0-? ms LABIEMAE GP MIgHiGAN

Thesis {or .3113 :‘ggrg’e. pf} My 5
MICHIGAN STAKE WWERSETY

Ann Herrick Waterman
1959

 

"L “5395

L I B R A R Y
Michigan State
University

mom AND POW GRAIN HORPHOLOGY 03' THE
LLBIATAI OII' MICHIGAN

by

Ann He rri ck Wat e man

AN ABSTRACT

thitted to the College of Science and Arte

Michigan State University of Agriculture and.

Applied Science in partial fulfillment of the
requimente for the degree of

“SEER or same:
Department of Botany and Plant Pathology
1959

Mill”

ABSTRACT

The taxonow and pollen grain morphology of the Michigan species of
the Labiatae have been investigated.

Approximately 3.1460 specimens at 15 herbaria have been examined. A
key to 29 Michigan genera is provided. and a species treatment of the 61
recognized species contains keys. discussions of geographic distributions.
habitats. and flowering periods. 01' the 61 Michigan species. 20 have
been introduced from the Old Iorld. 1+ are cosm0politan in temperate
regions and the remaining 37 are native in various portions in North
America. A comparative analysis has been made of the species of Lab-
iatae in Michigan with those in Wisconsin. Illinois. and Indiana. A
check list for each county in Hichigau gives the names of the species
collected in that county. Species distributions have been indicated by
symbols on outline maps of Ilichigan.

Pollen grain studies have been made of 52 species of the Labiatae
occurring naturally in Michigan and of 3 cultivated species. rho char-
acteristics of pollen grain morphology are discussed and are also sum-
marized in a table. For a number of species. a determination has been
made of nuclear content of pollen grains at anthesis by a technique
developed in this study. Two major morphological groups of grains have
been recognized: those that are binucleate and tricolpate and those
that are trinucleate and hexacolpate. This observation confirms a
previous report of this condition. Variation in size and increase in
number of colpi within individual plant specimens have been observed in
5 naturally occurring species. Comparisons have been made of untreated
plants with a cultivar which was irradiated or treated with colchicine.

Pollen grains with more than 6 colpi also have been found to be tri-

nucleate. A.key to 48 naturally occurring species and 2 cultivated
species (based on acetylated pollen grains) has been constructed.
Photomicrographs of pollen grain nuclei of lb species and acetylated

grains of #1 species are included.

TAXONOMY AND POLIM GRAIN MORPHOLOGY OF THE
LABIATAI OF MICHIGAN

by

Ann Herrick Waterman

A THESIS
Submitted to the College of Science and Arts

Michigan State University of Agriculture and
Applied Science in partial fulfillment of the

requirasents for the degree of

MASTER OF SCIENCE
Department of Botam and Plant Pathology

1959

TABLE OF CONTENTS

.Acknowledgements. . . . . . . . . . . . .
List of Distribution Maps. . . . . . . . .
Introduction. . . . . . . . . . . . . . .
Taxonomy. . . . . . . . . . . . . . . .
Historical Aspects . . . . . . . . . .
Comments on.Distribution and Habitats.

Material Studied . . . . . . . . . . .

nomenclature . . . . . . . . . . . . . .

Key to Michigan Genera . . . . . . . .

Treatment of Species . . . . . . . . .

Check List of Species Recorded in.iach.nichigan County . .

Pollen Grain Studies. . . . . . . . ......

Material and.Methods . . . . . . . . .

Diacmsion O O C O O O O O O O O O O O C O I O O O O O 0

Key to Species or Groups Based on Pollen Grain Characteristics

Table I. Pollen Grain Characteristics

Table II. Documentation of Pollen Grain Studies . . . . . .

Platefi.xrnoo................o...

References. . . . . . . . . . . . . . . . . . . . .

ii

11

55
55

63

7h

78
9h

1
ACKNWLEDGRMENTS

The taxonomic advice. critical suggestions. and inspiration given by
Dr. John B; Seaman. my manor professor. have made possible this investi-
gation. He also made arrangements for me to study with Dr. llso
Barghoorn at Harvard.University. I am indebted to Dr. Barghoorn for his
assistance with the photomicrography. for provision of equipment and for
working space in his laboratory and for the priviledge of examining the
Harvard.microscope slide collection of acetylated pollen grains.

Helpful assistance has been given me by the curators of herbaria where
collections of Michigan specimens of Iabiatae were studied. The herb-
aria are listed at the beginning of the taxonomic treatment.

To Dr. D. G. Voss of the University of Michigan. I am especially
grateful for allowing me to examine his recently collected specimens and
the Isle Royals National Park Herbarium (which.was temporarily at.Ann
[Arbor in connection with the Michigan.rlora Project). He also helped me
obtain the outline maps of Michigan from the Zoology Department of the
university of Michigan.Museum. and gave advice on various technical
matters. A A

I should like to thank Dr. llizabeth.McClintock of the California
Academy of Sciences for her notes on Michigan Physostegia.

Dr. Ray Nelson of the Department of Botany and Plant Pathology of
Michigan State University has kindly let me examine plants which were
irradiated with cobalt-60 or treated with colchicine.

I wish to thank my graduate committee members. Drs. G.‘l. Prescott and

Leo M. Mericle. for their professional criticism.

Map no.

H00 mVOxU't 3U) NH

Stat—-

I-‘H
«PU

HH
mm

8555

21.

23

25
26

30
32
33

35
36

11

List of Distribution Maps

Teucrium canadense

Scutellaria parvula. S. leonardi. S. elliptica var.
hirsute

S. lateriflora

S.galericulata................... .

Marrubium vulgare. Dracocephalum parviflorum

Agastache nepetoides

A. soropmlariaefolia
Mepetacataria.....................

Glechoma hederacea

Prunella vulgaris

Physostegia virginiana var. virginiana. P. virginiana
var. speciosa. P. aboriginorum

Leonurus cardiaca. L. marmbiastrum. . . . . . . . . . .

Galespsis tetrahit. Stachys mssopifolia

Lamium amplexicaule. L. purpureum. L. maculatum
Stachys palustris var. palustris. S. palustris var.
homotricha

s. m.p1d8e O O O O O 0 O O O O O O O O O O O O O O O O

Stachys tenuifolia

Monarda didym. M. punctata var. villicaulis

M. fistulosa
Blephiliaciliata...................

Blephilia hirsuta

Hedeoma pulegio ides

H. hispida

Acinos arvensis. Satureja hortensis. . . . . . . . . .

Calamintha glabella var. angustifolia. Collinsonia
canadensis

Clincpodium vulgere

Thymus sermllum. Hyssozns officinalis

Pycnanthemum flexuosum. P. verticillatum, P. pilosum. .

Pycnanthemum virginianum

chopus uniflorus ‘

L. rubellus
L.aSper.L.V1rginlcuS.................

chOpus americanus

Mentha piperita

M. spicata. M. gentilis

M.arvensissubsp.canadensis. . . . . .. . . . . . ..

Page

11!

20

21+

32

35

38

1

INTBOCUCTIOH

This investigation has developed out of an initial interest in pollen
grain morphology in the Labiatae. Pollen grains of species of this fam-
ily in the New World have received until now practically no attention from
botanists. In the initial phases of this investigation it was considered
advisable to examine a wide range of species in the family. To this end
the Michigan species were chosen as a base. Since the Iebiatae in
Michigan have received no over-all taxonomic investigation more recently
than M. J. Beal's (19010 flora of Michigan. it was necessary that the
taxonomic studies be carried out in conjunction with those on pollen
grains.

In making available the results of the combined taxonomic and pollen
grain studies it is heped that contributions have been made to several
diverse fields. Certain taxonomic problems in the Michigan species of
this family have been clarified. and much information on the geographic
distribution of these species within the state has been brought together.
Comparative studies of pollen grain morphology and cytology have produced
data that can be applied to the understanding of the taxa above the rank
of genus and on the specific and infraspecific level also. Paleobotanical
investigations have until the present done little with pollen grains of
the Iebiatae. but information from studies such as this one may aid in
subsequent investigations of various fossil deposits. In Michigan. as
new other areas of the United States. the honey industry is important;
knowledge of these pollen grains should be useful to that industry.

Several botanical problems still await investigation in the Labiatae

of Michigan. Slightly more than one-fourihof the naturally occurring

2

species are known from five or fewer collections. Additional field work
would make available more specimens of the little hown species and also
provide data on distributions and habitats. Improvement in the taxonomic
understanding of these species would then be possible. Purther studies
of pollen grain morpholog should be made of additional species. and
comparisons of plants within and between populations of the same species
are needed. Additional techniques for preparation of pollen grains
should be employed such as those used or discussed by Wodehouse (1935).
Paegri (1956). and Ikuse (1956). Time has not been available in the
present study to permit examination of chromosomes. Such studies would
be helpful in explaining several peculiarities that have been encountered
in the pollen grain work. They probably would be helpful also in clari-
fying certain taxonomic problems. Experimental approaches along several
lines are in order. although these might more profitably be applied on a

broader geographic basis to generic or smaller subgroups of the family.

TAXOMOM!
Hiltoncal Asmctg.

The first reports on the Iebiatae in Michigan were by John wright
(1839) who included 23 species from the two southernmost tiers'of counties
in the state. Cooley's (181419) Catalogue listed two more species 'in the
primitive region south of Lake Superior". and Coleman (187% enumerated
42 species from the lower Peninsula. Beal's (1901+) flora of Michigan has
been for many years the most detailed listing of Michigan species of
Iaabiatae. Since that publication. however. 12 more species in the mint
family have been added to the Michigan flora.

Ploristic studies in the Douglas Lake region (Voss. 1956 and unpublished

check list). Kalamazoo Co. (flames and flames. 191W). and Isle Royals

3
(Brown. 1937) provide complete lists of species now known from these

areas. The studies in Berrien Co. (Billington. 1925). Gogebic Co.
(Darlington. 1921. 1923). Oakland Co. (Bingham. 19115. and Washtenaw Co.
(Walpole. 1921+: Hermann. 1936a. 1937) are of interest but do not have
full listings of species of those particular areas. 0. A. Farwell's
reports of species of the Labiatae in papers from 1896 to 194.5 (of. McVaugh
et a1.. 1953) have been confirmed in most but not all cases. Of the
numerous county check lists published by C. X. Dodge between 1900 and
1921. only the posthumous paper (1921) (including the eastern and north-
ern parts of the Lower Peninsula along Lake Huron. and Chippewa. School-
craft. and Tuscola Counties) has been examined in this study. Unfor-
tunately herbarium specimens have not been found to verify his species

reports.

Comments on Distribution and hbitgtg.
In Driquet's (l895) world floristic treatment of the Labiatae. nine

subfamilies are delimited: three of these are represented by species in
Michigan. About one-third (20) of the 61 species in Michigan have been
introduced from the Old Morld and four are cosmopolitan in temperate
regions. The remaining species have more or less restricted distributions
in North America.

Although certain areas of Michigan are still poorly collected (especial-
1y Alcona. Baraga. Clare. Gladwin. Isabella. Lapeer. Mecosta. Ogemaw.
Osceola. Tuscola and Mexford Counties. each of which has only one to four
presently-known species) general patterns of geographic distribution can
be noted from the maps. More species occur in the lower Peninsula than
in the Upper Peninsula (59 species have been collected in the Lower

Peninsula and 33 in the Upper Peninsula). Only two species (W

it

fogniculum and Mpg; genezengig) of the total state flora occur in the
Upper Peninsula but not in the lower Peninsula.

On the basis of the present study 29 genera and 61 species of the
Labiatae are recognized in Michigan. Sufficiently recent detailed studies
of three contiguous states are available to permit comparisons between
these areas and Michigan (nomenclatural differences in these studies have
been taken into account). Koeppen (1957) accounted for 29 genera and
59 species in Misconsin. Of these. 2? genera and 148 species are comon
to both states. while two genera and 13 species are found in Michigan but
not in Wisconsin. and two genera and 11 species are found in Wisconsin
but not in Michigan. Jones and Puller (1955) treated 30 genera and 76
species in Illinois of which 26 genera and 52 species are comon to both
states. Three genera and nine species are found in Michigan but not in
Illinois and four genera and 214 species are found in Illinois but not in
Michigan. Deam (19110) listed 28 genera and 63 species in Indiana of which
214 genera and '46 species are comon to both states. rive genera and 15
species are found in Michigan but not in Indiana and four genera and 17
species are found in Indiana but not in Michigan.

Habitats of the Michigan species range from xeric sandy areas to mesic
Open woods. pastures. roedsides and lawns. river and stream banks. flood-
plains. and damp lake shores. None of the species are aquatic although

several species may be found with roots of the plants submerged.

to i S ed.

The taxonomic aspects of this study are based primarily on an examina-
tion of approximately 3.160 herbarium specimens. A month of field work
(July. 1958) was spent in counties known to (be poorly collected. This

field trip has helped augment the distributional data for a number of

5
species. to provide living material for pollen grain studies. and to con-
tribute to the knowledge about the living plants and their environment.

Only non-cultivated plants or escapes from cultivation are considered
in the taxonomic treatment although some cultivated species are consid-
ered in the pollen grain studies.

When specimans are cited in the taxonomic treatment and in the pollen
grain studies. the herbarium abbreviations are those listed by LanJouw
and Stafleu (1959). Specimens in the herbaria listed below have been
examined:

Albion College. Albion

Aquinas College. Grand hpids

Bailey Hortorium. Ithaca. New York

Chicago Natural History Museum. Chicago. Illinois

Cranbrook Institute of Science. Bloomfield Hills

Gray Herbarium of Harvard University. Cambridge. Massachusetts

Isle Royals National Park Herbarium. Houghton

Michigan State University. East Lansing

Missouri Botanical Garden. St. Louis. Missouri

National Museum. Smithsonian Institution. Washington. D. C.

University Herbarium. University of Michigan. Ann Arbor

University of Michigan Biological Station Herbarium. Douglas
Lake. Cheboygan

Wayne State University. Detroit

Western Michigan University. Kalamazoo

Wiegand Herbarium of Cornell University. Ithaca. New York

The symbols on the distribution maps show actual localities with the
exception of spen circles which indicate county records only. The latter
are from herbarium specimens with insufficient locality data. In certain
instances on these maps it should be noted that the known species distri-

butions illustrate much collecting activity in several local areas.

kmenclaturg.
Where possible the nomenclature employed here follows that of the most

recent comprehensive treatments such as monographs. revisions. or chromo-

somal studies. These are cited where pertinent in the text. In taxa

6
for which no intensive studies are available. the nomenclature of Fernald
(1950) or Gleason (1952) is employed. The keys to genera and species are
mostly adapted from Fernald although.in several cases aid in key construc-

tion has been obtained from the following papers: DeWolf (fiatureja.

glinOpgdium. Calgginthaq and.Agiggg. T95h). Bpling (§cutellaria, l9h2).
Epling and Stewart (m 1939). Grant and Eplins (Lcn_an_.t__hemm.__. 1943).
Hermann (lam. 1936b). Lewis (w. 1915). Lint and Ipling
(Agastaghg. 19h5). and.McClintodk and.lpling (Teucrium. l9h6).

no attempt has been made to list full synonymy. The synonyms that
are listed in the species treatment are names used by Fernald (1950) but
not employed in this study. The sequence of genera and species follows

Fernald's arrangement.

0.

CC.

G.

7
KEY TO GENERA

Ovary shallowly h—lobed; style not gynobasic; nutlet scar lateral
and large. . . . . . . . . . . . . . . . . . . . . . . . . . . B
Ovary deeply h—lobed; style gynobasic; nutlet scar basal and
small. . . . . . . . . . . . . . . . . . . . . . . . . . . . . D
B. Lobes of corolla nearly regular; flowers on axillary l- to 3-
flowsred pedunclss. . . . . . . . . . . . . l. Trichgstema
BB. Lobes of corolla irregular. lower lip 2 times as lange as
upper; flowers in close terminal verticils or spiciform
racemes. . . . . . . . . . . . . . . . . . . . . . . . . C
flowers in close terminal verticils; corolla with very short upper
lip. not cleft. blue; bracts obovate. leaflike. . . . . 2. 5,1252
flowers in spdciform racemes: corolla deeply cleft between the two
small lobes of upper lip. dark pink to white; bracts linear.
acuminate and not leaflike. . . . . . . . . . . . 3. Teugrium
D. Calyx with distinct protuberance on upper side. h. §ggtellaria
DD. Calyx without distinct protuberance on.upper side. . . . . . B
Calyx teeth 10. spiny pointed. spreading at maturity; plants
white tomentose. . . . . . . . . . . . . . . . . . . 5. Marrubigg
Calyx teeth.5. not spiny pointed nor spreading at maturity; plants
not white tomentose. . . . . . . . . . . . . . . . . . . . . . F
P. ‘Antherifsrous stamens 4. all with 2-locu1ar anthers: upper lip
of corolla galeate or concave. . . . . . . . . . . . . . . . G
FF. Antheriferous stamens 2 or b. upper pair shorter; upper lip of
corolla not galeate nor concave. . . . . . . . . . . . . . . P
Upper (posterior) stamens longer than lower (anterior). . . . . . B

Upper stamens shorter than the lower. . . '. . . . . . . . . . . . K

I.

II.

I.

