolis albo- vel luteo-variegatis].

VITEX NEO-CALEDONICA Gandoger

Additional bibliography: Mold., Phytologia 15: 311--312. 1967;
Mold., Fifth Summ. 1::343 (19710 ‘and 2: 724, 7/64) & 927. 1980:
Mold., Phytol. Mem. 22.332, & 592. 2960,

VITEX NLONAKENSIS Engl.

Additional bibliography: Mold., Phytologia 15: 312. 1967;
Mold., Fifth Summ. 1: 224 (1971) and 2: 927. 1971: Mold., Phytol.
Boe. 25205 & 592. 1980.

[to be continued]

NOTES ON NEW AND NOTEWORTHY PLANTS. CXLIX

Harold N. Moldenke

LANTANA GLANDULOSISSIMA £. PARVIFOLIA Mold., f. nov.

Haec forma a forma typica speciei laminis foliorum plerumque .
2--4 cm. longis 1.8--2.6 cm. latis recedit.

This form differs from the typical form of the species in hav-
ing its mature leaf-blades mostly only 2--4 cm. long and 1.8--2.6
cm. wide.

The type of the form was collected by J. Rzedowski (no. 7933)
in cactus-mesquite matorral 28 km. south of San Luis Potosi, San
Luis Potosi, Mexico, along the central road to Mexico City, at
1900 m. altitude, on August 3, 1956, and is deposited in the
Herbario Nacional of the Instituto de Biologia in Mexico City.

LANTANA URTICOIDES f. ACULEATA Mold., f. nov.

Haec forma a forma typica speciei ramulis argute aculeatis re-
cedit.

This form differs from the typical form of the species in hav-
ing its stems, branches, and branchlets conspicuously thorny-
aculeate.

The type of the form was collected by L. S. Smith and Y. Co-
rona (Mex.28) a few km. north of Tehuacdn, Puebla, Mexico, at an
altitude of about 6000 feet, on August: 4, 1966, and is deposited
in the Herbario Nacional, Instituto de Biologia, Mexico City.

LANTANA VELUTINA f. FLAVA Mold., f. nov.

Haec forma a forma typica speciei corollis flavis recedit.

This form differs from the typical form of the species in hav-
ing yellow corollas.

The type of the form was collected by Robert Bye (no. 7753)
at the base of Buenos Aires falls in the short-tree forest on the
north side of a barranca on the La Bufa to Quiraro road, Munici-
pality of Batopilas, Chihuahua, Mexico, at 1100 m. altitude, on
July 30, 1977, and is deposited in the Herbario Nacional, Insti-
tuto de Biologia, Mexico City. The collector describes the plant
as a small shrub, about 1 m. tall, with recurved spines on the
twigs [not evident on the type specimen], the leaves sticky, and
the flowers [corollas] yellow.

VITEX VESTITA £. UNIFOLIOLATA Mold., f. nov.

Haec forma a forma typica speciei foliis ut videtur unifolio-
latis recedit.

This form differs from the typical form of the species in ap-
parently having its leaves only unifoliolate, the leaflet appar-
ently usually borne at a considerable angle with the petiole.

The type of this form was collected by W. J. J. O. de Wilde and
B. E. E. de Wilde-Duyfjes (no. 13415 in a montane rainforest at
Camp. 2, ascent of Gunung Bandahara, Gunung Leuser Nature Reserve,
Atjeh, Sumatra, June 27, 1972, deposited in the U.S.National Her-

barium, Washington.
182

BOOK REVIEWS

Alma L. Moldenke

"NEPTUNE'S GIFT - A History of Common Salt" by Robert P. Multhauf,
xix & 325 pp., 37 b/w fig. , 9 maps, 36 tab, & 99 photo.
Johns Hopkins University Press, London & Baltimore, Maryland
21218. 1979 Second printing. $22.50.

