Key to the Quaternary pollen and spores of the Great Lakes region

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Life Sciences
Miscellaneous Publication

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Key to the Quaternary
Pollen and Spores of the
Great Lakes Region

John H. McAndrews, Albert A. Berti
and Geoffrey Norris

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ROYAL ONTARIO MUSEUM
LIFE SCIENCES
MISCELLANEOUS PUBLICATION

JOHN H. McANDREWS,

ALBERT A. BERTI,

GEOFFREY NORRIS

Key to the Quaternary
Pollen and Spores
of the
Great Lakes Region

Publication date: 9 November 1973

ISBN 0-88854-149-X

Suggested citation: Life Sci. Misc. Publ., R. Ont. Mus.

.739

ROYAL ONTARIO MUSEUM
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LIFE SCIENCES EDITORIAL BOARD

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Editor, d. barr

Associate Editor, J. c. barlow
Associate Editor, J. R. tamsitt

john h. mc Andrews is Associate Curator of the Department of Geo-
logy, Royal Ontario Museum and Associate Professor in the Department
of Botany, University of Toronto.

albert a. berti is a National Research Council Fellow at the Depart-
ment of Botany, University of Toronto and Research Associate of the
Department of Geology, Royal Ontario Museum.

Geoffrey norris is Associate Professor in the Department of Geology,
University of Toronto and Research Associate of the Department of
Geology, Royal Ontario Museum.

price: $2.50

©The Royal Ontario Museum, 1973

100 Queen's Park, Toronto, Canada

PRINTED AT THE UNIVERSITY OF TORONTO PRESS

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Contents

Introduction, 1

Materials and methods, 1

Using the key, 3

Key to pollen classes, 4

Key to spore classes, 4

Key to pollen grains, 5
Tetrads, 5
Vesiculate, 5
Poly plicate, 5
Inaperturate, 5
Monoporate, 6
Diporate, 6
Triporate, 6
Stephanoporate, 7
Periporate, 7
Monocolpate, 8
Dicolpate, 8
Tricolpate, 8
Stephanocolpate, 10
Pericolpate, 10
Tricolporate, 10
Stephanocolporate, 12

Key to spores, 13

Alete, 13

Monolete, 13

Trilete, 13
Glossary of morphological terms, 15
Figures, 19

Acknowledgments, 60
Literature cited, 60

Digitized by the Internet Archive

in 2012 with funding from

Royal Ontario Museum

http://archive.org/details/keytoquaternarypOOmcan

Key to the Quaternary Pollen and Spores
of the Great Lakes Region

Introduction

In recent years the method of fossil pollen and spore analysis (Faegri and
Iversen, 1964) has become important in both paleoecological recon-
structions of vegetation and climate and in biostratigraphic correlation of
sediments. Effective reconstructions and correlations depend upon accurate
fossil identifications.

Scattered information is available on pollen and spore morphology, but
relatively little has been organized in a form that can be used for fossil identi-
fication in the Great Lakes region. These aids include Sears (1930), "Com-
mon fossil pollen of the Erie Basin," which contains a key to only 33 taxa,
illustrated by drawings. Presently, pollen analysts commonly use the unil-
lustrated key to the northwest European pollen types (Faegri and Iversen,
1964), but it is not wholly applicable to our region nor does it include
spores. Likewise, the key of Kapp (1969), "How to know pollen and
spores," covers only the United States, is illustrated by drawings and does
not focus on fossil identification. Another geographically limited but useful
aid is Richard's (1970) "Atlas pollinique des arbres et de quelques arbustes
indigenes du Quebec," which contains excellent photographs and descriptions
but omits herb pollen and lacks a general key. The recent atlas of tree pollen
by Adams and Morton (1972) has similar limitations, but its scanning
electron micrographs are excellent for sculptural detail.

Fossil pollen assemblages are composed primarily of pollen of wind-
pollinated species, especially trees, and to a lesser extent shrubs and herbs.
Except for aquatic plants, pollen of insect-pollinated species are rare. Our
photo-illustrated key is designed for the identification of fossil pollen and
spores (microspores and homospores of pteridophytes and Sphagnum) com-
monly found in Quaternary deposits of the states and the province surround-
ing the Great Lakes. The key is most applicable in the middle Great Lakes
region, especially southern Ontario and Michigan, but is also useful in such
peripheral areas as the Atlantic Coast, and the subarctic and prairie regions
as shown in Fig. 1 .