8
H. Anther sacs parallel; stamens exceeding corolla. 6. Agastache
EH. Anther sacs strongly divergent; stamens not exceeding the
cmmun... ... ... ... ... ... ... ... .. I
Calyx distinctly Z-lipped. upper tooth much broader than the others.
notched between some of the sinuses; flowers blue. .9. Egaggcephglum
Calyx regular or oblique. . . . . . . . . . . . . . . . . . . . J
J. Stem erect; flowers many in each whorl. forming a terminal
inflorescence.white; leaves ovate to ovate—oblong; anther

locules merely divergent. . . . . . . . . . . . . . 7. 1222112

JJ. Stem weak. trailing; flowers 2-6 in each whorl. in the exile of

leaves. purplishpblue; leaves rounded to rsniform; anthers of
perfect flowers with locules at angles. each pair forming a
cross. . . . . . . . . . . . . . . . . . . . . . . 8. filgghgg§_
Calyx deeply bilabiate. lO-nerved. reticulately veined. closed in
fruit. upper lip flat and 3-toothed; pollen grains hexacolpate. .
10. Mile
Calyx not strongly bilabiate. obscurely 5-10 nerved. Open in fruit.
teeth subequal; pollen grains tricolpate. . . . . . . . . . . . L
L. Flowers Opposite; calyx obscurely nerved. membranaceous,
globose or enlarged after flowering. . . . . . ll. Physgstegig
LL. Flowers in compact spikes. whorls or panicles; calyx 5-nerved.
firm. not enlarged nor inflated after flowering. . . . . . M
Caulins leaves pahnately veined. deeply cleft or incised; calyx
teeth divergent; whitish upper lip of corolla densely woolly. . .
12. Leonurus
Cauline leavespinnately veined (except m) and crenate. dentate.

or serrate; calyx teeth ascending; upper lip of corolla glabrous or

00.

9
pilose. . . . . . . . ... . . . . . . . . . . . . . . . . . . . N
N. Anther sacs without Atrichomss; flowers interruptedly spicate.
the upper or all bracteal leaves reduced; upper lip of corolla
concave.......................l5.§_t_g:m
NN. Anther sacs with trichomes; flowers in exile of well developed
leaves; upper lip of corolla galeate. . . . . . . . . . . . O
O. Trichomes within the anther sacs; calyx lobes sharply spinose—
tipped....................... l3. filespgis
OO. Trichomes on outer surface of anther sacs; calyx lobes not sharply
I spinose-tipped....................lh.m
P. Antheriferous stamens 2. ascending and parallel; anther sacs
linear; corolla strongly 2-lipped. . . . . . . . . . . . . Q,
PP. Antheriferous stamens 2 or it; anther sacs ovoid; corolla
slightlyZ-lipped..................... S
Q. Anthers with lower locales deformed; flowers mostly interruptedly
spicate. with upper bracteal leaves reduced. . . . .16. alv
QQ. Anthers with both locules fertile; flowers verticillate. capitate.
withleafyouterbracts.................... B
B. Calyx equally 5-toothed. 15-nerved. . . . . . . . 17. Menard;
RR. Calyx 2—11pped. l3-nerved, upper lip with three awned teeth.

18. 16 11111
S. Corolla z-lipped. the upper lip erect. entire. or shallowly

nOtChed. (except EICMthQM)s s s o s s s s o s s s s o o s o T
SS. Corolla not bilabiate. less equally ll-lobed (upper lip of

Qollinsoniafringsd)...................... a

T. Fertile stamens 2; calyx gibbous on lower side. . l9. Hgdeoga

TT. Fertile stamens lb; calyx not gibbous (except Mm). . . U

 

 

 

 

5'1:
8t:

1'11

bb

lO
Calyx lS-nerved. naked in throat; inflorescence a dense spike . .
2a. m

Calyx 10 to l3-nerved. villous or naked in throat; inflorescence not
a dense spike. . . . . . . . . . . . . . . . . . . . . . . . . V
V. Flowers in dense or loose axillary and terminal cymes or verti-

cils; filaments converging. ascending parallel or curved. . W
W. Flowers in loose racemes or dense corymbs; filaments diverging.

notcurved........................ W
Calyx campanulate. lo-nerved. not bilabiate. . . . . 20. Saturejg
Calyx tubular. l3-nervsd and bilabiate. . . . . . . . . . . . . X
X. Calyxgibbous...................21. M1
xx. Calyxnotgibbous..................... Y
Calyx straight. bearded at throat; bracts of inflorescence leaf-
like.......................w..22. Calamintg
Calyx curved. slightly bearded at throat; bracts of inflorescence
'setaceous-hirsuts. . .1. . . . . . . . . . . . . . . 23. glinopodig
2. Calyx bilabiate. densely villous in throat; flowers in loose to

dense racemes; corolla pirple. . . . . . . . . . . . 25. m
22.. Calyx not bilabiate. naked in throat; flowers in dense corymbs;

corolla white with purple spots. . . . . . . . 26. ficnantm

Flowers solitary in loose teminally branched racemes. yellow.
stamensz......................29. W
l'lowers crowded in dense whorls or spicate; corolla whitish to pink.-
pirpleandbluish; stamensZorh. . . . . . . . . . . . . . . . b
b- Stamens 2; flowers whitish. not with mint fragrance. 27. ms

bb. Stamens h; flowers pinkish-purple to white; with mint fragrance.

28. Mengg

11

Treatment of Species

l. Trichostemg L. False Pennyroyal; Bastard Pennyroyal.
Reference: Lewis. 191:5.
a. Calyx nearly actinomorphic; stamens barely exssrted. nearly
straight.....................l. 1.22%
ea. Calyx zygomorphic; stamens long sxsertsd and strongly arched. . .
2- L W
1. .1. pragatum L. No map.
Known in Michigan from only two collections: Alpena Co.. Thunder Bay

Island. Whgglgr mu (flowering) 18 July 1895 (GE. MSC. US); and
Chippewa Co.. Drumond Island. shallow soil over limestone pavement.
w m1,(flowering)29 August 1958 ('01)).

These northern specimens are like the small depauperats plants de-
scribed as Iganthug brachiatgg var. linearig Iassett (1933). and Lewis
states. "These may represent a northern ecotype. but this can only be

determined by further investigation.”

2. 1. W L. Bastard Pennyroyal. No map.

Known in Michigan from four collections: Kalamazoo Co.. Texas Twp”
22.93 mu (unspened buds) 1“ August 1951 (WMU); Texas Twp.. mg; 5619,.
31 August 1950 (WMU); Oshtemo Twp.. gaps; 1232. 19 September 1939 (WMU);
Oakland Co.. Drayton Plains. Open grassy field. Chandler 5.3.. 15 Sept-
ember 1916 (B111. MICH. WUD). There are no habitat data with the Kalamazoo
Co. specimens. however. Hanes and Hanes (1947) report that their Oshtemo

Twp. collection is from sterile sandy fields bordering oak woods.

2. 51:35; L. Bugleweed.
a. Plants stoloniferous; leaves glabrous. . . . . . 1. A. reptw

12

aa. Plants not stoloniferous; leaves pubescent. . . 2. A. ggnevgnsig

1- i- r4329. 33- No nap-
Known in Michigan from only two collections: Berrien Co.. Brandywine

Acres. bridge over St. Joseph River. flgflert l+222. (flowering) 15 May
1951+ (MICH); Ionia Co.. Ionia. weed in lawn. m 3,3... (flowering)

22 May 1953 (MSC). Introduced from the Old World.

2. A. genevensig L. No map.

Known in Michigan from only one collection: Marquette Co.. Huron Mt.
Club. Lake Superior dunes. m g.n.. (no date) (BLH). Introduced

from the Old World.
3. 132.31g L. Germander; Wood Sage.

Reference: McClintock and Epling. 19146.
1. Teucrium ganadensg L. Amarican Germander. Map l.

Scattered throughout the state but not frequently collected in the
Upper Peninsula. In moist sand. clay. and swampy soils in cpen and
shaded areas. l'lowsring from June to September.

At the time of Beal's (1901!) Flora this species was known only from

the Lower Peninsula.

h. §mellgia L. Skullcap.

Reference: Bpling. 19152.
a. Flowers in terminal or axillary racemes. . . . . . . . . . . . . b
an. Flowers solitary in exile of ordinary foliage leaves. . . . . . c
b. l'lowsrs in terminal racemes; cauline leaves deltoid—ovate.

margins crenate; corolla galea plus tube lit-21 mm. long. . .

13

1. §. elliptica var. hirsutg

bb. Flowers in lateral racemes; caulins leaves ovate. margins

crenate-serrate; corolla galea plus tube 5-7 mm. long. . .
2. s. alanine
c. Galea and tube of corolla 114-21 mm. long; leaves cordate at base.
2.5-5 cm. long; stem 2-6 dm. high. . . . . . . 3. §_. galericulata
d. Basal hairs of leaves sglandular. leaves entire. rsvolute. .

4. §,. leonar

dd. Basal hairs of leaves glandular. leaves irregularly sub—

dentate....................5. immla

1. i. glliptica Muhl. var. hirsute (Short) Fern. Hairy Skullcap. Map 2.
The northern extent of the range of this variety is Kalamazoo. Case
and St. Joseph Counties. leing reports the variety (as subspecies) from
Berrien Co.. but no Berrien collection has been seen in this study. The

only recorded habitats are oak woods. Flowering from July to August.

2. §_. Lateriflog L. Mad-dag Skullcap. Map 3.
Abundant throughout the state. In swampy and wet sandy soils on lake
shores and river and stream banks. Flowering from May to September.
Occasionally white—flowered forms of this species are collected; var.
glbiflora Farwell is one of these. The type of this variety. not located

by McVaugh et a1. (1953). is at MICH. Farwell 3285.

3. i. galericulatg L. Common or Marsh Skullcap. Map 11'.
S. epilobiifolia A. Hamilton
A Eurasian and North American species. abundant throughout the state.
In the same habitats as S. lateriflorg but frequently flowering earlier.

Flowering from May to August.

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15
b. §. leonardi Epling. Smooth Skullcap. Map 2.
§. mrvula Michx. var. 1gonarg; (Bpling) Fern.

Known in Michigan from only two collections from Ingham 00.: East
Lansing. Grand Trunk RR east of Okemos crossing. Walpgle 1218 (fruits
only) 10 June 1922 (AIBC. BLH. use); and pilllggton M” (same locality
as the first collection). dry gravelly bank. (with fruits only) 9 July

1923 (31-3. WU. MUD).

5. _S_. mailichx. Small Skullcap. Map 2.
A widely distributed but poorly collected species in Michigan. In

dry limestone soils and along railroads. Flowering from June to July.
5. Marrubigg L. Horehound.

l. Mgrubium M13 L. Common Horshound. Map 5.
Introduced from the Old World. sometimes escaping from cultivation

and becoming a weed. Flowering from June to August. and in November.

6. Agastachg Clayt. Giant Hyssop.

Reference: Lint and Epling. 19145.
a. Calyx teeth ovate-obtuse. lobes at anthesis l-l.5 mm. long; corolla
yellowish................... lO. Quagtoides
aa. Calyx teeth deltoid-acute. lobes at anthesis 2-2.5 mm. long;
corolla rose to purple or blue. . . . . . . . . . . . . . . . b
b. Leaves and stems generally conspicuously pubescent; calyx teeth
glabrous; corolla rose to purple. 2. A. agrophulariaefolia
bb. Leaves and stems essentially glabrous. whitish beneath; calyx

teeth pubescent; corolla blue. . . . 3. A. foeniculpm

      

               
       

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17
l. A. nepgtoideg (L.) ths. Yellow Giant HyssOp; Hedge Hyssop. Map 7.
Found in the southern part of the Lower Peninsula. In open woods

and meadows. Flowering from July to September.

2. A. gcrophulariaefolia (Willd.) the. Purple Giant Hyssop. Map 7.
Distribution in Michigan same as A. nepgtoideg. Occasionally culti-

vated as an ornamental. Flowering from August to September.

3. A. fgsniculum (Pursh) the. No map.

Known from only one collection in Michigan: Houghton Co.. prairie.
19m}; 10323. 15 August 1931: (BLH). (Farwell. 1938).

7. ML. Catmint.

l. m gagarigI L. Catnip. Map 8.
Introduced from the Old World and established throughout the state

except on Isle Hoy'ale. In rich. Mp to dry soils. on roadsides. in

pastures and near buildings. Flowering from June to October.
8. Glechoma L. Ground Ivy.

1. Qlechog hederacea L. Gill-over—the-ground. Map 9.
Including var. gicrantha Moricand and var. pagiflora (Benth.) Farwell.
Introduced from the Old World and frequent throughout the state. In
rich. damp to dry soils. on roadsides. in lawns. cemeteries. and pastures.

Flowering from May to July.
9. Dracocephalum L. Dragonhsad; Lion's Heart.

1. Dragocephalum miflorum Nutt. American Dragonhead. Map 5.

Scattered throughout the state. In sandy. Open areas. Flowering

18

from June to August.

10. well; L. Heal-all; Selfheal; Carpenter-weed.

l. flagella Elgaris L. Map 10.
Including var. lanceolata (Bart.) Fern.

 

Abundant and widespread in Michigan. In wooded areas and as a weed
along roadsides and in pastures and lawns. Flowering from June to Octo-

ber.
11. Egysostegia Benth. False Dragonhead.

3. Leaf margin undulate. firm. . . . . . . . . 1. E. gboriginorum
aa. Leaf margin serrate. membranaceous. . . . . . . . . . . . . . . b
b. Cauline leaves subtending infloresence greatly reduced in size.
1.5-3.5 cm. long. 0.2-0.6 cm. wide; mid-cauline leaves 6.5-9.0
cm. long. l-2.5 cm. wide; corolla 1.5-2 cm. long. . . . . . .
2. 2. virginiana var. virginiana
bb. Cauline leaves subtending infloresence not greatly reduced.
5—ll.5 cm. long. 0.6-2.5 cm. wide; mid-cauline leaves 9-15 cm.
long. 1.6-9.0 cm. wide; corolla 1.6-2.0 cm. long (in cultivated

specimens 2.8 cm. long). . . . . .3. 2. virginiana var. spgcigsa

lo 20 gbgrginoggg Fern. Map 110
Known in Michigan from one collection'from Wayne Co.. Detroit. waste

ground. "probably an escape from cultivation“. Farwell 5.9.911 (determined
by him as Dracocemalum denticulatum Ait. and 2. virginianum var.
dgnticulatm Ait.). 11+ July 1918 (em. on).

This collection was annotated by Elizabeth McClintock in 191+? as _D_.

virginianpg. These plants. however. do not fit the description in

19
Gleason's (1952) treatment which is based on Dr. McClintock's findings;
the uppermost leaves are longer than the internodes which they subtend.

the calyx is 0.75 cm. long. and the corolla is 2.3 cm. long.

2. 2. virginiana (L.) Benth. var. virginiana Map 11.

2. parviflogg Nutt.

Known from a few collections scattered throughout the state; some—
times cultivated. On river banks and lake shores. Flowering from July
to September.

One specimen. Farwsll 621. July 1888 (BLH). has uppermost leaves
which are longer than the internodes they subtend. The other specimens

examined coincide with the character as stated in the key (lead b) above.

3. 3. W (L.) Benth. var. gmcigsa (Sweet) Gray. Map 11.
Scattered throughout the state; occasionally cultivated. In low

areas on damp. rich soil. l'lowsring from July to October.
12. Leonurus L. Motherwort.

a. Lower two calyx lobes strongly ref lexed; cauline leaves palmately
lobed to subentire or 3-cleft; corolla pale purple. tube bearded
within.......................‘l. L.cardiaca

ea. Lower calyx lobes not strongly reflexed; cauline leaves coarsely

toothed; corolla whitish. tube naked within. . 2. L. parrubiastrum

1. L. cardiaca L. Common Motherwort. Map 12.
Naturalized from the Old World. and abundant throughout the state.

In mucky to sandy soils. Flowering from June to October.

2. L. marrubiastrum L. Map 12.

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21
Introduced from the Old World. and known in Michigan from only one
collection: Ingham Co.. floodplain of Red Cedar River. garmelee 1221.

(flowering) 15 September 1953 (M80).
13. Galeopsis L. Hemp-Nettle

l. leeopsis tetra_hi_g L. Map 13.

Including var. b_1_f_i_d_§_ (Boenn.) LeJ. 8: Court. and forma albaFarwell
(Farwell. 1916).

Naturalized from the Old World; most of the Michigan records are from
the northern part of the state. In marshes. Open woods. lake shores.

and roadsides. Flowering from July to September.
114. @ium L. Dead Nettle.

a. Corolla 18-30 mm. long; perennials with creeping stems bearing
offshoots....................1. Laculatum
aa. Corolla 8-18 mm. long; winter or summer annuals. without creeping
stems. . . ... . . . . . . . . . . . . . . . . . . . . . . . . . b
b. Upper leaves sessile and clasping. reniform; corolla tube
naked within. . . . . . . . . . . . . . . . 2. L. amplexicaule
bb. Upper leaves petioled. cordate-ovate; corolla tube with a ring

ofhairswithin..............-3o 9w

1. L. Qaculatum L. Variegated Dead Nettle. Map 114.
Introduced from Maps and escaping from cultivation. scattered in the

southern part of the Lower Peninsula. Flowering from May to August.

2. _L_. mlexicaule L. Henbit. Map 1'14.

Naturalized from Europe; known from the Lower Peninsula only. In

22
cultivated fields. gardens. moist woods. and roadsides. Flowering from

April to June.

3. L. mgreum L. Purple Dead Nettle. Map 14.
Naturalized from EurOpe; probably common but poorly collected in
Michigan. On sandy lake shores. woods. and gardens. Flowering from

April to June.

15. Stage L. Hedge Nettle.

a. Upperstemspubescentonsides. . . . . . . . . . . . . . . . . . b
aa.Upperstemsglabrousonsides. . . . . . . . . . . . . . . . . . c
b. Hairs on angles longer than hairs on the sides of stem; leaves

not densely pilose. . . . . . . . 1. §_. Elustris var. Elustris

bb. Hairs of nearly equal length on both angles and sides of stem:
leaves densely pilose. . . . . . 2. fl. mlustris var. homotricg
c. I Calyx lobes hirsute or hirsute-ciliate; leaves more or less
pubescentabove. ....... . ... ....3. §.hispida
cc. Calyx lobes entire; leaves glabrous above. . . . . . . . . . . . d
d.. Principal leaves on petioles 2—3 cm. long. leaves elliptic-
lanceolata. . . . . . . . . . . . . . ll. §. tenuifolia

dd. Principal leaves sessile. oblong-lanceolate. 5. §. hissomfolia

1. §. mlustris L. var. pglustris. Map 15.
Introduced from Maps and occurring throughout the state except on

Isle Royals. On barrier beaches. damp roadside ditches. and in pastures.