Most botanical students, technicians, professors and aficio-
nados are aware of (cell theory fame with Schwann) Schleiden's
"Die Rose" 1873, but maybe not about his "Das Salz" 1874 where "we
see laid out the evidences of salt in history, literature, com-
merce, folklore, science and technology". Part I of Multhaus'
book covers this earlier Age of Culinary Salt with sources for
this universal necessity and primitive methods of production from
sea brine and rock salts in various countries. Part II deals with
the Era of Chemical Salt with exploitation of mother liquors, salt
geology, the Kali and petroleum industry rise, dye and chlorine.
Appendix I gives statistics on Salt Production and II on Produc-
tion of Artificial Soda, Sodium Hydroxide and Chlorinated Hydro-
carbons. The illustrations are of great interest historically as
is the text for reading. Of course, for many kinds of chemists
this is a needed book.

"BIBLE PLANTS AT KEW" by F. Nigel Hepper, 64 pp., 26 color
photo., 11 draw., 1 map & 1 fig. Her Majesty's Stationery
Office, London EC1IP 1BN. 1981. £2.95 net paperbound.

A few decades ago when strolling through the immaculately

kept Royal Botanic Gardens at Kew, I noticed how very many plants
scattered about were ones that were mentioned in the Bible under
one name or another. In this regard it must be remembered that
the King James Version (1611) predated Linnaeus' start of binom-
ial nomenclature by a century and a half. The map of these gar-
dens, the descriptive text, the beautifully clear color photo-
graphs, all help with identifying Biblical plants.

"WORLD ENERGY: THE FACTS AND THE FUTURE" by Don Hedley, 368 pp.,
154 b/w tab., 32 charts. Facts on File, Inc., New York, N.
7: 30019... 29822622. 30.

The author states "that the most pressing problem [facing man-
kind now] is to maintain the world's energy supply.....The ‘energy
crisis’ represents a threat to progress and to standards of living
in rich and poor countries alike which is inevitable and imminent.
Our fuels....will run out" soon. With clearly explanatory and
reliable text, effective and clearcut charts and tables the author

183

184 Peet Oo Cue Len Vol. 49, No. 2

"forecasts how the world energy economy will have changed by the
year 2000 and what is likely to happen beyond....internationally,
nationally, commercially and personally."

"LEWIS AND CLARK'S AMERICA - A Voyage of Discovery" by Willis F.
Woods, Director Seattle Art Museum, Bicentennial Illustra-
ted Catalogue, 96 pp., 13 color prints, 76 b/w prints, 3
maps.

"LEWIS AND CLARK'S AMERICA - A Contemporary Photo Essay", photo-
graphy by Paul Macapia, journal and sketches by Mary E.
Macapia. Seattle Art Museum, 96 pp., 58 b/w photos, 5 maps
& 4 draw. University of Washington Press, Seattle, Washing-
ton 98105. 1976. Both $12.50 paperbound & slip-covered.

These books make a lovely combination; each is attractive in
its own right. The paintings of American and European artists
made after the expedition are beautiful, accurate and very well
printed for the scenery, the wildlife, the Amerinds and some of
their artifacts. The small expedition left on May 14, 1804,
reached the Pacific coast and returned to St. Louis on September
23, 1806. "Lewis was a born naturalist. He saw and described
178 plants.....117 are present today in the Lewis and Clark
Herbarium at the Academy of Natural Sciences in Philadelphia. The
vertebrate animals....totaled 122 species and subspecies."

Woods' introduction also mentions the artists. Macapia's beauti-
ful photographs, carefully avoiding Coca Cola signs, Russian
thistle growing and airplanes flying, are arranged along the ex-
pedition's route as indicated by his wife's journal and sectional
maps.

"YOUR BIBLICAL GARDEN - Plants of the Bible and How to Grow Them"
by Allan A. Swenson, xxii & 217 pp., 8 color pl., 24 b/w
photo. pl. & 25 draw. Doubleday & Co. , Garden City, New York,
or New York, New York 10017. 1981. $13.95.

"Many of the most beautiful , more striking plants of the scrip-
tures are available to you for your outdoor home gardens or for
growing indoors as house plants and in containers on porch, balcony
or rooftop in the city. Here, for easy reference, are the names of
the plants that you will find in this book, the fruits and flowers,
herbs, vegetables and trees.'"' Alternate choices, similar species
or forms, growing directions and items of special interest comprise
the easily read text which is enriched by excellent photographic
prints. The drawings, like that of the pomegranate, offer little
embellishment.

UU

85 00288 2676

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