Materials and Methods

The 144 entries in the key were selected from our analyses of sediments of
late Pleistocene (Sangamonian, Wisconsinan) and Holocene age and on
fossils reported by others. The majority of taxa (76) are rare, that is, they
usually represent less than 1 % of a total fossil spectrum. Aquatic taxa are
also rare, but some may be locally abundant. Only 46 taxa are common
enough to occur consistently with values greater than 1 % . Not all grow in
the Great Lakes region today, but most grew in this region sometime in the

Fig. 1 — Zonal vegetation map of the Great Lakes region (modified from Braun,
1950, and Rowe, 1959). The key is most applicable in the conifer-hardwood
forest zone.

Quaternary. Fossil Ephedra and Sarcobatus are exceptions, in that they
appear to have been blown in from beyond our region (Maher, 1964).

The key is in two parts, one for the sixteen classes of pollen and the second
for the three classes of spores. In our fossil flora, spores (Figs. 16-21) tend
to be morphologically simpler than pollen (Figs. 2-15). Spores have a slit-
shaped aperture (laesura) that lacks a membrane. They may have a single
laesura (monolete, Fig. 16c) or three triradiate laesurae (trilete, Fig. 18c).
The only exception treated here is the alete Equisetum (Fig. 16a).

In pollen grains apertures may be absent (Fig. 5e), simple (elongate
colpi, Fig. 10k, or circular pores, Fig. 5m), or compound (colpi with pores,
Fig. 13n). Usually multiple apertures are present, and in fossils the aper-
tures often retain a membrane (Figs. 9c, s; 10l). Spores commonly have a
relatively thick, structureless wall, and many taxa possess a loose-fitting sac
or perine that is often lost from fossil spores. Pollen have walls of varying
thickness and complexity and may have easily distinguished columellae and
a tectum (Fig. 13e); pollen never have a perine. The size of spores and
pollen is generally similar except that the minimum size of pteridophyte
spores is about 25 jum, whereas many pollen are smaller. Fungal spores (not
considered in the key) can be confused with pollen and spores of vascular
plants but are distinguishable by their small size (5-20 fim), general lack
of apertures, and simple wall structure.

We attempted to keep terminology to a minimum and have defined terms
in the glossary (p. 15). Spore terminology follows Harris (1955), and
pollen terminology is based on Faegri and Iversen (1964). More exhaustive
and detailed treatments of pollen and spore morphology are found in texts
such as Erdtman (1957, 1965, 1966), Erdtman and Sorsa (1971), Kremp
( 1 965 ) , and Tschudy and Scott ( 1 969 ) .

The key is illustrated with reference pollen and spores collected from
herbarium sheets or living plants, treated according to the method of Faegri
and Iversen (1964) with KOH and acetolysis solution, and mounted in sili-
cone oil. Due to the inflated and uncompressed nature of the pollen and the

shallow depth of focus of high magnification microscope objectives, it was
not possible to illustrate all morphologic features in one photograph. In
general, spores and pollen are best illustrated either in "optical section" to
show features of the internal wall structure (Fig. 6f) or in surface view to
show details of the external sculpture pattern (Fig. 6a). Occasionally, both
the optical section and the surface pattern of the exine are illustrated in the
same photograph (Fig. 6e). Only the most useful diagnostic features are
illustrated in this work allowing the user to "key-out" an unknown grain
with the aid of a light microscope.

Both polar and equatorial views have been used to illustrate the various
morphologic features in this key; these views are indicated in the Figure
captions by the abbreviations "pv" and "ev" respectively. Photograph mag-
nifications are at XI 000 except for the vesiculate genera Pinus, Picea, and
Abies (Fig. 2) which are at X500.