Flowering from July to September.

2. s. Elustrig var. homotricha Fern. Map 15.

Known in Michigan from Cheboygan. Emet. and Keweenaw Counties. On

23

damp. sandy lake shores. Flowering from July to August.

3. 5,. hispidg Pursh. Map 16.
S. tenuifolia Villd. var. mug (Plush) Fern.
Reference: Gleason. 1952.
Scattered throughout the Lower Peninsula. and in three counties in the
Upper Peninsula. On wooded river banks. lake shores. and in swamps.

Flowering from June to September.

1+. 5. Mlfllld. Hedge Nettle. Map 1?.
Known only in the southern half of the Lower Peninsula. On moist
ground in wooded areas and on floodplains. Flowering from July to

September.

5. i. msgopifolig Michx. Map 13.
Primarily an Atlantic Coastal Plain species. found in Michigan in the

southwestern counties of the Lower Peninsula. In abandoned fields. on

lake shores. and in marshes. Flowering from June to August.

16. filvia L. Sage.

a. Plants perennial; calyx slightly bilabiate; corolla strongly exssrt—
ed from the calyx. . . . . . . . . . . . . 1. _S_. azurea var. grandiflora
aa. Plants annual; calyx strongly bilabiate; corolla slightly exserted

from the 08131. e o s e s s e e s s s e e e e s s 2. £0 reflexa

1. _S_. gzureg Michx. gt Lam. var. grandiflora Benth. No map.
Known in Michigan from only two collections: Kalamazoo Co.. North of

Vicksburg. mg m gh2§, (flowering) 15 September 193“ (WMU); Kent

Co.. Grand Rapids. waste place. probably introduced from the Southwest.

      
    

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Cole 5,3,. (flowering) September 1898 (MSC).

2. §_. reflexa Hornem. No map.
Introduced from the western United States and known from Kalamazoo,

Monroe. and Wayne Counties. Flowering from August to September.
17. Monarda L. Horsemint; Wild Bergamont.

a. Flowers in axillary and terminal glomerules; corolla creamy-yellow
with purple spots. . . . . . . . . 1. fl. pu_nctat§ var. villicaulis
aa. Flowers in terminal solitary glomerules; corolla red. purple or
pink.............................b
b. Cauline leaves deltoid-ovate; corolla pink or rarely white.
2. fl. fistulosa
bb. Cauline leaves dsltoid—lanceolate; corolla red-purple. . . c
c. Dracts vividly red; corolla vermillion to scarlet; calyx without
dense crown of stiffly divergent bristles. . . . . 3. L4,. $13121;
cc. Bracts dark purple; corolla deep red-purple; calyx with a dense

crown of stiffly divergent bristles. . . . . . . . a, 91. media

1. 21. mctata L. var. zillicagig (Pennell) Shinners. Horsemint. Map 18.
Reference: Shinners. 1953.
Videly collected in the western part of the Lower Peninsula; known

also from southeastern Michigan and from one station in Chippewa Co. On

sand dunes and in dry pastures. Flowering from July to September.

2. M. {istulosg L. Wild Bergamont. Map 19.
Occurring throughout the state except on Isle Royals. In darnp and
dry muck to light soils. in open woods and grassy areas. Flowering from

July to September.

26
3. )1. m L. Oswego Tea; Bee-Balm. Map 18.
Introduced from the eastern United States. and scattered in the south-
ern part of the Lower Peninsula. Along roadsides and in cemeteries.

Flowering from July to August.

'4. 93,. media Willd. No map.
Known in Michigan only from St. Clair Go. from C. K. Dodge collections

(at MSG). Flowering from July to August.

a. Lobss of lower calyx lip extending beyond the sinuses of the upper
calyx lip; cauline leaves sub-sessile to sessile. . 1. 1;. 11 1:
aa. Lobss of lower calyx lip not extending to the sinuses of the upper

lip; cauline leaf petioles 1-2 cm. long. . . . . . 2. 13, hirsuta

l. h. giliatg (L.) Benth. Downy Blephilia. Map 21.
Occasional in the southern half of the Lower Peninsula. In dry open

woods and prairies, on light sandy soil. Flowering from June to August.

2. a. hirsuta (Pursh) Benth. Woodmint; Hairy Blephilia. Map 21.
Fairly common in the southern part of the Lower Peninsula and known

also from one collection in Ontonagon Co. (Darlipgton in 1923. MSC). In

swamps. beech-maple woods. floodplains and river banks. Flowering from

June to September.
19. Hgdsgma Pursh. Mock Pennyroyal.

Reference: Epling and Stewart. 1939.
a. Cauline leaves petiolate. lanceolata and serrate; calyx lobes entire;

sterile stamens obvious. . . . . . .' . . . . . l. g. pglegioides

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aa. Cauline leaves sessile. linear and entire; calyx lobes ciliate;

sterile stamens degenerate. . . . . . . . . . . . 2. g. hispida

l. g. pulegioideg (L.) Pers. American Pennyroyal. Map 22.

Occasional in the Lower Peninsula. Epling and Stewart report that
the range of this species extends through Dickinson. Menominee. and
Delta Counties in the Upper Peninsula. but no collections from this area
have been seen in the present study. In dry sandy soils. in pastures.

swamps. and open woods. Flowering from June to September.

2. E. gapida Pursh. Rough Pennyroyal; Hock Pennyroyal. Map 23.
Scattered throughout the Lower Peninsula and known from one collection

in Dickinson Co. (Igrnald m. GB, MICE) in the Upper Peninsula. Epli’ng

and Stewart reported this species from the southern lower Peninsula only.

In sandy loam fields and on sandy roadsides. Flowering from June to

AuglISts
20. Satureja L. Savory.

1. §atureja hortensig L. Summer Savory. Map 214.
Introduced from EurOpe. cultivated in gardens and rarely an escape.

Flowering from August to September.

’\

21. Aging; Mill. Mother-of-thyme.
Reference: Defiolf. 1951+.
1. MM mensig (Lam) Dandy. Map 214.
Naturalized from EurOpe and found in both the Upper and Lower
Peninsulas; most collected at the northern tip of the Lower Peninsula
and adjacent Upper Peninsula. On calcareous and sandy soils. Flowering

from July to September.

a ' ,’/_1'1"1|
\ .
-” ” / u «summonses '

   

.. ‘ ‘ _ ‘l " '-. ‘
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ll}? 00 ‘ H 7 b3 u \ ' ' 'r ,) . ‘ . . z - r
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30

22. Calamintg Mill. Calamint.

References: DeWolf. 1951+. 1955.
1. W W (141::th Benth. var. wstifolig (max-r.) newolf.
94‘? 25.
Satureja arkansana (Ruth) Briq.
Frequent in the northern counties bordering Lakes Michigan and l

Huron. On wet limestone and sandy shores. Flowering from July to

August .

23- Wk Basil.

Reference: Deflo lf . 1951».

l. Cliggpgdium My; L. wild Basil. Map 26.
w E5232! (1..) Fritsch var. ngogaea Fern.

A circumboreal species found throughout most of the state in
calcareous and alluvial soils. On lake shores. in deciduous and coni-

ferous woods. and in meadows. Flowering from May to September.
21h m 1» Hyssop-

1. stsomg gfgiginalig 1.. Map 27.

An escape from herb gardens. known from collections in five counties.

Flowering from July to September.
25. MB, L0 Thyme.

1. 1m; gemllum 1.. Wild Thyme: Creeping Thyme. Map 2?.

Introduced from the Old World and established in areas scattered over
the state: planted by the State Highway Dept. on sandy road banks in the

central part of the state. Flowering from Juen to September.

31
26. Wmchx. Mountain Mint.

Reference: Grant and Fpling. 19%.
a. Calyx teeth l—l.5’mm. long. lanceolate to subulate. the tips sharp-
pointed......................l. Lglexuosum
aa. Calyx teeth 0.7-1.0 mm. long. deltoid. pubescent. the tips not
sharp-pointed......................... b
b. Sides of stem glabrous. angles pubescent; leaves sessile. 6-11
mm.broad.................l. LW
bb. Sides of upper half of stem pubescent: cauline leaves short-
petioled.lo-18mm.broad................. c
c. Lower surface of leaves pubescent chiefly on veins: heads pedicillate.
a. 2. magnum
cc. lower surface of leaves pubescent over most of the surface; heads

nearlysessile.................. l4%. 2.2ilosum

l. 3. {leggmm Walt.) DSP. Narrow—leaved Mountain Mint. Map 28.
Scattered in the Lower Peninsula: its northern range in Michigan was
previously reported by Lint and Fpling as Allegan (erroneously printed
Allegany) and Wayne Counties. Recent collections extend its range to
hmet (1991 m. MICE) and Leelanau (Ihgmmon 1:33.59 BIB) Counties. On

roadsides. golf links. and old fields. Flowering from June to August.

2. 2. zirginigum (L.) Durand 8: Jackson. Mountain Mint. Map 29.
Abundant in the southern half of the Lower Peninsula: known also from
Leelanan and Menominee Counties. In dry. sandy areas. Open woods and

marshes. Flowering from July to September.

3. 2. zerticillatum (Michx.) Pers. Map 28.

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33
Collected on Saginaw Day in Huron Co.. and along the southeastern
side of Lake Michigan. In Open. sandy areas. Flowering from August to

September .

u. 2. 1112M Butt. Map 28.
Known in Michigan from only three collections: Berrien Co.. Niles.

shaded bank of St. Joseph River. Edge 53g, 25 August 1917 (MICK);
Monroe Co.. In Salle. upland roadsides and fields. 11119}; m. (flower-

ing) 26 July 1922 (GB. MICK); Mashtenaw Co.. five miles NW of Dexter.

Beggrg églz. (flowering) 8 August 19148. (MUD).
27. m L. Mater Horehound.

Reference: Hermann. 1936b.
a. Calyx teeth shorter or equalling nutlets. calyx lobes broadly
triangularwithoutmidnerve................... b
as. Calyx teeth surpassing nutlets. lobes narrowly triangular with
prominentmidnerve....................... c
b. Leaf margins concave from lowest tooth to stem; main stem
usually tuberous; usually large robust plants; tsp of nutlets
with numerous. acute tubercles. nutlets when mature conceal-
ingthecslyces. . . . . . . . . . . . . .1. lazirginig!
bb. Leaf margins usually convex to straight (infrequently concave):
main stem usually tuberous: plants not large and robust; top
of nutlets with 3 to 5 rounded tubercules. nutlets when nature

not concealing the calyces. . . . . . . . . . 2. L. mil-2!!!

c. Leavessessile...................3.19m

cc. Leases petiolate. . . . . . . . . . . . . . . . . . . . . . . . . d

d. Corolla surpassing calyx lobes by 2-3 mm.; leaves shallowly

34
serrate; ridge of nutlets not entire. verrucose to tuber—
culate...................l&. Labellus
dd. Corolla surpassing calyx lobes by 1 mm. or less; leaves

irregularly cleft or pinnatifid; ridge of mitlets entire. .

5. L. gericanus

l. L. zirginicgg L. Map '32.
.Known in Michigan from one collection: Ingham Co.. Meridian Twp..
Red Cedar floodplain forest. Parmelee 39.53.- (mature nutlets only) 20

September 1952 (M86). This species is easily confused with L. gn__ifloru .

2. L. uniflorus Michx. Map 30.
Abundant throughout the state. In wet areas. Flowering from July to
September. The tuberous stolons of this species are infrequently collect-

. ed.

3. L. m Greene. Map 32
Local along the southern margins of Lakes Huron and Michigan and also

from Cheboygan Co. On sandy or swampy lake shores. occasionally with

the roots submerged. Flowering from July to September.

it. L. Menu! Moench. Mater Horshound. Map 31.
Scattered throughout the state. but not known from Isle Royals. In
swamps. marshes. and on wooded river banks. Flowering from July to

September.

5. L. geriganus Muhl. Hap 33.
Including var. loggii Banner and var. ggabrifolig Fern.
Midely distributed and the most comon species of this genus in

Michigan. On sandy or swampy lake shores. river banks. and in roadside

I I
I" ,4 \ y ,-
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----‘ *’ , 31. moms RUBELLUS . L. vmslmcus +
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36
ditches. Flowering from July to August.

28. Mentha L. Mint.

a. flowers in crowded terminal spikes or in terminal interrupted
spikem........................... b
a. Flowers in axils of leaves in subglobose clusters. . . . . . d
b. Spikes interrupted; leaves petioled. . . l. 31. piperita
bb. Spikes densely crowded. slender and leafless; leaves sessile
or nearly so. . . . . . . . . . . . . . . . . . . . . . c
c. Plants soft-pubescent; leaves round-ovate to broadly elliptical.
2. a. new
cc. Plants essentially glabrous: leaves oblong or ovate-lanceolata.
3. !. ate.
d. Stem glabrous or with few scattered hairs; calyx glabrous or
upper half pubescent. . . . . . . . . . . . it. 31. gentilig
dd. Stem retorse-pubescent on angles: calyx pubescent. . . . . .

5. 11. mengig subsp. canadensis

l. E. 11.212129; L. Peppermint. Map 314.
rhe non-cultivated specimens of this species have sterile anthers.
Introduced from Europe and considered by some to be a lubrid between
'5. m L. and 91. sgcata L. (cf. Olapham. Tutin. and Warburg, 1952).
Scattered throughout the state. Along swampy roadsides and borders of

cultivated fields. l‘lowering from July to September.

2. £1. gotundifolia (L.) Suds. Apple-scented Mint. No map.
Introduced from Europe and known in Michigan from only three col-

lections: Kalamazoo Co.. Schoolcraft Twp.. Portage Greek. damp ground.

37
Rapp .9. Hang; 21466. 21 August 1936 (WU); Kent Co.. Alpine Twp.. 901; m”
8 October 1899 (AQC); Grand Rapids. Ogle m" 1899 (M50).

3. E. M L. Spearmint. Map 35.

All material examined of this species had only sterile anthers. Intro-
duced from Europe and scattered throughout the Lower Peninsula. 0n
beaches, damp roadsides. and along rivers. Flowering from August to

September.

h- E- m 1‘- Map 35.
Introduced from Europe and thought to be a hybrid between M. 223339.11

and fl. spicata (cf. Clapham. Tutin. and Mai-burg. 1952). Scattered in

the Lower Peninsula. In moist or wet areas. flowering from July to

August.

5. 21. arvgggig L. subsp. gaggdensis (L.) Kara. Mild Mint. Map 36.
Reference: Hare. 1956.

H. mm; var. M Fern. and var. 11.11212 (Benth.) S. 3.
Stewart.

This is a circumboreal species; the range of subsp. My};
extends from Alaska to Iabrador, south to Calif ornia. Arizona. and
Virginia. It differs from the European subsp. mensis and the Asian
subsp. pimrascgng in chromosome number. leaf and calyx-lobe shape. and
has pulegone instead of menthol.

In Michigan specimens there is variation in amount of stem and leaf
mbesence.

Comon throughout the state. Along lake shores and river banks and

in marshes. Flowering from June to August.

3.3 .. .
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39
29. Qollinsonia L. Horse Balm.

1. mm L. Stone-root; Rich—weed. Map 25.
fairly common in the southern part of the Lower Peninsula. In

woodlands. flowering from July to October.

Check List of Species Recorded in Each Michigan County

ALCORA COUNT!

Olinopodium vulgare
Draoocephalum parviflorum
Lchpus americanus
Scutellaria.galericu1ata

ALGEB COUNTY

Blephilia ciliata

Olinopodium vulgare

Galeopsis tetrahit

Lycopus americanus

L. rubellus

L. uniflorus

Mentha arvensis subsp.
canadensis

Prunella vulgaris

Scutellaria galericulata

S. lateriflora

ALLEGAN COUNTY

Agastache nepetoides
Blephilia hirsuta
ClinOpodium vulgare
"Glechoma hederacea
Iemium maculatum
Mentha arvensis subsp.
canadensis
M. piperita
M. spicata
Monarda.fistulosa
M. punctata var.
villicaulis
Physostegia virginiana var.
speciosa
Prunella vulgaris
Pycnanthemum flexuosum
P. virginianum
Scutellaria lateriflora
Stachys hyssOpifolia
Teucrium canadense

ALPBRA COUNTY

Oalamintha glabella var.
angustifolia

Clinopodium vulgare
Leonurus cardiaca
Lycopus americanus
Mentha arvensis subsp.
canadensis
Trichostema brachiatum

ANTRII COUNTY

Oalamintha glabella var.
angustifolia
Clinopodium vulgare
Lycopus americanus
Prunella vulgaris
Scutellaria galericulata

AREEAC COUNTY

Olinopodium vulgare
Lycopus americanus
Pycnanthemum virginianum
Stachys hispida
Teucrium canadense

BARAGA COUNTY

Olinopodium tulgare
Galeopsis tetrahit
Prunella vulgaris

BARRY COUNTY

Glechoma hederacea

Leonurus cardiaca

Lchpus uniflorus

Mentha arvensis subsp.
canadensis

M. spicata

Monarda fistulosa

M. punctata var. villicaulis

Nepeta cataria

Prunella vulgaris

Scutellaria lateriflora

Stachys hyssOpifolia

BAY COUNTY

Clincpodium vulgare

Hedeoma pulegioides

Lycopus americanus

Mentha arvensis subsp.
canadensis

M. spicata

Physostegia virginiana var .

virginiana
Pycnanthemum virginianum

3111213 COUNTY

Clinepodium vulgare

Leonurus cardiaca

Lycopus americanus

Mentha arvensis subsp.
canadensis

Monarda punctata var.
villicaulis

Repeta cataria

Scutellaria.ga1ericulata

313mm! comm _y

.Agastache nepetoides

AJuga reptans

Blephilia hirsita

Clinopodium vulgare

Glschoma hederacea

Hedeoma pulegioides

Lycoyus americanus

L. asper

L. rubellus

Marrubium vulgare

Mentha arvensis subsp.
canadensis

M. gsntilis

M. piperita

M. spicata

Monarda f istulosa

M. punctata var.
villicaulis

Nepeta cataria

Physostegia virginiana var.

speciosa

1+1

Prunella vulgaris
Pycnanthemum pi lo sum

P. virginianum

Satureja hortensis
Scutellaria galericulata
S. lateriflora

S.
Stachys hispida

S. hyssopifolia

S. tenuifolia
Teucrium canadense

BRANCH COUNTY

Agastache nepetoides
Collinsonia canadensis
Hedeoma pulegioides
Lycopus rubellus

L. uniflorus

Monarda f istulosa
Pycnanthemum virginianum
Scutellaria galericulata
Stachys temiifolia

CALHOUN COUNTY

Blephilia ciliata

ClinOpodium‘vulgare

Collinsonia canadensis

Glechoma hederacea

Leonurus cardiaca

Lycopus americanus

L. uniflorus

Mentha arvensis subsp.
canadensis

M. gentilis

M. piperita

M. spicata

Monarda fistulosa

M. punctata var. villicaulis

Nepeta cataria

Prunella vulgaris

Pycnanthemum virginianum

Scutellaria galericulata

S. lateriflora

Stachys hyssOpifolia

Teucrium canadense

1/ The nine species of the Labiatae noted in Billington's (1925) study

of Warren Moods have been seen.