Pollen and spore measurements in the Figure captions were made with
an ocular micrometer on 10 undistorted grains from a single slide. Measure-
ments are of the greatest dimension excluding sculptural projections and the
perine. Measurements of vesiculate grains are of the body in equatorial view.
The average size is given in fim with the range in parentheses.

In addition to reference slides, the most effective use of the key requires
a high quality microscope, and good fossil preparations. Fossil pollen are
often folded, broken or shrunk, and size measurements of reference pollen
must be used with caution. Apertures are often obscured, and we recom-
mend the use of a liquid mounting medium (silicone oil or glycerine) so that
grains can be rolled to reveal key features.

In some cases the pollen or spore class to which a grain belongs (e.g.
Equisetum, Ambrosia, Tilia or Cyperaceae) is either uncertain or difficult
to determine. This problem has been overcome by entering the pollen or
spore in more than one class. In addition, because the key is basically for
identification rather than taxonomy, Equisetum and Selaginella selaginoides
are entered in both the pollen and the spore classes.

Using the Key

Use of the key involves several steps. First the unknown is classified as a
pollen grain or spore. Then it must be keyed to class with either the "Key
to pollen classes" or the "Key to spore classes." The third step is to identify
the unknown within the proper class in the "Key to pollen grains" or "Key to
spores." Finally, the unknown specimen should be compared with the photo-
graphs and with reference material, to confirm or reject the identification.

Nomenclature follows the 8th edition of Gray's Manual of Botany (Fern-
aid, 1950). Species names and authors that do not appear in that manual
are Sarcobatus vermiculatus (Hook.) Torr., Iva axillaris Pursh, Ephedra
aspera Engelm., E. coryi Reed, E. nevadensis Wats., E. torreyana Wats., E.
trijurca Torr., and E. viridis Coville.

Key to Pollen Classes

A. Grains in groups of 4 1 Tetrads p. 5

AA. Grain single (monads)

B. Apertures absent (or indistinct)

C. With two bladders 2 Vesiculate p. 5

CC. Without bladders

D. With ridges separated by folds 3 Polyplicate p. 5

DD. Without ridges and folds 4 Inaperturate p. 5

BB. Apertures present

E. Apertures simple, i.e. either pores or colpi
F. With pores

1 pore 5 Monoporate p. 6

2 pores 6 Diporate p. 6

3 pores 7 Triporate p. 6

More than 3 pores equatorially arranged

8 Stephanoporate p. 7

More than 3 pores, some or all not equatorial

9 Periporate p. 7

FF. With colpi

1 colpus 10 Monocolpate p. 8

2 colpi 11 Dicolpate p. 8

3 meridional colpi 12 Tricolpate p. 8

More than 3 meridional colpi 13 Stephanocolpate p. 10

More than 3 colpi, some or all not meridional

14 Pericolpate p. 1 0

EE. Apertures compound, i.e. pores in colpi

3 colpi meridionally arranged 15 Tricolporate p. 1 0

More than 3 colpi meridionally arranged

16 Stephanocolporate p. 12

Key to Spore Classes

A. Spore without aperture 1 Alete p. 13

AA. Spore with slit-shaped aperture (laesura)

B. Spore with single laesura 2 Monolete p. 13

BB. Spore with triradiate laesurae 3 Trilete p. 13

Key to Pollen Grains

1. Tetrads

A. Reticulate, tetrads usually rhombohedral, each grain monoporate.

Typhaceae Typha latijolia 2a, b

AA. Not reticulate, tetrahedral tetrads
B. Echinate

C. Spines 5-8 fim long and spaced 5-10 /xm apart. Rare to locally

abundant. Selaginellaceae Selaginella selaginoides 2c; 19e

CC. Spines 2-3 /xm long and more closely spaced. Rare. Drosera-

ceae Drosera 2d

BB. Psilate or scabrate, each grain tricolpate or tricolporate

Ericaceae 2e- j

2. Vesiculate Bladders with internal reticulum. Pinaceae

A. Bladders hemispherical (not constricted at attachment with body of
grain) , cap of body (proximal surface) of even thickness, body 50-80
/xm broad. Reticulum of bladder finer than in Pinus (Fig. 3d) or

Abies (Fig. 3f) Picea 3a-d

AA. Bladders more than hemispherical (constricted at attachment to
body), cap of uneven thickness
B. Body of grain 60-90 /urn broad, bladder 45-70 /xm broad, cap
thick (5 /xm) but often appearing thinnest in centre Abies 3e, f
B.B. Body 40-55 jam broad, bladder 24-40 /xm broad, cap thinner (2
/xm) and of even thickness except along margin. Pinus. Species
separation by size statistics is not reliable (Whitehead, 1964).
C. Distal verrucae present and about 1 fim broad. (Smaller and
more scattered verrucae occur on Picea). Section Haploxylon

Pinus strobus 3k

CC. Distal verrucae absent. Section Diploxylon (P. banksiana, P.

resinosa and P. rigida) Pinus banksiana type 3g-j

3. Polyp! icate (with meridional folds), prolate, grain c. 25 X 45-65 /xm.
Ephedraceae

A. Grain with c. 15 straight folds. Includes among others E. aspera and

E. trijurca. Rare Ephedra torreyana type 4a

AA. Grain with c. 6 branched folds. Includes among others E. coryi and
E. viridis. Rare Ephedra nevadensis type 4b

4. Inaperturate

A. Grain banana-shaped, with an imperfect reticulum, grain c. 35 X 70

xim. Rare. Zosteraceae Ruppia 4c

AA. Grain spheroidal (or pear-shaped in Cyperaceae) but often dis-
torted by splits and folds
B. Grain 55-85 /xm. Pinaceae

C. Psilate Larix 4d

CC. With irregular, hollow vervucae Tsuga 4e

BB. Grain 20-45 /xm except some Cyperaceae as large as 60 /xm

5

D. Gemmate with gemmae distinct but scattered, fossil grains often
angularly broken. Cupressineae Thuja or Juniperus 5a-c

DD. Psilate, scabrate or reticulate
E. Psilate or scabrate-verrucate

F. Psilate, often with a wrinkled perine. Rare to locally abun-
dant. Equisetaceae Equisetum 5d; 16a

FF. Scabrate-verrucate

G. Spheroidal, no poroids and often folded or flattened.

Rare to locally abundant. Salicaceae Populus 5e

GG. Pear-shaped with 3 lateral poroids and 1 poroid on

broad end. Most fossil grains are probably aquatic

species of Carex, Scirpus, Eleocharis or Cyperus. See

Faegri and Iverson (1964) for more detailed indenti-

fication Cyperaceae 5f, g; 7l, m

EE. Reticulate

H. Reticulum broken (muri incomplete). Includes the other
members of the subgenus Coleogeton, i.e. P. filiformis

and P. vaginatus. Zosteraceae

Potamogeton pectinatus type 5h

HH. Reticulum unbroken. Zosteraceae or Juncaginaceae ....

Potamogeton natans

type (subgenus Eupotamogeton) or Triglochin 5i, J

5. Monoporate

A. Pore with distinct annulus, psilate or scabrate, grain c. 20-50 jiim

(larger in cereals, e.g. Zea Fig. 5m) Gramineae 5k-m

AA. Pore without annulus, grain 1 8-30 ^m

B. Reticulate. Sparganiaceae or Typhaceae

Sparganium or Typha angustifolia 6a, b

BB. Echinate. Rare. Lemnaceae Lemna 6c

6. Diporate

A. Psilate, pore circular and annulate, exine thin, grain spheroidal, 1 8-20

fxm. Rare. Moraceae Moms 6d; 7d

AA. Psilate, but with protruding pores, exine thicker

rare grains of Corylaceae

7. Triporate Grains of Ostrya and Carpinus occasionally have 4 pores.
A. Pores distinctly protruding, psilate

B. Grain 45-80 fxm. Rare. Onagraceae Epilobium 6e

BB. Grain 16-35 fxm

C. Inner surface of pore margin rough. Myricaceae

Comptonia or Myrica 6f
CC. Inner surface of pore margin smooth. Corylaceae

D. With separation of endexine at base of pore, exine thick

Betula 6g-i

DD. Without separation of endexine, exine thinner

Ostrya or Carpinus 6j-k

A A. Pores scarcely protruding

E. Reticulate, endexine conspicuously thickened around pore, oblate.

Tiliaceae Tilia 6l; 14m

EE. Psilate-scabrate
F. Annulate, spheroidal

G. Pore sunken, i.e. tectum swells outward at pore and bends in-
ward at inner margin, psilate. Rare. Cannabinaceae

Humulus 6m

GG. Pore not sunken, scabrate. Rare. Ulmaceae .... Celtis 6n, o
FF. Not annulate, suboblate-oblate