CASS COUNTY

Agastache nepetoides

A. scroplmlariaefolia

Blephilia hirsuta

Clinopodium vulgare

Collinsonia canadensis

Hedeoma.pulegioides

Leonurus cardiaca

Lycopus americanus

L. rubellus

Le muloma

Mentha arvensis subsp.
canadensis

M. piperita

M. spicata

Monarda fistulosa

Repeta cataria _

Physostegia virginiana var.
virginiana

Prunella vulgaris

Pycnanthsmum virginianum

Scutellaria elliptica var.
hirsuta

S. galericulata

Se lateriflora.

Stachys hyssOpifolia

CEARLEVOIX COUNTY

Acinos arvensis
Calamintha glabella var.
- angustifolia
Clincpodium vulgare
Leonurus cardiaca
Lchpus americanus
Mentha arvensis subsp.
canadensis
Prunella vulgaris
Scutellaria galericulata
Teucrium canadense

CHARLEVOIX COUNTY: GARDEN ISLAND

Acinos arvensis

Calamintha glabella var.
angustifolia

_2/

b2

Leonurus cardiaca
Nepeta cataria

OHARLEVOIX COUNTY: BEAVER ISLAND
Calamintha glabella var.
angustifolia
Clincpodium vulgare
Lamium purpureum
Lycopus americanus
Mentha arvensis subsp.
canadensis
Monarda fistulosa
Nepeta cataria
Prunella vulgaris
Scutellaria lateriflora

cmomm comm _z/

Acinos arvensis

Calamintha glabella var.
angustifolia

Clinopodium vulgare

Galeopsis tetrahit

Glechoma hederacea

Hedeoma hispida

Hedeoma pulegioides

Lamium maculatum

Leonurus cardiaca

Lycopus americanus

L. asper

L. uniflorus

Marrubium vulgare

Mentha arvensis subsp.
canadensis

M. gentilis

M. piperita

M. spicata

Monarda fistulosa

Nepeta cataria

Prunella vulgaris

Pycnanthemum flexuosum

Scutellaria galericulata

S. lateriflora

Stachys palustris var. palustris

S. palustris var. homotricha

Teucrium canadense

In the history of the floristics in Douglas Lake Region. Voss (1956)

has given an account of the rejected records.

CHIPPEMA COUNTY

Acinos arvensis

Calamintha glabella var.
angustifolia

Clinopodium vulgare

GaleOpsis tetrahit

Glechoma hederacea

Lycopus americanus

Le uniflom

Mentha arvensis subsp.
canadensis

M. piperita

Monarda fistulosa

M. punctata var. villicaulis

Nepeta cataria

Prunella vulgaris

Scutellaria galericulata

S. lateriflora

Se parvula.

Stachys palustris var. palustris

Trichostema brachiatum

CLARE COUNTY

Scutellaria lateriflora
Thymus sermllum

CLINTON COUNTY

Blephilia ciliata

Clinopodium vulgare

Leonurus cardiaca

Lycopus americanus

L. uniflorus ‘

Mentha arvensis subsp.
canadensis

M. piperita

Monarda fistulosa

Prunella vulgaris

Pycnanthemum.virginianum

Scutellaria galericulata

5. lateriflora

CRAWFORD COUNTY

Draco cephalum parviflorum
Lchpus americanus

_3/ See note: Cheboygan.County

“3

L. uniflorus

Mentha arvensis subsp.
canadensis

M. gentilis

Monarda fistulosa

M. punctata var. villicaulis

Prunella vulgaris

Scutellaria galericulata

S. lateriflora

DELTA COUNTY

Calamintha glabella var.
angustifolia

Clinopodium vulgare

Lchpus americanus

L. uniflorus

Prunella vulgaris

Scutellaria galericulata

S. lateriflora

Stachys hispida

Teucrium canadense

DICKINSON COUNTY

Dracocephalum parviflorum

Hedeoma hispida

Lycopus americanus

Mentha arvensis subsp.
canadensis

Monarda fistulosa

Prunella vulgaris

Stachys hispida

EATON COUNTY

Blephilia hirsuta ,
Glinopodium vulgare
Collinsonia canadensis
Glechoma hederacea
Prunella vulgaris
Pycnanthemum virginianum
Teucrium canadense

mm comgl

Acinos arvensis

Oalamintha glabella var.
augustifolia

Olinopodium vulgare

Galeopsis tetrahit

Glechoma hederacea

Hedsoma hispida

H. pulegioides

Leonurus cardiaca

Lycopus americanus

L. uniflorus

Mentha arvensis subsp.
canadensis

M. piperita

M. spicata

Monarda fistulosa

M. punctata var. villicaulis

Prunella vulgaris

Scutellaria galericulata

S. lateriflora -

Stachys palustris var. palustris

S. palustris var. homotricha

Thymus serpyllum

GENISRR COUNTY

Blephilia hirsuta
Collinsonia canadensis
Galeopsis tetrahit
Lycopus americanus
Mentha piperita
Monarda.fistulosa
Prunella‘vulgaris
Scutellaria galericulata
S. lateriflora

Stachys hispida

S. tenuifolia
Teucrium canadense

GLADHIN COUNTY

lepeta cataria
Scutellaria galericulata

comm comm _n/

ClinOpodium vulgare

an

Dracocephalum parviflorum

Galeopsis tetrahit

Lycopus uniflorus

Mentha arvensis subsp.
canadensis

Prunella vulgaris

Satureja hortensis

Scutellaria galericulata

S. lateriflora

Stachys palustris var. palustris

GNEND TRAVERSE COUNTY

Clinopodium vulgare
Leonurus cardiaca
Lycopus americanus
Mentha arvensis subsp.
canadensis
Menarda fistulosa
Nepeta cataria
Physostegia virginiana var.
virginiana
Prunella.vulgaris
Teucrium canadense

GRATIOT COUNTY

Agastache scraphulariaefolia

Blephilia hirsuta

Clinopodium vulgare

Glechoma hederacea

Lycopus americanus

L. rubs llus

Leonurus cardiaca

Mentha arvensis subsp.
canadensis

M. piperita

M. spicata

Monarda fistulosa

Nepeta cataria

Physostegia virginiana var.
speciosa

Prunella vulgaris

Scutellaria galericulata

S. lateriflora

Stachys hispida

Tsucrium canadense

_5/ Burlington (1921) reports fileghoma hedgragga. Teucrium canadense.

and (1923) Lyggpgg,amgricang§.

These specimens are not in the Michigan

State‘University Heriarium but undoubtedly occur in this county.

«- VIM—...“. .——.—r— ._ ‘_ _

N COUNTY

Lopodium'vulgare
nopus officinalis
turns cardiaca
'pus amerioanus
Lniflorus

rubium vulgare

:ha arvensis subsp.
.nadensis

lanthanum verticillatum
rirginianul

sellaria galericulata

Lateriflora

 

 

 

 

 

 

HILLSDALE COUNTY

Agastache nepetoides
Collinsonia canadensis
Glechoma hederacea
Monarda fistulosa
Teucrium canadense

ROUGRTON COUNTY

Agastache foeniculum

Clinopodium vulgare

Dracocephalum parviflorum

GaleOpsis tetrahit

Glechoma hederacea

Leonurus cardiaca

Lycopus americanus

L. uniflorus

Mentha arvensis var.
canadensis

Monarda fistulosa

Nepeta cataria

Physostegia virginiana var.
virginiana

P. virginiana var. speciosa

Prunella vulgaris

Scutellaria galericulata

S. lateriflora

Stachys palustris var. palustris

Thymus sergyllum

INGRAM COUNTY

Agastache scraphulariaefolia

Blephilia hirsuta

Collinsonia canadensis

Dracocephalum parviflorum

Glechoma hederacea

Hedeoma pulegioides

Lamium amplexicaule

L. maculatum

L. purpureum

Leonurus cardiaca

In marrubiastrum

Lycopus americanus

L.'uniflorus

L. virginicus

Marrubium vulgare

Mentha arvensis subsp.
canadensis

N. spicata

Monarda fistulosa

Nepeta cataria

Physostegia virginiana var.
speciosa

Prunella vulgaris

Pycnanthemum flexuosum

P. pilosum

P. virginianum

SatureJa hortensis

Scutellaria galericulata

8. lateriflora

S. leonardi

S. parvula

Stachys hispida

S. tenuifolia

Teucrium canadense

Thymus serpyllum
Trichostema brachiatum

IONIA COUNTY

AJuga reptans
Blephilia ciliata
B. hirsuta
Clinopodium vulgare
Lamium amplexicaule
Leonurus cardiaca
Lycopus americanus
Monarda fistulosa
Nepeta cataria
Prunella,vu1garis
Scutellaria galericulata

S. parvula

IOSCO COUNTY

Clinopodium vulgare

Lycopus americanus

L. uniflorus

Mentha arvensis subsp.
canadensis

Scutellaria galericulata

IRON COUNTY

Clinopodium vulgare
Glechoma hederacea
Lycopus americanus
Mentha arvensis subsp.
canadensis

Prunella vulgaris
Scutellaria galericulata

ISABELLA COUNTY

Nepeta cataria

JACKSON COUNTY

Agastache nepetoides

Blephilia ciliata

Collinsonia canadensis

Glechoma hederacea

Hedeoma hispida

H. pulegioides

Lamium maculatum

Leonurus cardiaca

Lycopus uniflorus

Mentha arvensis subsp.
canadensis

M. piperita

Monarda fistulosa

Prunella vulgaris

Pycnanthemum virginianum

Scutellaria galericulata

S. lateriflora

Teucrium canadense

Thymus serpyllum

KALAMAZOO COUNTY

Agastache nepetoides
A. scrophulariaefolia
Blephilia ciliata

B. hirsuta
Clinopodium vulgare
Oollinsonia canadensis
Dracocephalum parviflorum
Glechoma hederacea
Hedeoma hispida

H. pulegioides

Lamium amplexicaule

L. purpureum

Lycopus americanus

L. rubellus

L. uniflorus

1+6

Marrubium vulgare

Mentha arvensis subsp.
canadensis

M. piperita

M. rotundifolia

M. spicata

Monarda didyma

M. fistulosa

M. punctata var. villicaulis

Nepeta cataria

Physostegia virginiana var.
speciosa

Prunella vulgaris

Pycnanthemum flexuosum

P. virginianum

Salvia azurea var. grandiflora

S. reflexa

Scutellaria elliptica var.
hirsuta

S. galericulata

S. lateriflora

Stachys hispida

S. hyssopifolia

S. palustris var. palustris

S. tenuifolia

Teucrium canadense

Thymus serpyllum
Trichostema dichotomum

KALKASKA COUNTY

Hedeoma pulegioides

Lycopus uniflorus

Monarda fistulosa

M. punctata var. villicaulis
Prunella vulgaris
Scutellaria galericulata
Teucrium canadense

mm 00mg]

Agastache nepetoides
A. scrophulariaefolia
Blephilia ciliata

B. hirsuta

ClinOpodium vulgare
Collinsonia canadensis

_j/ The three species indicated by an asterisk (*) have been collected

since the study of E. J. Cole (1901).

Glechoma hederacea

*Hsdeoma hispida

H. pulegioides

HyssOpus officinalis

Leonurus cardiaca

Lycopus americanus

L. rubellus

L. uniflorus

Marrubium vulgare

Mentha arvensis subsp.
canadensis

M. piperita

M. rotundifolia

M. spicata

Monarda didyma

M. punctata var. villicaulis

Nepeta cataria

Physostegia virginiana var.
speciosa

Prunella vulgaris

Pycnanthemum virginianum

Salvia asurea var. grandiflora

‘Satureja hortensis

Scutellaria galericulata

S. lateriflora

Stachys hispida

S. hyssOpifolia

S. palustris var. palustris

S. tenuifolia

Teucrium canadensis

‘Thymus serpyllum

KENEENAN COUNTY

Clinopodium vulgare

Dracocephalum parviflorum

Galeopsis tetrahit

Glechoma hederacea

Leonurus cardiaca

Lchpus americanus

L. uniflorus

Mentha arvensis subsp.
canadensis

M. piperita

Nepeta cataria

Physostegia virginiana var.
virginiana

Prunella vulgaris

Scutellaria galericulata

S. lateriflora

Stachys palustris var. palustris

S. palustris var. homotricha

1+7

KEMEENAW COUNTY: ISLE ROYALB

Clincpodium vulgare

Dracocephalum parviflorum

Galeopsis tetrahit

Glechoma hederacea

Lycopus americanus

L. unif lorus

Mentha arvensis subsp.
canadensis

Prunella vulgaris

Scutellaria galericulata

S. lateriflora

Thymus serpyllum

LAKE COUNTY

Mentha gentilis
Prunella.vulgaris
Scutellaria galericulata
Stachys hyssopifolia
Teucrium canadense

Thymus serpyllum

LAREER COUNTY

Monarda fistulosa
Prunella vulgaris
Teucrium canadense

LEBLANAU COUNTY

Acinos arvensis

Calamintha glabella var.
angustifolia

Clinopodium vulgare

Redeoma hispida

H. pulegioides

Lamium amplexicaule

Leonurus cardiaca

Lycopus americanus

L. rubellus

L. uniflorus

Marrubium vulgare

Mentha arvensis subsp. canadensis

M. piperita

M. spicata

Monarda fistulosa

M. punctata var. villicaulis

Nepeta cataria

Prunella vulgar is
Pycnanthemum flexuosum
P. virginianum
Scutellaria galericulata
S. lateriflora

Teucrium canadense

Thymus serpyllum

LENANEE COUNTY

Agastache nepetoides
Blephilia hirsuta
Collinsonia canadensis
Leonurus cardiaca
Lycopus uniflorus
Monarda fistulosa
Nepeta cataria

Prunella vulgaris
Pycnanthemum virginianum
Scutellaria galericulata
S. lateriflora

Stachys hispida

S. palustris var. palustris
Teucrium canadense

LIVINGSTON COUNTY

Redeoma hispida

R. pulegioides

Iamium amplexicaule

L. purpureum

Leonurus cardiaca

Lycopus americanus

L. uniflorus

Mentha arvensis subsp.
canadensis

M. piperita

Monarda fistulosa

Nepeta cataria

Prunella vulgaris

Pycnanthemum virginianum

Scutellaria galericulata

S. lateriflora

Stachys tenuifolia

LUCB COUNTY
ClinOpodium vulgare

Lycopus americanus
Mentha arvensis subsp. canadensis

Prunella vulgaris
Scutellaria galericulata
Stachys palustris var. palustris

MACKINAC COUNTY

Acinos arvensis

Calamintha glabella var.
augustifolia

GaleOpsis tetrahit

Glechoma hederacea

Lamium maculatum

Leonurus cardiaca

Lchpus americanus

L. uniflorus

Mentha arvensis subsp. canadensis

M. piperita

Monarda fistulosa

Prunella vulgaris

Scutellaria galericulata

S. lateriflora

Stachys palustris var. palustris

Teucrium canadense

MACOMR COUNTY

Galeopsis tetrahit
Hedeoma hispida
Leonurus cardiaca
Lycoyus americanus

L. uniflorus

Mentha gentilis
Monarda didyma

M. fistulosa

Nepata cataria
Prunella vulgaris
Pycnanthemum flexuosum
P. virginianum
Scutellaria galericulata
S. lateriflora