H. Grain 40-55 ju,m, slightly heteropolar. Juglandaceae

Carya 7a

HH. Grain < 40 jum

I. Grain 20-30 ^m, pores equatorial and slightly protruding,
exine thick. Corylaceae Corylus 7b

II. Grain 12-20 /xm, psilate, pores not protruding, exine thin
(0.5 /mi). (Exceptional triporate Moms, Fig. 7d, will key
out here.) Rare. Urticaceae Urtica 7c

8. Stephanoporate

A. Arci (internal thickenings) connecting the 4 or 5 (rarely 3 or 6) pro-
truding pores. Corylaceae Alnus 7e-g

AA. Without arci

B. Rugulate-reticulate, 4-6 non-protruding and non-annulate pores,

oblate. Ulmaceae Ulmus 7h. i

BB. Psilate-scabrate, 4 (3-6) non-protruding but strongly annulate
pores. (Ostrya and Carpinus occasionally have 4 protruding
pores). Haloragaceae

C. Annulus 2-3 fim high, pores asymmetrically situated. Rare

Myriophyllum alterniflorum 7 J

CC. Annulus 1-2 fxm high, pores symmetrically situated. Includes

among others M. farwellii, M. spicatum, M. verticillatum

Myriophyllum exalbescens type 7k

9. Periporate

A. Heteropolar (pores mostly concentrated on one hemisphere)

B. Pear-shaped, scabrate, with 3 (4-6) lateral poroids (thin areas)

and 1 poroid on broad end Cyperaceae 5f, g; 7l, m

BB. Oblate, psilate, 7-18 annulate pores. Juglandaceae

C. 7-11 pores Juglans cinerea 7n, o

CC. 12-18 pores Juglans nigra 7p

AA. Not heteropolar, spheroidal

D. Echinate with spines 2 pan long and regularly spaced, pores indis-
tinct. Alismataceae Sagittaria 7q

DD. Not echinate

E. Reticulate, grain 30-50 />im

F. Reticulum coarse (lumina 5-10 /on), pores at bottom of
lumina. Includes among others P. hydropiper, P. hydropiper-
oides, P. pensylvanicum, P. punctatum and P. sagittatum.
Rare. Polygonaceae Polygonum lapathijolium type 8a

FF. Reticulum finer (lumina 1 pan). Hamamelidaceae

Liquidambar 8b

EE. Psilate-scabrate or verrucate

G. Distinctly annulate, grain 15-30 pan

H. More than 16 pores, exine thick. Rare

Caryophyllaceae 8c

HH. Fewer than 16 pores

I. About 14 pores, psilate. Rare. Chenopodiaceae

Sarcobatus 8d

II. 8-14 pores, scabrate. Mostly restricted to sediments of

the past century. Plantaginaceae

Plantago lanceolata 8e

GG. Not distinctly annulate

J. More than 30 pores. Includes Amaranthaceae and Cheno-
podiaceae except Sarcobatus. For method of distinguishing
species by pore number see McAndrews and Swanson

( 1 967 ) Chenopodiineae p.p. 8f, g

JJ. Fewer than 30 pores

K. Pore margin distinct, exine very thick, grain > 30 pan.

Rare Caryophyllaceae

KK. Pore margin indistinct, grain 15-30 p,m

L. Psilate, 6-12 pores. Rare. Ranunculaceae

Thalictrum 8h

LL. Verrucate, 4-11 pores. Mostly restricted to sedi-
ments of this century. Includes among others P.
virginica and P. pusilla. See also Bassett and Cromp-
ton (1968). Rare Plantago major type 8i