Stachys tenuifolia
Teucrium canadense

MANISTEE COUNTY

Clinopodium vulgare

Lchpus americanus

Mentha arvensis subsp. canadensis
M. gentilis

Monarda fistulosa

, Prunella vulgaris
Scutellaria galericulata
S. lateriflora

MARQUETTE COUNTY

AJuga genevensis

Clinopodium vulgare

Galeopsis tetrahit

Leonurus cardiaca

Lycopus americanus

L. uniflorus

Mentha arvensis subsp. canadensis

Monarda fistulosa

Nepeta cataria

Physostegia virginiana var.
speciosa

Prunella.vulgaris

Scutellaria galericulata

S. lateriflora

MASON COUNTY

ClinOpodium vulgare

Lchpus americanus

L. asper

L.'uniflorus

Mentha arvensis subsp. canadensis
Monarda fistulosa

M. punctata var. villicaulis
Prunella‘vulgaris
Scutellaria galericulata

S. lateriflora

Stachys mesopifolia

MECOSTA COUNTY

Clincpodium vulgare

Monarda punctata var.
villicaulis

Stachys hyssopifolia

MENOMINEE COUNTY

Leonurus cardiaca

Lycopus americanus

L. uniflorus

Mentha arvensis subsp. canadensis
Monarda fistulosa

Prunella vulgaris
Pycnanthemmm‘virginianum
Scutellaria galericulata
S. lateriflora

Stachys hispida
Teucrium canadense

MIDLAND COUNTY

Agastache nepetoides

Glechoma hederacea

Hedeoma.h18P1da

Lycopus americanus

L. uniflorus

Mentha arvensis subsp. canadensis

Physostegia virginiana var.
speciosa

Prunella vulgaris

Pycnanthemum flezuosum

P. virginianum

Scutellaria galericulata

S. lateriflora

Stachys hispida

Teucrium canadense

MISSAUNEE COUNTY

Clinopodium vulgare

Lycopus americanus

.Mentha arvensis subsp. canadensis
Prunella vulgaris

Scutellaria galericulata

MONROE COUNTY

Blephilia hirsuta

Glechoma hederacea

Leonurus cardiaca

Lycopus americanus

L. as per

L. uniflorus

Mentha arvensis subsp. canadensis

M. gentilis

M. spicata

Menarda fistulosa

Nepeta cataria

Physostegia virginiana var.
speciosa

Prunella vulgaris

Pycnanthemum flexuosum

Physostegia virginiana var. virginiana

50

P. pilosum Pycnanthemum verticillatum
P. virginianum P. virginianum

Salvia reflexa Scutellaria galericulata
Scutellaria galericulata Stachys twssopifolia

S. lateriflora S. palustris var. palustris
Stachys hispida Thymus serpyllum

S. palustris var. palustris
S. temifOlia

Teucrium canadense NEWAYGO COUNTY
Blephilia ciliata
MONTCALM COUNTY Hedeoma hispida
Monarda fistulosa
Agastache scrophulariaefolia M. punctata var. villicaulis
Blephilia ciliata Nepeta cataria
ClinOpodium vulgare Prunella vulgaris
Lycopus americanus Scutellaria galericulata
L. uniflorus Stachys hyssopifolia

Mentha arvensis subsp. canadensis Teucrium canadense
Monarda fistulosa ‘
Pycnanthemum virginianum
Scutellaria galericulata 0mm commj/
Stachys palustris var. palustris
TAcinos arvensis
Agastache nepetoides

MONTMORENCY COUNTY A. scrophulariaefolia
Collinsonia canadensis
Clinopodium vulgare Glechoma hederacea
Lycopus uniflorus Hedeoma hispida
Mentha arvensis subsp. canadensis H. pulegioides
Monarda fistulosa Lsmium amplexicaule
Scutellaria galericulata Leonurus cardiaca
Lycopus americanus
L. rubellus
'MUSKEGON COUNTY L. uniflorus
Marrubium vulgare
Agastache nepetoides Mentha arvensis subsp. canadensis
Lycopus americanus I"Mentha gentilis
L. rubella; M. piperita
Mentha piperita M. spicata
Monarda punctata var. villicaulis Monarda fistulosa
Prunella vulgaris M. punctata var. villicaulis

_fl l'rom M. Bingham's (19145) list the following species were not seen:

Elemilia giliata. g. hirsuta, SathJa glabella (synonym for Acinos

arvensisf). Satureja vulgaris (synonym for glinomdium Iglgarg),
officinalis, Ezcnanthemum ilosum, Stachys tenuifolia and

§.- miflora (7). Perilla grutescens (L.) Britt. is also reported.

he two specimens with this determination are: Biggham m” middle
beach, I. Bloomfield Twp.. August 1936 (BLH 7281+). and Stoutamire 23,53,
wet woods near Horseshoe Lake. Oxford Twp. (BLH 32101). These two
cOllections are Cgllingonia canadeggis. The additions to Bingham's county
flora are indicated by and asterisk (*).

Nepeta cataria
I"Physostegia virginiana var.
virginiana

Prunella vulgaris
Pycnanthemum flexuosum

P. virginianum

Scutellaria galericulata
S. lateriflora

Teucrium canadense
Trichostema dichotomum

OCRANA COUNTY

Lycopus americanus

Harrubium vulgare

Mentha gentilis

Monarda fistulosa

M. punctata var. villicaulis
Prunella vulgaris

Stachys palustris var. palustris

OGIMAN COUNTY

Mentha arvensis subsp. canadensis
Prunella vulgaris
Scutellaria galericulata

ommoon COUNTLZ/

Blephilia,hirsuta

Clinopodium vulgare
Dracocephalum parviflorum
Galeopsis tetrahit

Lycopus americanus

In uniflom

Mentha arvensis subsp. canadensis
Prunella vulgaris

Scutellaria galericulata

OSCEOLA COUNTY

Monarda punctata var. villicaulis
Prunella vulgaris

51

Scutellaria galericulata
Stachys hyssOpifolia

OSCODA COUNTY

Lycopus uniflorus

Mentha spicata

Monarda punctata var. villicaulis
Prunella vulgaris

Scutellaria galericulata

OTSEGO COUNTY

ClinOpodium vulgare
Lyc0pus'uniflorus

Mentha arvensis subsp. canadensis
Nepeta cataria

Teucrium canadense

OTTAIA COUNTY

Blephilia hirsuta

ClinoPodium vulgare

Iamium purpuremm

Leonurus cardiaca

Lchpus americanus

L. rubellus

L. uniflorus

Mentha arvensis subsp. canadensis

M. spicata

Monarda fistulosa

M. punctata var. villicaulis

Physostegia virginiana var.
speciosa

Pycnanthemum verticillatum

Scutellaria galericulata

S. lateriflora

Stachys hispida

S. hyssopifolia

Teucrium canadense

_2/ Undoubtedly occurring in this county are the species 1122912 2115—3113
and Sggtellaria lateriflora (Darlington, 1937), but no herbarium

specimens have been seen.

PRBSQUE ISLE COUNTY

Calamintha glabella var.
angustifolia
ClinOpodium vulgare
GaleoPsis tetrahit
Hedeoma pulegioides
Lycopus americanus
L. uniflorus
Monarda.fistulosa
Prunella.vulgaris
Scutellaria galericulata
S. lateriflora
Stachys hispida
Teucrium canadense

ROSCOMMON COUNTY

Dracocephalum parviflorum
Lycopus americanus

L. uniflorus

Mentha arvensis subsp. canadensis
Monarda fistulosa '

Nepeta cataria

Prunella vulgaris
Scutellariargalericulata

S. lateriflora

SAGINA' COUNTY

Leonurus cardiaca
Monarda fistulosa
Scutellaria lateriflora
Teucrium canadense

Thymus serpyllum

SANILAC COUNTY

Glechoma hederacea
Lamium amplexicaule
Lycopus americanus
L. uniflorus
Prunella vulgaris

SCROOLCRAFT COUNTY

Calamintha glabella var.
angustifolia

Clinopodium vulgare

Galeopsis tetrahit

52

Glechmma hederacea

Lycopus americanus

L. uniflorus

Mentha arvensis subsp. canadensis
Monarda fistulosa

Prunella vulgaris

Scutellaria galericulata

S. lateriflora

Stachys palustris var. palustris

SHIANASSNE COUNTY

Blephilia hirsuta
GaleOpsis tetrahit
Leonurus cardi aca
Lycopus americanus
Mentha piperita
Scutellaria galericulata

ST. CLAIR COUNTY

Agastache nepetoides

A. scrophulariaefolia
Blephilia ciliata

B. hirsuta

Clinopodium vulgare
Collinsonia canadensis
Glechoma hederacea
Hedeoma pulegioides
Hyssopus officinalis
Lamium amplexicaule

I» maculatum

Leonurus cardiaca
Lycopus americanus

L. asper

L. rubellus

L. uniflorus
Marrubium.vulgars
Mentha arvensis subsp. canadensis
M. gentilis

M. piperita

1‘. spicata

Monarda didyma

M. fistulosa

M. media

M. punctata var. villicaulis
Nepeta cataria

Prunella vulgaris
Pycnanthemum virginianum
SatureJa hortensis
Scutellaria galericulata
S. lateriflora

53

Stachys hispida
S. palustris var. palustris
Teucrium canadense

Thymus serpyllum

ST. JOSEPH COUNTY

Agastache nepetoides

Clinopodium vulgare

Collinsonia canadensis

Hedemma pulegioides

Leonurus cardiaca

Lycopus americanus

L. rubellus

L. uniflorus

Mentha arvensis subsp. canadensis
M. piperita

M. spicata

Monarda punctata.var. villicaulis
Prunella vulgaris

Pycnanthemnm virginianum
Scutellaria galericulata

Stachys hispida

S . hyssOpifolia

Teucrium canadense

TUSCOLA COUNTY

Physostegia virginiana var.
virginiana
Pycnanthemum virginianum

VAN RUBEN COUNTY

Agastache nepetoides

Clinopodium vulgare

Collinsonia canadensis

Hedeoma pulegioides

Leonurus cardiaca

Lycopus americanus

L. rubellus

L. uniflorus

Mentha arvensis subsp. canadensis
M. spicata

Monarda fistulosa

M. punctata var. villicaulis
Prunella vulgaris
Pycnanthemum virginianum
Scutellaria galericulata
Stachys hispida

S. hyssOpifolia

Teucrium canadense

WASHTENAM county

Agastache nepetoides

A. scraphulariaefo lia
Blephilia ciliata

B. hirsuta

Collinsonia canadensis
Dracocephalum parviflorum

’Galeopsis tetrahit

Glechoma hederacea
Hedeoma hispida

*Hyssopus officinalis

Lamium amplexicaule

*L. purpureum

Leonurus cardiaca

Lycopus americanus

L. rubellus

Le uniflorus

Marrubium vulgare

Mentha arvensis subsp. canadensis

M. gentilis

M. piperita

M. spicata

Monarda didyma

M. fistulosa

M. punctata var. villicaulis

Physostegia virginiana var.
virginiana

P. virginiana var. speciosa

Prunella vulgaris

Pycnanthemum pilosum

P.‘virginianum

Scutellaria galericulata

S. lateriflora

SatureJa hortensis

Stachys hispida

Teucrium canadense

_5/ Species marked with an asterisk (*) indicate species not included

inflwalpole's (192k) and Hermann's (

1936a, 1937) additions to the flora.

Stachys ngustris var. pglustris is reported by Walpole as being common.
No specimens of this'species were seen.

Sh

Thymus serpyllum
Trichostema brachiatum

WAYNE COUNTY

Agastache nepetoides

A. scraphulariaefobia
Blephilia hirsuta ‘
Collinsonia canadensis
Dracocephalum parviflorum
Hedeoma hispida

H. pulegioides

Glechoma hederacea
Lamium amplexicaule
Leonurus cardiaca
Lycopus americanus

L. asper

L. rubellus

L. uniflorus

Marrubium vu1gare

Mentha arvensis subsp. canadensis
M. piperita

M. spicata

Monarda fistulosa

M. punctata var. villicaulis
Nepeta cataria
Physostegia aboriginorum
P. virginiana var. virginiana
Prunella vulgaris
Pycnanthemum flexuosum

P. virginiamm

Salvia reflexa
Scutellaria galericulata
S. lateriflora

Stachys hispida

S. tenuifolia

Teucrium canadense

“BOND COUNTY

. Clinopodium vulgare
Lycopus americanus

Le uniflorus
Scutellaria galericulata

55
POLLEN GRAIN STUD ms

The pollen grains of the Labiatae are relatively uniform morphoIOgi-
cally. but have a rather conspicuous pattern of variation in number of
colpi. Differences in number of colpi were first recognized by Pritzsche

(1832) and Mohl (1839). Evidence from pollen grain morphology and
cytology has been little used for classification. although Erdtman (1916)
suggested that such data may support a tentative division of the family
into two groups possibly deserving subfamily rank. It will be shown
subsequently in this paper that some species and even varieties also

can be distinguished on the basis of minor technical characters of
pollen grain morphology such as exine sculpturing and shape and size of
the grains (cf. discussion. pp. 57-62 and key to species or groups of

species based on pollen grain morphology. pp. 63—68).

Materi_a1 and Metmdg.
hr the majority of species studied. anthers were removed from herb-

arium material at Michigan State University. the Gray Herbarium, and the
University of Michigan. Some examinations were also made of pollen
grains obtained from fresh material growing in the Deal-Garfield Botanic
Garden of Michigan State University. from field collections. and from
cultivated m plants grown by Dr. Ray Nelson of Michigan State
University. The latter are plants of M. pipgrita cv. Mitcham which had
no special treatment or were treated in one of the following ways:
Irradiated with cobalt-60 at 5,000/r, colchicine treated, or crossed
with 3,. m.

The following species occurring naturally in Michigan have not been

considered in the pollen grain studies because of insufficient material:

56
Agastaghe foeniculum..AJgg§.genggensis, yyggpgggvirginicus. Mentha
rgtundzfolia, Eygpggthemum zerticillatum, Salvia reflexa. §cutellaria

leogggdi. and Trichostema dichotgggg.
Voucher specimens of living material investigated.have been deposited

 

in the Michigan State University Herbarium. A set of microscope slides
of acetylated pollen grains also has been filed in the Michigan State
University Herbarium. Specimens from which pollen grain studies have
been made are listed in Table II. pp. 714—77. for each specimen there

is included in the table the collector and number. herbarium. slide
number. condition of plant (either from fresh or from herbarium material).
method of preparation (either acetocarmine or acetylation). and reference
to illustration in the plates.

Three types of slide preparations were usually made for each specimen:
(1) anthers crushed in.glycerine Jelly for examination of pollen grains
in their 'natural' condition, (2) anthers crushed in glycerine Jellyb
acetocarmine mixture for determination of number of pollen grain nuclei
at anthesis. and (3) pollen grains acetylated (and sometimes chlorinated)
using Erdtman's (l9h3,1952) process for examination of the exine sculptur-
ing. shape class. size measurements and other characteristics. The
second.method above has been modified.from Leitner (1937) and is de-
scribed in detail as follows. The anthers are first crushed in.me1ted
glycerine Jelly on a.microsc0pe slide. A drop of acetocarmine is mixed
with the Jelly and the cover glass applied. The slide is then heated
gently over an alcohol flame for 5 to 10 minutes. The preparation is
then allowed to stand for one to five (or more) hours while the nuclei
absorb the stain. Slides thus prepared are semi-permanent. Por deter~

mination of nuclear content of the pollen grains. the preparation is then

57
examined under LL30 x with very brilliant illumination. This method is
not satisfactory for anthers which have been stored previously in acetic
acid.

Plowers nearest the base of plants at the beginning of their flower-
ing period were the most satisfactory sources for pollen grains.

Measurements of acetylated pollen grains were made at a magnification
of 970 I with a calibrated ocular micrometer in a standard Bausch and
Lamb monocular research microscope. Unless otherwise indicated. the
measurements given are the average of ten grains.

Photomicrographs were made with a Zeiss research microscOpe and Leitz
pillar-type bellows camera with 3}“ x hi" plates. Ansco Type 282 (how)
panchromatic film was used. As reproduced in this study. the photo-
micrographs are at a magnification of 1020 1 unless otherwise indicated.
A duplicate set of photomicrographs of acetylated pollen grains has been

filed in the Paleobotanical laboratory of Harvard University.

Mien.

The results of the examinations of pollen grains are summarized in
Table I. In the following paragranis. inportant features of the pollen
grain studies have been indicated. The terminology employed is that
of Erdtman (191.3, 1952) and Brdtman and Vishnu-Mitter (1958).

Pollen grains of investigated Michigan species are radiosymmetrical
(having more than two planes of symmetry) and sonicolpate (the apertures

are arranged parallel to the polar axis). _2/ The apertures are longi—

_9_/ Mentha gentilis. M. pipgrita cv. Mitcham (both non-irradiated and
irradiated). and Monarda m have both sonicoopate and pancolpate
pollen grains ( the latter are grains with 10 or more apertures that are
not all arranged parallel to the polar axis).

58
tudinal and technically referred to as colpi (sing. colpus) and normal
grains of all Michigan species are either tricolpate or hexacolpate.
The species with tricolpate grains are very constant in respect to this
character. although one species. W brachiatum contains many
deformed grains. This may be due to the depauperate specimen from
which the anthers were taken. In some species ordinarily having hexa-
colpate grains. exceptional cases have been observed in which 8. 10.

and 12 or more colpi occur in plants from natural populations. The

species m; w. W virginiana. and Sgturejg hortengig

have hexa- and octacolpate grains. m dim and Mentg gentilig
have 6—. 8-. 10-. and 12— (or more) colpate grains. Both non-irradiated

and irradiated plants of 3.9.9.529. m cv. Mitcham have 8. 9. 10. 12.
and other polycolpate grains. while colchicine treated plants of this
cultivar have only octacolpate or occasional nonacolpate grains. In

5. piperita cv. Mitcham and also in 5. man; the maJority of the
grains have more than six colpi, but in the other species the hexacolpate
grains outnumber those with more than six colpi.

An outline (after Briquet. 1895) of subfamilies. tribes. and sub-
tribes in which the Michigan genera of the Labiatae have been placed is
provided below. The number of colpi occurring in the members of each
tribe and subtribe is indicated. It should be noted that both tri-

and hexacolpate types are found in the subfamily Stacknroideae (Lamioideae).

Subfamily AJugoideae. tribe AJugeae: M. geyrium.