10. Monocolpate Nymphaeaceae

A. Colpus circular, baculate Nymphaea 8j, k

AA. Colpus linear

B. Echinate, spines 5-6 ^m long Nuphar 8l

BB. Psilate-scabrate. Rare Brasenia 8m

11. Dicolpate, scabrate-verrucate, 50-72 pan. Rare.

Pontederiaceae Heteranthera 8n

12. Tricolpate

A. Sculpturing elements elongated parallel to surface
B. Striate or rugulate
C. Striate

D. Colpus without membrane, > 30 pan. For detailed treatment
of Acer pollen see Helmich (1963).

Aceraceae Acer rubrum 9a, b

DD. Colpus with verrucate membrane, < 30 ^m. Rare. Saxi-

f ragaceae Saxifraga oppositifolia 9c, D

CC. Rugulate

E. Grain > 30 yu,m, delicately rugulate-reticulate, with colpus
membrane. Aceraceae Acer saccharinum 9e, f

EE. Grain < 30 pan

F. With colpus membrane, endexine thin. Rare. Rosaceae

Dry as 9g-i

FF. Colpus membrane absent, endexine thick. Rare. Acer-

aceae Acer negundo 9j-l

BB. Reticulate

G. Colpus with psilate-scabrate margin, polar area small
H. Subprolate-prolate, without operculum. Salicaceae

I. Reticulum unbroken (muri complete) Salix p.p. 9m-p

II. Reticulum broken. Rare Salix herbacea 9q, r
HH. Prolate-perprolate, grain c. 25 x 45 ixm, with operculum,

reticulum fine, Rare. Leguminosae

Petalostemum purpureum 9s

GG. Margin absent, i.e. reticulum of exine extends to edge of
colpus
J. Tectate, polar area small

K. Exine thick (2.2 yum), lumina 1.5 pcm, colpus membrane

absent. Aceraceae Acer saccharwn 9t, u

KK. Exine thinner, lumina smaller. Includes among others
Stachys and Scutellaria. Rare Labiatae p.p. 10a, b
JJ. Intectate

L. Polar area small, exine thick (2.5 ^m), lumina 1.5 ju.m,

Rare Cruciferae 10c, D

LL. Polar area medium to large, exine thinner (< 1.5 ju,m),
lumina smaller (< 1.5 /xm)

M. Colpus membrane absent, 20-25 /xm. Oleaceae

Fraxinus nigra or F. quadrangulata 10e-g

MM. Colpus with scabrate membrane, spherical, grain 16-

20 fim. Platanaceae Platanus 10h

AA. Sculpturing elements absent or isodiametric, or elongated perpen-
dicular to surface (not striate, rugulate or reticulate)
N. Scabrate, spheroidal-subprolate

O. Irregularly scabrate, no colpus membrane, large polar area, grain

20-35 /Lim. Fagaceae Quercus 10i-k

OO. Very regularly scabrate, colpus membrane scabrate, small polar
area, grain 22-26 />tm. Rare. Ranunculaceae Caltha 10l
NN. Not scabrate

P. Echinate, spines short (c. 1 /xm), polar area large, spherical,
grain 18-23 fxm. Compositae Ambrosia 10m; 12m, n

PP. Verrucate or clavate

Q. Verrucate, grain 26-32 /xm. Rare. Includes species of the

section Batrachium. Ranunculaceae

Ranunculus flabellaris type 10n

QQ. Clavate

R. Colpus with margin of smaller clavae, prolate, grain 22-32

fxm. Rare. Aquifoliaceae Ilex or Nemopanthus lOo, P

RR. Colpus without special margin, clavae-verrucae of various

sizes, subprolate, grain 30-36 fim. Rare. Rosaceae

Rubus chamaemorus 1 0q, r

13. Stephanocolpate

A. Psilate-scabrate

B. Small polar area, 6-10 colpi. Rare. Rubiaceae Galium lOs-u
BB. Large polar area, 4 (5-6) colpi each with a broad membrane.