Trichosteg THICOLPATB

Subfamily Scutellarioideae: M TRICOLPATE
Subfamily Stachyoideae (Lamioideae)

Tribe Marrubieae: mm THICOLPATE

Tribe Nepeteae: Agastache. Daggceghglum. Qlechog.
ta
Tribe Stacmeae (laminae)

mACOLPATN

59

subtribe Brunellinae: Prunella HEXACOLPATE
Subtribe Lamiinae: galeOQsis. Lamium, Leonurus.

thsostegia, Stacgyl TRICOLPATE

Tribe Salvieae: Salvyg HEXACOLPATD

Tribe Monardeae: ‘flgggrgg, Blephilig HIIACOLPATE

Tribe Saturieae:
Subtribe Melissinae (Saturinae): Aginos, gedegma,

galamintha. glinomdium. Suture]; HBXACOLPATI

Subtribe HyssOpinae: gygggpug HBXACOLPATS
Subtribe Thymineae: Exchanthgggm. £hymug_ HEXACOLPATN
Subtribe Menthainae: nggpgg, Mentha HEIACOLPATE
Subtribe Perillinae: Collinsonia HEIACOLPATB

Leitner (l9h2) reported the occurrence of bi- and trinucleate pollen
grains at anthesis in the Labiatae. Erdtman (19h5) noted that some of
the species Leitner had found to be binucleate were tricolpate and some
that she found to be trinucleate were hexacolpate. In the present study
this correlation of the binucleate condition in the tricolpate grains
also has been found.lQ/ In the species with pollen grains having more than
six colpi. the grains are also trinucleate. The hexacolpate grains of
Mgnth§,ggntilis and M, pipggitg_cv. Mitcham (P1. I. Fig. 13s) contain no
protoplasm but the grains with more than six colpi (rig. 13a) do some-
times contain protoplasm and have the trinucleate condition.

In lrdtman's tentative division of the Labiatae into two groups based
on pollen grains (the first group containing species with binucleate,
tricolpate grains and the second group species with trinucleate, hexas
colpate grains). the Lamiinae was not considered. In the present study.
this subtribe has been found to contain only members with binucleate.
tricolpate grains. Consequently it is suggested that this subtribe be

included with ertman's first tentative group.

19] The vegetative nuclei do not take as deep a stain as do the generative
nuclei. In the photomicrographs. therefore. only the generative nucleus
can be seen in the tricolpate grains (eg. Pl. I. Pig. 1), and only the

two sperm nuclei are visible in the hexacolpate grains (eg. Pl. I. Fig. 7).

60

In this study three classes of amine sculpturing have been recOgnized:
faintly reticulate. fine reticulate and reticulate.l;/ The reticulate
pattern is made up of muri (ridges) and lumina (spaces). The hexacolpate
species have only the reticulate sculpturing pattern. One tricolpate
species. Marrubium m differs in its reticulate pattern from the
other species in that the muri seemingly are larger than the lumina (Pl.
III. rig. 6a'). The lumina of zgzsgstegia virginiana var. virginiana
(P1. V. Fig. 13a) and var. gpgcioga are conspicuously larger than those
of other reticulate species. They range in diameter from two to four
microns in contrast to lumina of other species which are less than two
microns in diameter.

The exines of Teucrium ggngdgnse (Pl. II, Fig. 1a' and lc').
Trichgstemg prachiggmm (Pl. II, Fig. 3c). and.§@yeostegig virginiana var.
virginiana and var. spgciosa are two microns thick or thicker. The other
tricolpate species have grains with thinner exines.

In each species the margins of the colpi have been found to have a
ragged appearance. At the region of the colpus the margins of the exine
end either abruptly or with gradual thinning as seen in P1. 11, fig. 6a
or P1. IV. rig. llb'. The only exception is Collinsonia ggpgdensis (Pl.
IV. Pig. h7a') in which the exine thins abruptly at 2 microns from the
edge of the colpus and thus appears half-umbonate.

The shape classes of the pollen grains of Michigan species of the

Labiatae are subprolate. prolate spheroidal. and oblate spheroidal (the

11/ .A cultivated species.‘L§migm,album is psilate (smooth) by ordinary
light microscopy (P1. VI, Fig. 17). However. by the carbon replica tech-
nique used with electron microscopy. Bradley (1957) found a pollen grain
of this species with the exine finely punctate and the colpus warty.

61

shape class subprolate is determined as a ratio of length of polar axis
divided by the equatorial diameter and ranges between 1.33 and 1.1% the
prolate spheroidal shape ranges numerically between 1.114 and 1.00; and
oblate spheroidal has a numerical range from 1.00 to 0.88). Shape class
variation is slight in these species. The shape class oblate spheroidal
is found only in gall-is; 9mg var. grandiflora. This shape is similar
to that of the cultivated species. §_. coccineg (P1. VII, Fig. 23) which
is oblate (numerical ratio of 0.75 to 0.50).

The polar diameters of normal pollen grains range between 18 and
56 microns (based on measurements of 10 grains). The polar diameters of
most species. however. range between 30 and (+0 microns. Generally the
tricolpate grains are smaller than those that are hexacolpate. w
m has the smallest tricolpate grains (polar diameter 18p.) and

and Mania 122m var. 2221222 has the largest (5211). In the
hexacolpate species. the smallest grains occur in $192212. uniflorus

(polar diameter 27:1) and the largest in Monarda gm (56p). M;
semllum, an escape from cultivation. has two distinct means in the
range of polar diameters (P1. XI. rigs. 35a and 35c) in the same plant.
The photomicrographs of octacolpate grains of m s r. c the
‘W- and SatureJa hortensis (P18. 1:. II. and 111) show that these
grains are larger than the hexacolpate grains from the same plants. But
then an average polar diameter of 10 octacolpate grains is taken (see
Table I), this is found to be about the same as the average polar diame.
ter of 10 hexacolpate grains. This similarity in size of hexa- and octa-
colpate grains is contrary to the reports in the literature (such as
Maurizio, 1956). In M m. however. the grains having more than

six colpi are larger. Mentha gntiln (probably of hybrid origin. see

62
p. 37) has a great variation in number of colpi and size (from 10p to
60);) similar to 11. aperita cv. 141th (P1. XIII. Fig. 1+2). although
the latter generally has larger polar diameters.
In the key to species based on pollen grain characteristics certain
groupings consider only polar and equatorial diameters. These differences
are more or less arbitrary however, and may be influenced in part by the

acetylation preparation. Yet when the measurements of acetylated grains

of four species in this study (Balsa Mo 3:. w- m
BEL-£12139 and m fiigttilosa) are compared with measurements of
glycerine Jelly mounts stained with gentian-violet or fuchsin in Ikuse's

work (1956). there is considerable agreement in size measurements in the
two studies. The acetylated preparations show a greater range in size
variation which may be due to the collapsed or expanded condition of
some of the grains measured.

Further study is needed especially in species in which irregularity
occurs in the number of colpi and size of pollen grains. These species
also should be investigated chromosomally. experimentally. and taxonomies].-

1: throughout their entire geographic range.

63

Key to Species or Groups Based on Pollen Grain Characteristics

Some of the Michigan species that occur in natural pepulations or
are cultivated can be keyed out to species while others can only be
placed into groups as follows: Group I includes two tricolpate. faint-
ly reticulate species with polar diameters between 2151 and 2611; Group II
includes Ll tricolpate. fine reticulate species with polar diameters
between 2143:. and 36p; Group III includes 15 hexacolpate species with
polar diameters between 27p and 3611; Group IV includes 7 hexacolpate
species with polar diameters between 3911 and 11611; and Group V includes
species with both hexacolpate and octacolpate grains.

The sizes of pollen grains are the average measurements of 10 grains
prepared by the acetylation method. It should be noted that distinc-
tions of species and groupings based on size alone are more or less
arbitrary.

l'or additional characteristics of the '49 species in the key. see

Table I. and the illustrations.

A. Pollen grains tricolpate, sculpturing reticulate. fine or faintly
reticulate. or psilate. . . . . . . . . . . . . . . . . . . . .

M. Pollen grains hexacolpate. sculpturing reticulate. . . . . . .

B

Q

B. hinereticulate.....................c
BB. kins fine or faintly reticulate, or psilate. . . . . . . G

C. PolardiameterlessthanBOu................. 1)

CC. Polar diameter greater than 3011. . . . . . . . . . . . . . . . E
I). Equatorial diameter 27p. 1: 22p. polar diameter 261:. shape

class subprolate, muri seemingly larger than lumina. exine

2p. in thickness (Pl. III. fig. 6). . . . flagrubium m

II.

6%

DD. Equatorial diameter 26.7]: x 23.511. polar diameter 21+.5u,
prolate spheroidal. muri not larger than lumina. exine less
than 2p. in thickness (Pl. VI. Fig. 19). . Stachyg hispida

Polar diameter greater than 50p. prolate spheroidal (equatorial

diameter 56.3u x 50p, polar diameter 51.511). . . . .msostegig

virginiana var. smciosa

Polar diameter less than 5031. subprolate. . . . . . . . . . . . 1'

1'. Equatorial diameter 16.71). x 37u. polar diameter 142.831 (Pl. V.

Fig. 13). . . . . . . msostegia WV”. virginig
FF. Equatorial diameter 142.91.: x 32.511. polar diameter 214); (P1. V.

l'ig.16).................Qa_1_e_op§1._§_§_e_t_r§h_i1
Exine psilate (equatorial diameter 27;). at 25p. polar diameter 26):.
prolate spheroidal; Pl. VI. Fig. 17). . . . . . . mm
Exine fine or faintly reticulate. . . . . . . . . . . . . . . . H
H. Exinefaintlyreticulate. . . . . . . . . . . . . . . . . I
EH. Exinefinereticulate...................J’
Shape class prolate spheroidal. polar diameter 24p. to 26p. Group
I: equatorial diameter 26}: x 23.511. polar diameter 2%. . . . .
W elliptica var. hirsuta; equatorial diameter 26.511 x
ZhJ-Hi. polar diameter 25.711. . . . . . . . . . . Mmcflatm
Shape class subprolate. polar diameter 26.7}1 (equatorial diameter
30.7u I 21:11; P1. III. Fig. 5). . . . . . . . Scutellaria lateriflora
J. Polar diameter greater than 3611 (equatorial diameter 02.8}1 x

3111. subprolate. Pl. II, Pig. 8). . . richestema brachigtum
JJ’. POlardiameterlessthan36u. . . . . . . . . . . . . . . .K
Polar diameter less than 20p (equatorial diameter 19.1w. 1 16.511.

polar diameter 18u. subprolate; Pl. V. Fig. 14). Leonurus ms;

00.

Q.
9.9..

65
Polardiametergreaterthan2011. . . . . . . . . . . . . . . . L
L. Exine Zn or more in thickness (equatorial diameter 38.5u x
28.911. polar diameter 3311. subprolate; Pl. II. Fig. l. . .
WW
LL. ExinelessthanZuinthickness. . . . . . . . . . . . . . ll
Polar diameter greater than 30u (equatorial diameter 30.8u x 28.9n.
polar diameter 31.511. prolate spheroidal). . . w W
PolardiameterlessthanBOu.................. N
H. Polardiameterlessthan23u............... 0
1m. Polar diametergreaterthan23u. . . . . . . . . . . . . . P
Polar diameter 21.511. equatorial diameter 231: x 2111. prolate
spheroidal...................MM
Polar diameter 22.1111. equatorial diameter 26): x 21.711. subprolate,
(P1.V.1'1g.15).............. eonuru W
P. Polar diameter 2911 (equatorial diameter 35.711 x 26.611, sub-
prolate. (P1. VI, Fig. 18). . . . . . . . gala-gm
PP. Polar diameter between 2131 and 2711. Group II: equatorial
diameter 26.511 x 2311. polar diameter 214-11. subprolate, Pl. II.
Pig. (1+. . . . . . Sgutellarig galericulata; 2811 1: 22.1111. 21111.
subprolate. Pl. VII. Fig. 21. . . . . . §§aghys w var.
Emotricg; 27.511 x 2211. 26.511. subprolate. P1. VI. Fig. 20. .
. . . . §§§£hz§_hzsggp§§olia; 27.5u x 27p. 27u. prolate spheroidp
a1. . . . . . m palustrig var. w; 3011 x 21411. 2711.

subprolate. Pl. II. Fig. 2. . . . . . . . . . . .m rgptang
Shape class subprolate to prolate spheroidal. . . . . . . . . . 8

Shape class oblate to oblate spheroidal. . . . . . . . . . . . R

R. Shape class oblate. equatorial diameter 39.611 1: 511.1411. polar

66
diameter 66.311 x 5111 (P1. VII. Fig. 23). . 5.322151 coccinea
RR. Shape class oblate spheroidal. equatorial diameter 2511 x 26.911.
polar diameter 2811 x 21111. . . Salli; 222.1222 var. grandiflora
Exine at colpus margin abruptly thinning at 211 from the edge and
appearing half-umbonate in polar median section (equatorial diameter
11,211 x 35.511. polar diameter 141.511. subprolate. P1. XV. Fig. ’47. . .
921112521112 W
Exine at calms margin not thinning abruptly at 211 from the edge.
the same thickness throughout or gradually thinning as seen in
mid-opticalsection...................... T
T. Pollengrainshexacolpateonly. . . . . . . . . . . . . . . U
TT. Pollen grains hexap. octa.. nonap. or polycolpate. . . . . . V
Pollen grains with two distenct sizes in the same plant (equatorial
diameter 33.611 1: 27.1111. polar diameter 3111. subprolate; 25.311 x
2211. 211.61. subprolate, P1. 11. Fig. 35. . . . . m germllum
Pollen grains more or less uniform in the same plant. . . . . . V
V. Polar diameters between 2711 and 361.1. Group III: polar diame-
ter 2711. equatorial diameter 28.811 x 25.611. prolate spheroidal.
P1. 111. 113. 1+1. . . . . . mm: 27.311. 28.1411 1
25.711. prolate spheroidal. Pl. XI. Fig. 3b. . . . . . Hyssom
gfficinalig; 2911. 29.711 x 2811. prolate spheroidal. . . . . .
Lycoms rubellug; 30.311. 31.111 x 27.511. prolate spheroidal. Pl.
XII. figs 38 and ’40. . . . . . Lycoms gericanus; 30.311.
311.911 1: 31.711. prolate spheroidal, Pl. III. Fig. 7. . . . . .
égastachg nepetoides; 32.611. 35.811 x 28.711. subprolate. P1. IX,
Fig. 28. . . . . . Blephilia hirsuta; 33.711. 33.711 1: 33.1411.
prolate spheroidal. . . . . . ficnanthemum flexuosum; 33.211.

67

35.2u x 26.6u. subprolate. P1. IV. Fig. 10. . . . . . Egpgga

gataria: 33.5p. 37.5u x 3hu. prolate spheroidal. Pl. IX. Fig.

30. . . . . . mm: 33.711. 36.511 x 33.711. prolate

spheroidal. P1. X. Fig. 33. . . . . . Clinopgdium zglgagg;

33.5p. “0.7p x 35.9p. prolate spheroidal. P1. I. Fig. 31. .

. . . . Galamintha glabella var. agggggggglia; 35p. 37.2p x

29.7p. subprolate. . . . . . Hedeoma pulegioides; 35.9p. 38p 3

31.8u. subprolate. P1. XIII. Pig. #2. . . . . . flg§£h2_§gzensig

subsp. ggpgdensis; 35.9u. 38.6u x 30.hu, subprolate. Pl. IV.

Fig. 12. . . . . . Eggpella vulgaris: 30.3u. 3h.9u x 31.7u.

prolate spheroidal. Pl. III. Pig. 8. Agastachg scr0phu1ariaefolig
‘VV. Polar diameter between>39u and b5.6u. Group IV: polar diameter

39p. equatorial diameter h0.5p.x 35.6p. subprolate. P1. 11.

Fig. 27. . . . . . Elephilig giliata: 39.up. h0.7u x 33.1p.

subprolate. P1. IV. Fig. 11. . . . . . Dracocephalgn pgrvifloggg;

110.311. 39.711 x 35.611. prolate spheroidal. P1. x1, Fig. 29. . .

. . . Eggggga,hispida; “2.2u, hh.5u x h2u, prolate spheroidal.

. . . . . Mgnarda media; hh.5u. h6.hu x 38p. subprolate. Pl. VIII.

Fig. 25. . . . . . Ignarda figtuloga: “5.3u. h6.lu x 37.6u.

subprolate, P1. IV, Fig. 9. . . . . . Glechoma hederagea; h5.7u.

h7.hp x 3h.3p. prolate spheroidal. P1. VIII. Fig. 26. . . . . .

Monarda pgpctatg var. villigaulis

Pollen grains hexa— and octacolpate. Group V: hexacolpate. 33.3p.
3h.5u x 30.1u. subprolate. octacolpate grains (average of 3) 32.hp,
Pl. 1. Fig. 32. . . . . . Satureja hortensis; hexacolpate. 33.6p.

39.5u x 35.2u. prolate spheroidal. octacolpate grains (avg. of two)

38.5p. P1. XI. Figs 36 and 37 ..... . Ezgnanthemum virginiapum:

68
hexacolpate. 35.5u. 37.9u x 31.9u. subprolate, octacolpate grains
35-511.P1.XII.Fig.39........ .. . .. .. Mm
.Pbllen.grains hexa-. octa-. and polycolpate. . . . . . . . . . x
:1. Polar diameter of hexacolpate grains 27.9u. subprolate. polar
diameter of octacolpate grains zhup35u. decacolpate grains
37upb0u. also with very small grains between lap and 15P° .
Mgentilis
1!. Polar diameter of hexacolpate grains 55.9p. of octacolpate
grains (avg. of h) 59.hpn of decacolpate ( one grain) 56.7n.

dodecacolpate (avg. of two) 61.511. no small grains. P1. VII,

Fig. 214'. e e s e e s e e s e s e e e e e 0 'MM

AaeummV
Amesnmv
Ame:aev
Awm1a~v
AaeummV
3::va

Ammumwv

Ammuamv
Amnnzmv
Ammuamv
AmmaomV
lam.ewv
A021~mv

«eschews may

om.mm
ma.om
om.me
m~.mm
am.em
Nn.om
Hm.mm

Amnummv mm.om x Amenamv mm.mm
Amm10mV «H.mm x Ameuamv no.0:
Aa=1~mv He.am u Aomraev ma.oe
Aamummv Ne.em u Amm1m~v a~.mm
Amn1mmv eo.m~ u “Henemv Ho.am
Ammuomv no.an s Ammumnv Ho.en

Ammuowv HH.~N w Awmunwv mo.am

“sea .mo .mmfimaw Hesse hump was pony

NN.:N
35.0N
wo.:~
ua.mm
NN.5N

mo.om

aeooaman
amaom

Aemuaav oo.m~ u Ammnmmv ~n.o~

Ammvoav mo.e~ x Aenummv ma.om

Ammuauv am.m~ u “augemv oo.o~

Ammummv ~m.m~ w “Henemv cm.mm

Ammuomv wo.:~ w Awmnnmv N~.om

Anm.awv ma.an u Awesome om.~e
nae; x £33

Amaoao«a_aav
uoaeaman
asaaoomaua

modueanouomamno dunno noaaom

opmaoamndu
oumaouanam
emsnoanpdm
mammoampdm

Heeaoeona.
opmdoum
23893.
epsaoaa

eamHoaanum

eamaoaapam
mamaouandm
SB 283 a

sundown
opmaoaaeam
opmaoaapsm

camaoanpam

Shape Class

opmasoupoa
mamasoaaon
epmaauaaoa
osaaeoanou
epmaaeapon
ovmaaowpoa

eumflsodaon

epmddoupou
onaa
eamaaoapou
3953
opmagoupou
aqpqaem
camaaoapoa
oaaa
oaaaeoason
cane
opmasoupou
«cam

Sculpturing

Ho.ofColpimnmmmmmmxo\0\oxo\oxo

.H magma

mmmmmm

N

no. of Nuclei
at Anthesis

awamdeb maaoashm
asaoamupama anammaeoooman
essences: machooaa
.3933 mnemoz
maaomemaamaanmoaom .4
mouaoaoaea omompmmmd

enemas» asapsaamz

madbhmn .m

mamaaoauoamm .m

euoaaaeonaa .m

damnaam

.uss soaomaaao manganeseom
ouaodmamo.asauodma
easemou swshd

aaaeamoman «anaconouaa

ue«009m

7O

Ammnmmv
“3&8
Ammammv
Ammuemv
Aemuamv

Swine

“amummv
nemuomv
“omnaav
833
Ameummv

nauseous may

oa.2m
3H.em
mm.a~
:~.am
mm.Hm
mm.mm
Hm.m~
oa.em
mm.-
:H.ma
mm.am
om.~a

nepoamHn
amapm

Ammnawv
Ammumav
58..an
“Hmnmmv
nem.e~v
Ammummv

homnemv
nemuomv
Ammumav
iamuanv
Ameaamv

Hm.mm
::.NN
Nw.mN
.o.w~
mm.mm
no.3m
mH.mm
om.mm
Ha.am
mm.mH
oH.om
$0.5m

#333 N upwcmn

Amm.amv
Amnummv

Hm.mm
mo.m~

A883 mmém

Ammummv
Ammuemv
Am~-mmv

318
8?an
“nausea
Ammaamv
Aomummv

Amaoaowa adv
nepeamun
HmaHOpmde

ma.mm
Hm.on
me.o~
no.am
mm.me
oo.m~
ae.oH
m~.om
oa.me

ammuouomam
oumaoam

sumaoaapmm

$3833
oamaoun

opmaoaaasm
AddaoamgAm
opsaeam
Hmduoammmm
oumaohn
Hmdaoaomaw
epmaoaa

mamaoaapam
eumHoaandm
emmaoampmm
33 83am

emmaoaa

oamaoaapsa

Shape Glass

opmaaeapoa
mamauoauoa
Guam
canadeaaou
Odd.“
assesses.“
«can
opmaduapoa
OQHM
opeaaoascs
handed“

oamfluum
opmadoamon
camasouuoh
mafia
oamaduaaoa
scam
oumamuwpoa
epmamuaaca

ousanoapmn

Sculpturing

AdoduaonoOV nodamahoaosammo manna aoaapm

mmmmmmmnmmmnm
N

N

N
.1 .u
n. .1.“
1“ CB
.0 .m.m

t
.m .M.m
m .nm
.N .m
.H OHQGB

seaanaa .m
mmoaapoaom
.um» manuuuamm .m
manpmaamm
.ama uuapmaamm manompm
goaaaa ...H
oasmofiwoaaam .4
Sudanese .A
sagas asaama
pandas.» aaaaooaee

asuummdasaama .A

condense adadaoca

N mmoaoomm .am» mamaadmaw> .m

mamanwmaaa
.amb mqsdnuwaab .m

annomfimaaopm mwwopmouhnm

weaoomm

 

‘wl-U 1
Ar :cd‘CQV\\ ‘1 “‘1

 

71

Am31emV
“amsmmv
Ammuomv
Aoenmmv
AweuamV

Ammumev
“weunav
Ame-~ev
oa.a~ x
oo.Hm u
mm.em w
“amueuv

Ammuomv

em.oe
ao.mm
ee.Nm
mm.mm
om.~e
oa.em
oa.om
oa.om
mm.mm
ma.ee
me.me
a~.o~
om.oo
aH.m~
~m.w~
m:.a~

Renegade may
uoaoasan
Adana

“He.env em.mm

Aemuemv
Aannmmv
Aoaummv
Ameuoev

AmmuomV
neeunmv
Awesomv

Ammumuv
Aemnomv

Ammlonv

Ha.m~
am.mm
mn.nm

NH.Nd

oa.ma
mo.mm
em.em
e~.m~
am.en
am.om
me.a~
om.H~

53: s names

Aeaoaoua_a«v

N

K

H

Aa31mmV
“oeuwnv
Aging
Ae3-ame
Aeeumev

Ammuaev
Ammuaav
Ammtmev

A Siam v
Ammnamv

Ammummv

nepmamqn
Hmuaoamavfl

Avouuummoov eoaoeaaoaomssmo manna aoHHpm

nu.mm
ma.um
an.mm
mm.o:
Hm.d:

mm.em
ee.me
am.ae
mm.o~
mn.om
mo.m~
ww.n~
mn.m~

SE 223.
epmaoaa

opmaoumpsm
mamaoaagaw
epmaeumaum

anewouomam
oamaoaa

opmaoaapmm

sumaoampam

Hedgehognu
opmaoaa

oumaoamgam

cheapo
33893.
eamano

ommaoannum

Heeaoaomae
eamaoum

Shape Glass

unmadeapma
opmadouaou
camasudpoa
somadowuou
mamaaoaaeu
mamadowpma
camadowaen
mamadowaoa
mpdeOWucu
oumamofiuma
mamaaoauoa
oumadoaaon
oumaaoauoa
oamdaowpeu

mamamoaaou

seam
manageapoa

OddH

Sculpturing

m m
o n
o m
o m
m m
NH
OH
m m
m m
o m
o m
m
m
m n
m N
m N
.1 .u a
D. 111.
1. c 8
_w «M.»
+v
.u .u.u
.m a.»
.N .u
.H capes

seamen: .m
modaoamoasa esoomom
nausea: .m

epaaaao eaaaeaoam

waves .2

«sauna .x

muoaaouam .:
maammoaaaab

.amp mamaoadn manage:

manoeoaamn .m

meqaoooo .m

muoamucamum
.amb moaned mabamm

eaaomaaoooae .m

«neonaaeo» .m

nausea»

72

homummv oa.mm

“He1Nnv me.mn
Ammuemv am.mm

Amm.emv am.e~
om.oe .me.om

AneuemV mm.am
“amnomv Hm.mm

Ammlomv om.mm

nemnomv we.e~
amm.emv oH.~m

“omummv mm.am
AmmuomV oa.mm
“Haummv ma.em
Ammuwmv n:.mm

oa.mm
Ammuamv m~.mm

Aesouoaa adv
noueasan

amaom

“omuamv mo.mm a Ammnmmv

hamummv am.am a “HaummV

Ammaomv om.m~ a nam10~v

“magnum somv

Ammnmmv
Ammaonv

Ammummv

AeN1mHv
Ammummv

Amuummv
Ammnomv
AcestV
Ammsmmv
Ammummv

oe.mn x an31mnv
Hm.mm x Ameumnv

hm.mm

mm.am
mm.a~

as.mm
m©.mm

H

Amnnauv

Ammomav
Ammummv

Aanaamv
Ammnmmv

ma.mm x Aeenamv
oa.em w “He.mnv
ma.om w Awn1~mv

fie; w 52::

«eschews adv
uoaoaman
Hmauoamavfl

ma.a~

oo.am
mm.m~

nn.on
nm.um

on.nm

:m.m~
mo.mm

on.mm
mm.en
ea.oa
ma.an
Hm.em

Hmdwoamnnm
oumdonm

opmsoamnam

Susana.
epmaonm

3383a.
opmaoam
opmaouaaam

aseaoeoem.
sunflona

camaeamgum
mamaoanpdm
Hmvaoaonmm
eumaoaa
Heeaouenm.
opmaoan
Hmmaoaognm
oamaoum
Hmduonomam
cannomn

sumaoaaamm

Shape Glass

comadoapeu

confiscapoa
camadoaumn

opmadoapon
osaaeeasea
opmaaeaaon
eneaeoanoa
opeasuaoon

onsaeuanou
upmaaoapoa

eamamoapou
mpsaaoasou
manageauou
ovensouuoa
ovmamuapoa
eomHsUuaou

'

Sculpturing

Acesnaaaoov eoaauaaeaomammo manna meadow

3:33 ..H

momma .A

mmmm

3...» 0H and mace?

Hm .oz oeaamv aaqmanamuao .m
0: .oz oewamv adamamdwua> .m
Esmeaam .m

admosxoam adammuamaozm

asHHbaaoe unease

maamaaoumwo manommhm

mamwasb aaaeoaoaaao
maaomaawsM¢m

.ua» seaweeds eanaaaafiso

nauseous nodded

m A
n A
m
m
m
n
n
m
m
m
m

namesake: mnoanamm

newcomm

at.Anthesis

m.ficum 0mm 0 m 0 mm o 0 mm mm mm m

Nb. of Nuclei

.H 0.33...

73

.opmmaoomxon and moan) meoaofia 0H cam 0H mmmamcp mammam moHHoa was omHm coafiemam ocean wane \Mfl

Amauemv aa.ae Aaeummv ma.mm x Awaaemv mm.ma opeaosaeee oneHaOauee m m mameeemeso manomeaafioo
mausoemmmo
Aneuemv mm.mm Ammummv ma.am a Annuamv ao.mm onaaoaaeum oueaeoasos m n .aaeam newness» .2
oztam comasoapoa 0H
omndm \Mfldmaamaw opmaaoapoa m
Aamuemv em.au away me.w~ x mm.:: mneaoaaeeu canadense“ m meanness .2
Humaouomam unease .z N
no.0m Ha.:~ w mm.mm cameosa onmaaoasoa e sweep“: .po mensweaa .2
Amnampm Aswmam m Aoafloazoaoov
swampy mm.me ocoV ma.mm N w.m: eamaonansm oumaaoauen w n semen“: .>o mpaamawa .3
mm.ownom.wm oaoa no NH
opmaaeapma ca m
om.mmlmm.mm camaaoaaoa m m
om.emumo.HN camaaowpoa m neopmaumaaav
Banana: .>o eaaaomwa .2
moummlza.m: oamadoaaou NH
on.weuom.oe eneasoanos ca n
ou.me.mm.m~ onsasoapea m n genus“:
o:.~muqz.oH oneaeuasos e .>o seasoned ensue:
Hmcaoaonmm
Ammummv wmuom homsamv om.am s Ammummv mfi.am oaeaoae oaeaeoapma e m neaeoaeoee .A
1 a a
£33 H news!" a m. h. d .n
a .1 O u e
1.. c N h
Annoyed: a3 Annoyed... a: C m f .... .m
house—man gossamun m. .m. o o A 0300mm
Hmaom Enoamdun m m m m M
as as «u “u

AcosnapseoV moapmaueaonammo manna moaapm .H capes

71

an .mHa .Hx .Hm

mm .mHa .MH .3
a .mHm .>H .Ha
eH .mHa .> .Hm

HH .maa .aH .Hm
a: .mHa .ax .Hm

mm .mHm .N
.Hm “a .mHm .H .Hm
Hm .mHm .N_.Hm

mm .mHm .NH

.Hm “w .mHa .H .Hm
am .mHa .xH .Hm
.mHa .HH .Hm

m .mHm .HHH .Hm
~.:Ha H.Hm
.Maa HHH .Hm
on .mHa .HH .Hm

.oz .wfih m .02 madam

Fl} <m<m4m§§§§

sn1<

<N§4m4¢l§§ <3

Acetocarmine (A)
Acetylation (B)

Preparation:

Presh.(P)

Condition of
or Herbarium (H)

Plant:

mummmmhmm n: ma. mmmmmmmmmm

.oz odaam

omzaoomH aaae aHv eHOHeeam

om: mom nooamm
om: AmHmHaase any smHeaeno
om: a: HHanmnm:
om: mmom omHoaesm
mu mmH mumHna
om: AmmmH ease.eH .eoqev
m6 mmmo anmaaom
om: mm: ooHoasam
mu Aomma pmdwdd aav hmafimm
om: mom assumes:
can own: seen
me mmmmH mouse
om: ammH ooHoaeam
um: one neeaumm

“unease oHeanom eHeHaaeeuHsom.qu

om: ARE as: a: nudism

umz ommn eoHoanmm
5:33 3.93 as tween
om: aHmm oeHoaaem
om: com Hangman:
me name HHosnea

ganmpnem .02 a. .Houooaaoo

goddamn madam soHHom Ho medpmodeaaoon .HH manna

mwamcuoammo manomwhm
wmuaoawmasu .m
madam“: maoocom
sensuous: maonomaw

nHeeanop eHmaooHee
asaoama>hmn asamnmeoooman
mammmdmamo manomaaflaoo
enemas» asaeoaoeHHo

maaouapmdwem
.amb mHHogmHm amaaHsmdmo

ensuaHn .m
epeHHHo eHHanamHm

«cannon amend

mdflomomwamaanaoaom .4
seaflopoaom omomvmmm4

mansosam eoaHo¢

meaoonm

75

me .mHa .5”

.E “9 .maa .H .8 E. a mm om: awn $53.13 $81.32.":
3 GE .2: .8 m4 a 2 8: mmw gaseous: 33...: 832303
n: .mHa .HHHx .8 me a R om: mam $53»: ennui: .8 2283 .2

m4 m we om: RH Seamus: 223mm .2
,2 a; .HHHn .8 m m an em: mam Sensing senescence
NH .mHa .H .E 4 HH 3. me can 3353 $33 3802a «58:
e a; .HHH .8 an m cm 8: i. 8&8 93m?» 333.3:
3 a; .HHx .E m m «a om: SR 3380:
4 HH R me 38 edema «83.25 ..H
m m 2: me 83H page a: swoon
4 m 8H um: 28H seems... a: canon 328.8 ..H
an as; .HHx .H.H me a 3 om: mmmm 3830:
HH .mHa .H .Hm e m mm me an .38 8%.. :H
3 6; .HHx .E m m S om: SH Sake
mm .mHa .HHx .3 m m mm om: RR 3880:
OH .wah .H .Hm d m mm mm wad mommeau ¢.aommeaw adamOfiaoam manoohq
mH .mHa .> .3 m... m an om: 9mm 83.58 5332552 ..H
3 .mHa .> .3 m4 a m cm: mmam scan 32er .3203
S .wHa .2 .E m a HH 8: SN assumes:
m .mrH .H .E < m 8 cm: ENE as: a: Soda ascended ..H
m4 n so um: :32 as: as 58am 53302 ..H
m m 8
4 m H om: mew unapopms eHdsoaHoHaam .A
S a: .Ha .E m .H a 2.28 3538 33.2.8133 n3 39H... ages
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um (m
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e a a m m3
m .... .... a m u

Adednapaoov moamaum manna aoaaom no sodasaaoanoon .HH edges

76

mm .ma .5, .E m u .2 um: m3 35363 «8308 .m
whoawauaauw
m4 u do om: Anoma umpaoumom adv oaoo .uflp magnum «abamm
R .mE .E .8 m4 m R um: “and 832.8
mm .mE .E .5 m4 m ma om: Nu 28am asaflfimh.’ .m
4 m on ma $8 Smfiam 2533 .m
m m mo mon mmoa ago»
4 m 02 omiomS $94 a: 203.5 5882.2 5.85385.“
NH .mE .3 .E m a mu om: Bu 5&3“:
: .E .H .8 ¢ m pm cm: :2: .33. :3 3.33m gamma? 3385
m m NS om: S: gangs: 303QO
4 m 3 me 8? 22:9” ..Ep 235?: .m
quaaaawumb
9 ......E .> .8 m4 m 9... om: :3 295 38> “32%."? .m
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m .mE .5 .E m m um um: mm 3233.
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5 m ca om: 38a #3. a: .38 «33 .:
mm .mE .2: .E m m mm om: H c833
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am .mE .2: .8 m4 m on om: 88a 2%“: a: 038 3.38 3.33:
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on mg a on 33m 3 mm .m .u on «3%. 523."on on a 93828 833m
wmw umm
a a e m a
h m m m m u
Avenueaqoov moacsum awake doanpm mo aoaumaamauoon .HH oaaoa

n .mam .HH .Hm m4 m mm om: “mama hash adv noaoos: adamanofiup mampmogoapa
mm .mfin .HN .Hm m4 m wm om: Amama .oamaom oHuH ..aonmv asflazmpom asshza
n .me .HH .Hm m m mm on: emu Hangman:
< m ma om: mmw umaoahmm omqmvmcmo aduuudma
m m no mon “mama hash gay nonquaaflm
4 m moa om: Ahmad aflau adv amflooo «aflomflsnop .m
gowhuoaon
HN .mfim .HH> .Hm m4 m an om: mm: Hamnnnax .um» adupmsawm .m
m a HoH om: mo: awaken“: manawsamg
m .mam .H .Hm 4 m mo ma aom commode .ua» afiupmsamn .m
on .mam .H> .Hm m m mm om: mmm ooamsuam
4 m cu mo «ca amgaso aflaomfimommhg .m
mH .mam .H> .Hm m m mm om: Ammma “fish adv uupan
4 w as me Noun magmamg mvflamfin mhnowpm
m m um moan Noam.gmsm>oz
4 m mm mm “puma adv naflsm masbhmm .m
m .mdm .HH .Hm m m mm um: mm gauoaaom
H .wdm .H .rm < m #0 ma mum nomaofia a commoau auonHuopmH .m
: .wam .HH .Hm m< m afl om: mmflm vflanono: upmasofluoamm .m
m m mm mon mum gnaw «pamuan
4 m 30H om: nmma ooaosham .ua» modpmaaao «Humaaogsum
mm .wfim .x .Hm m< m on om: naom :Opmaflanmn a.momuom .Hmncpuon anmnspam
mm .wflh .HH> .Hm m m m Acouuaa udaupom afloumuaoiauom adv «anamopamn .m
\I F
A) I\
(a h)
m m n .M flmw
O .1 O r
Mum mrm
.02 .mg a 62 33m r mfl. u u m 02 33m Stanton 62 a “3328 333m
mu“ an» .
a .... u .w m m.

“dosaaanoov nofludam sauna condom mo dawuaocoasoon .HH manna

78

PLATES I-IY

for specimen documentation of pollen grain illustrations. see Table
II. All photomicrographs unless otherwise indicated are at a magnifica-
tion of 1020 X. Each letter (i.e. a. b, etc.) indicates an individual
pollen grain for which there may be two or more photomicrographs made

at different focal levels (except in illustration hfie).

79

Plate I

Nuclei in mature pollen grains photographed in glycerine jelly-

acetccarmine mixture.

1.

9.
10.
11.

12.

13.

1n.

Scutellaria lateriflora. Generative nucleus, 8p in length; tri-
colpate pollen grain, 29p. Faint outline of generative cell mem-
brane is visible (indicated by dashed line on overlay).

Agastache gcroghulariaefolia. Two sperm nuclei; hexacolpate; #60 I.
Egpgga,cataria. Two sperm nuclei; hexacolpate; #60 X

Prunella vulgarig. Two sperm nuclei visible in #b; hexacolpate;
#60 X.

ngigg,pggpgreum. Generative nucleus visible in 5b; tricolpate;
#60 X.

Stacgzs palugtris var. palustris. Generative nucleus. tricolpate;
#60 X.

Monarda gistulogg. Two sperm nuclei. 3p in length; hexacolpate.
#8u x #lp.

Elephilia hirsuta. Two Sperm nuclei; hexacolpate; #60 X.
Clinopodium vulgare. Two sperm nuclei; hexacolpate; #60 X.

chgpgs americanus. Two sperm nuclei; hexacolpate; #60 X.

L. m. Two sperm nuclei; hexacolpate; #60 x.

Eggtg§_arvengig subsp. canadensis. Two sperm nuclei; hexacolpate;
#60 X.

Eggth§_gipgritg cv. Mitcham (irradiated). Two sperm nuclei visible
in 138 which is poly- and pancolpate; b-- lZ-cci pate; c and d-- octa-
colpate; e-- hexacolpate, without protOplasm; all #60 X.

Collinsonia canadensis. Two sperm nuclei; hexacolpate; #60 X.

 

 

 

 

i~ - \
,' " ‘9 Q
*L4L: ‘\e/ '

 

6b

 

   

I ‘ I \\
\ \ /' '\ /:\\ _
'. i \ 'r jg:
\
l,/' ‘ / \ II' \ “I _)
m/ / \"

   

PLATE I

1.

2.

#.

80
Plates II-XV
Acetylated pollen grains (some also chlorinated. i.e. bleached).
Plate II

Teucrium canadense. Tricolpate; ap- oblique equatorial surface,
exine reticulate; a'-- mid-Optical section of same. 35H x 32p;

b—- oblique Optical section of polar view. 3#u x 3#u; b'-— surface
view of same. showing ragged colpi margins; c- polar surface;

c‘-- mid-optical section of same showing thick exine.

éjggaIEeptang. Tricolpate; a-- optical section of polar view, 28p;
b-— mid-Optical section of equatorial view just below surface, 28p x
2#u; c- surface view of equatorial plane showing ragged colpus

margin.

Trichostema braghiatum. Tricolpate; all equatorial views; a- ##u
x 3#u; b—- #Zu x 32p; c-— #fiu x 32p. Second—largest-grained species

of the tricolpate species.

Scutellaria galericulata. Tricolpate; a—— oblique polar view, fine

reticulate; a'- mid-optical section of same, 25p; b and b'--
equatorial views slightly below the surface. 25p 1 21p; c and c'—-
equatorial mid-optical sections, 25u; d-- oblique polar view slightly

below the surface; d'-- mid-optical section of same. 22p 1 21p.

 

PLATE I I

6.

I

6.

7.

8.

81

Plate III

Scutellaria lateriflora. Tricolpate; faintly reticulate; a-

equatorial view, 29p 1 2#u; b—- equatorial view 31p x ZQu.

Marrubium zulgare. Tricolpate, reticulate, muri seemingly larger
than lumina. walls thick; ap- mid-optical section of polar view,
26p; a'- polar surface; b- equatorial midpoptical section. 29p

x 25p; b'-— equatorial surface.

Agastache nepetoides. Hexacolpate, width of exine about qu. width
of lumina l-Zp, reticulate; a—- polar surface; a'- polar mid—optical
section, 35p 1 3#u; b—— equatorial view sligtly below surface; c-—
equatorial surface of collapsed grain; c‘- lower focus of same.

36p 1 21p; d-- ruptured pollen grain, 31p.

Agastache scraphulariaefolia. Hexacolpate, reticulate (grains

chlorinated). thickness of exine ca. 2p. width of lumina 2‘3P3
a-- polar surface; a'- mid—optical section of same, 36p; b——
equatorial view; c—— equatorial view showing 3 narrow colpi, an

exceptionally large grain for this species, #Zu x 31p.

 

” X. \\ '.: 2: .° '
' - K V N 3
8a /

8a' 8b 8c

PLATE III

9.

10.

11.

12.

Plate IV

Qlechoma hederacea. Hexacolpate, reticulate; a-w polar view.
mid-Optical section, 52p; a'-¢ polar surface of same; b--
equatorial surface, colpi margins showing granular nature of the

exine, 52p x #Ou. a-~ is an exceptionally flattened grain.

Nepeta cataria. Hezacolpate. reticulate; a and a'-- oblique polar
sub-surface views, 25u.x 23p; b-— subsurface polar view; b'--
mid-Optical section of same, 26p x 2#u; c- polar view, grain

greatly eXpanded, 3hp,

Dracocephalum parziflorum. Hexacolpate, reticulate. lumina seeming-
ly punctata; a-- equatorial view #Ou x 32p; b-- subsurface polar
view; b'-- mid-optical section of same, greatly expanded. #6u; c—-

oblique polar view.

Prunella gulggris. IHexacolpate, reticulate; a-- polar view; a'--
mid-optical section of same. 38p x 36p; b-~ equatorial surface

showing ragged colpus margin; b'- mid¢0ptical section of same.

39p X 33“-

x?)

I

-....-z '12a' 12b

PLATE IV

3
at
‘3
3.
N"
.‘

eu"'

  

13.

l#.

15.

16.

83

Plate V

msogtggia yirgigisna var. virginiag. Tricolpate. reticulam
pattern very large, 2 to # microns; a-- polar surface. colpi

nearly meeting, broken, perhaps caused by acetylation; a'-- mid-
optical section of same; b, c. and d-- #60 I. largest-grained

species of the tricolpate species.

Leoggus ELM- Tricolpate. fine reticulate; ap- equatorial
surface; a'a— Optical section Of same. 21;: x 1811; b and b'-—
mid-optical section and surface of oblique polar view. 19p; c and

c'-- mid-Optical section and surface of oblique equatorial view.

w garrubiastm. Tricolpate. fine reticulate; a—-- surface
polar view; a'-- subsurface Optical section of same, 26p. 1 21m;

b— polar surface; b'-- mid-Optical section Of same. 21p.

gleonsis tetrahi . Tricolpate. reticulate; am- oblique polar
surface; a'-- mid-Optical section Of same. 35): x #311; b--

equatorial surface. 37p. 1: 319.

 

17.

19.

20.

Plate VI

1.92.1.9; m. Tricolpate, psilate; 8,- polar view, 25].: at 23p;

b. c. d. and e— equatorial views. each showing one colpus.

Iamium m. Tricolpate, fine reticulate. thin exine;
a-- equatorial subsurface; a'-- mid-Optical section of same.

35p; b-- equatorial subsurface. 35p. x 27p.

m hispida. Tricolpate. reticulate; a-- equatorial sub-
surface view. 33}: z 22p; b— polar surface; b'-- mid-Optical
section of same. 2911; c and c'-- surface and mid-Optical views;

d and d'--- mid-Optical and surface equatorial views.

We mssOnglia. Tricolpate. fine reticulate; a-n- equatorial
surface; a’-- mid-Optical section of same. 25):; b- equatorial

subsurface. c-- equatorial subsurface. 30p 1 22p.

f ‘ ”" "‘ "\
\ ' , / ‘
k 1 x > /
‘ si/ ~ \\ J -
17a 17b 17c 17d l7e
‘ A

 

PLATE VI

85

Plate VII

21. Staggyg Elugtris var. homotricha. Tricolpate. fine reticulate;
s.— Oblique polar surface; a'-- mid-optical section of same. 26):;
b—- equatorial surface; b'-~ midpoptical section Of same. 26p x

23p; c- equatorial surface; c'- mid—Optical section of same.

22. Salvia sxlvestrig. Hexacolpate. reticulate; ap- subsurface
section Of polar view. ca. ##u; b—- subsurface section Of

equatorial view; #60 x.

23. film goccinea. Hexacolpate. reticulate, oblate (i.e. the equa-
torial axis is greater than the polar diameter); ap- subsurface
section of polar view; b-- equatorial surface showing 3 colpi.
grain ca. #011 z 5#u; c-- subsurface section Of equatorial view;
#60 X.

2#. Monarda.digzgg. Hexacolpate (occasional octacolpate grains are
found in the same specimen). reticulate. largest-grained species;
a—— Oblique polar surface; a'-- mideptical section of same. 53H x

59p; c-— subsurface section of octacolpate grain. 591.1 x 63p.

 

PLATE VI I

86

Plate VII I

25. M {igtulosb Hexacolpate. reticulate; a-- oblique polar
view; a'-- mid-Optical section of same. #9u; b.- equatorial

surface; b' and b"-- mid-Optical section and subsurface of same.

26. film W var. zilligaulig. Heracolpate, reticulate; a.---

polar view. faint bumm texture in region of the colpi; a'-..
mid-optical section Of same. pollen grain at a slightly Oblique

angle. #6u; b-- equatorial mid-Optical section. #7}: x 3711.

 

26a

25b "

25b"

 

PLATE VI I I

2'7-

28.

29.

30.

87

Plate 11

Blephilia ciliata. Hemacolpate. reticulate; a.— subsurface
Optical section in polar view. #21:; b—- mid-Optical section of
equatorial view. #011 x #5u; c and d- same pollen grains as an-

and. bl“, ”‘60 Xe

W hirsuta. Hexacolpate. reticulate; a»- surface of polar
view, a'-- mid-optical section of same. 36p; 13-— subsurface

section Of equatorial view. 39}: x 30p.

Hedgoma hispida. Hexacolpate. reticulate; a-- surface of polar

view; a'-- mid-Optical section of same. #311.

£533.! m. Hexacolpate. reticulate; a-- surface of Oblique
polar view; a'- mid-Optical section of same. 39):; b-- subsurface

section in equatorial view. 3#u at 32p.

 

PLATE IX

31.

32-

33.

Plate 2

Qalmintha Mm. mtifolia. Hencolpate. reticulate;

a— surface in polar view; b-- subsurface section in polar view;

#Ou x 38u; c-- mid-Optical section Of equatorial view. 3“? x 38):.

Satureja hortggig. Hexacolpats (occasional octacolpate grains
from the same specimen). reticulate; ao- oblique polar view;
a'-- mid-Optical section of same. 3211 x 3111; b- equatorial view;
b'-- mid-Optical section of same. 35;: x 31):; c- octacolpate

grain. polar view; c'—- mid-Optical section of same. 39p. x 38p.

Cligmdgg gum. Hexacolpate. reticulate; a- oblique polar
view; e'- mid-Optical section of same. 36p; b— polar view;

b'-- mid-Optical section of same. 3311 x 3#u.

31b

 

 

33b'

33b

PLATE X

3a.

35-

36.

37.

Plate 11

m gffiginalis. Bexacolpate. reticulate; subsurface section

in equatorial view; pollen grain ruptured.

m W0 Hexacolpate. reticulate; surface section of 95 '
Oblique polar view: a'- mid-optical section of same. 3141; x 30p;
b— equatorial mid-optical section. 39}: x 21411; b'- surface view
Of same slightly collapsed grain; c-- subsurface polar view; d-- E“;

 

 

subsurface equatorial view. 26;: x 231.1.

agnanthemug vugigigum (slide no. #8). Bexacolpate. reticulate;
a-- surface Of Oblique polar view; a'-- mid-Optical section of same.

3#u; b-- surface Of equatorial view. faint bumpy texture in the

region of the colpi; b'-- mid-Optical section of same. #011 x 321.1.

newthem W (slide no. 31). Hexacolpate (occasional
octacolpate grains from the same plant). reticulate; a-— surface

Of polar view; a‘-- mid-optical section Of same. ##u: b-

octacolpate grain. subsurface polar view. #211.

 

 

. l , K“ ‘ ‘fl
4/ "“ ... ' .’

37a W ' 373'

PLATE X I

38.

39-

111.

90

Plate II I

m americanus (slide no. 32). Hexacolpate. reticulate:
a— surface of polar view: a'--- mid-optical section of same,

Bun: b-- mid-optical section in equatorial view. 299. x 29p.

m m. Hemolpate (occasional octacolpate grains from
the same plant). reticulate; a,» subsurface in polar view: a'--
mid-optical section of same. 38p; b-— mid-optical section in

polar view. 38}: 1 Ban: b'-- subsurface of same: o- octacolpate

grain, mid-optical section in oblique polar view. “60 1.

MW (slide no. 1+1). Hexacolpate. reticulate: a—-

polar surface; a'-- lid-Optical section of same (double exposure).
3211; b—- equatorial surface, 31411 x 21m; c and d- equatorial and

polar mid-optical sections. 1160 x.

m uniflorug. Hexacolpate. reticulate; a-u- mid-optical
section in oblique polar view. 29p; a'-- subsurface of same; b

and b'-- two different optical sections, 27): x 2711.

, -'._ 5‘ : rg'flT"

 

 

39b'

39b

39a'

39a

 

 

 

41b'

 

 

PLATE XII

1+2.

143.

M,

91

Plate XIII

m arveglil subsp. W. Hexacolpate, reticulate;

sp- nid-optical section of polar view, 37p: surface of oblique

polar view: b and c-- equatorial view. #60 I.

m pigrifig cv. Kitchen. Hexacolpate. reticulate grain
(indicated by arrow). ca. 20p; remaining grains are pancolpate.

many are ruptured and collapsed; 1460 x.

Mentha M cv. Mitchell (colchicine treated). Octacolpate.
reticulate: a.— surface of oblique polar view; b—- mid-optical

section of polar view. 53}: x 51p.

 

 

 

PLATE XI I I

92

Plate XIV

us. ‘flggtgg,pipgrita cv. Hitchan (irradiated). Hexa-. octa—. dodeca-.
and polycolpate grains. reticulate: ap- optical section. ca. hBu;
b—- subsurface view. ca. 35p; c-- surface view: c'-- midpoptical
section of dodecacolpate grain. ca. 55p; c"- subsurface view of
same; d- polar surface of hexacolpate grain; dl- midpoptical
section of same. ca. 33p: e- microscope field showing variations
in nwnber of colpi. arrangement of colpi. and collapsed or deform-

ed grains (note octacolpate grain left of center); #60 X.

 

45a ’ 45b

 

 

PLATE XIV

93

Plate XV

h6. flgntha pipgrita cv. Mitcham X.§, crispg. Hexacolpate. reticulate;
ca. 28p x 25p (equatorial diameter); #60 I.

#7. Collinsonia canadensis. Hexacolpate. reticulate. exine shows
aburpt thinning near the colpi; ap- oblique polar view; a'-
mid-Optical section.of same. ”Zn; b and b'- portion of chlorinated
grains; c—- equatorial surface: c'—- mid-optical section of same.

“3P X 35)” do .s and f-- 1460 I.

46b
~46c '
'5 ~ ' '\
l
,1:
\C .; ”ad
46s

 

PLATE XV

9“

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It,

 

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