Rare. Hippuridaceae Hippuris 11a

AA. Reticulate or echinate

C. Reticulate

D. 6 colpi, small polar area. Rare. Includes among others Lycopus,

Mentha, Monarda, and Prunella Labiatae p.p. 11b, c

DD. 4 colpi, large polar area

E. Subprolate, reticulum fine, grain 20-24 /u,m. Oleaceae

Fraxinus pennsylvanica or F. americana 1 1 D-F

EE. Grain with shape of rectangular prism, reticulum medium-
coarse, 4 short colpi, grain c. 32 ^m. Rare. Balsaminaceae

Impatiens 1 1g, H

CC. Echinate, 3 long and 3 short colpi. Rare. Loranthaceae

Arceuthobium 111, J

14. Pericolpate

A. Reticulate, with clavae at bottom of lumina. Rare. Polygonaceae

Polygonum amphibium or P. coccineum 1 1 K, L

A A. Echinate. Rare. Portulacaceae Portulaca oleracea 11m

15. Tricolporate This is the largest class and somewhat difficult because
pores are often obscure. Reference slides are especially important.

A. Echinate. Compositae

B. Exine structure consisting of lacunae separated by echinate ridges

arranged in a geometrical pattern. Rare

subfamily Liguliflorae 1 2a-c

BB. Exine without lacunae or ridges. Subfamily Tubuliflorae

C. Spines > 1.5 fxm polar area small. Includes all genera except the

Ambrosia group and Artemisia Tubuliflorae p.p. 12d-g

CC. Spines short (< 1.5 yu,m), pores indistinct, grains 14-26 fim.

Ambrosia group

D. Columellae of tectum distinct, colpi indistinct, polar area large

E. Exine thick, spines c. 1 fxm long. Rare to locally abundant
Iva ciliata 1 2h, i

EE. Exine thinner, spines shorter. Rare Xanthium 12j

DD. Columellae indistinct

F. Colpi long and distinct, small polar area. Rare to locally
abundant Iva xanthijolia 1 2k, l

FF. Colpi short and indistinct, large polar area. The pre-
dominantly western Iva axilaris is identical

Ambrosia 10m; 12m, n

AA. Not echinate
G. Psilate-scabrate

H. Grain < 16 /u,m, prolate, pores equatorially elongated. Fagaceae
Castanea 1 2o, P

10

HH. Grain > 18 fim

I. Oblate, triangular in polar view, pores protruding. Elaeagna-
ceae

J. Colpi short (c. 8 fim). Rare Elaeagnus 13a

JJ. Colpi long (c. 20 fim). Rare. Shepherdia argentea 13b

II. Spheroidal to prolate or perprolate
K. Spheroidal to prolate

L. Equatorial thickening of endexine. Includes among others

P. cristatwn and P. achoreum. Rare. Polygonaceae

Polygonum aviculare type 1 3c, D

LL. No equatorial thickening
M. Grains spheroidal

N. Exine thick (3-4 /xm) but thinning toward colpus,
columellae distinct, micro-echinate, grain 20-30

fim. Compositae Artemisia 13e, f

NN. Exine thinner and of uniform thickness

O. Pore protruding slightly, circular and annulate,

grain 30-34 fim. Rare. Nyssaceae

Nyssa 1 3g-i

00. Pore not protruding

P. Grain 35-43 fim, scabrate, pore often elon-
gated along meridian and often not very dis-
tinct. Fagaceae Fagus 1 3 J, k

PP. Grain 25-30 fim, pore circular and distinct.

Rare to locally abundant. Polygonaceae

Rumex 13l, m

MM. Grains prolate

Q. Pore circular and slightly annulate, grain 24 X 35

fim. Rare to locally abundant. Elaeagnaceae

Shepherdia canadensis 13n, o

QQ. Pore not circular

R. Pore not elongated but characterized by a con-
striction of the colpus. Cornaceae

S. Grain < 25 fim, psilate. Rare

Cornus canadensis 13p, Q

SS. Grain > 35 fim. Rare. Includes among others
C. racemosa, C. alternijolia and C. rugosa

Cornus stolonijera type 14a-c

RR. Pore elongated equatorially, exine thicker at
poles than at equator, grain c. 35 X 50 /mi.

Rare. Polygonaceae

Polygonum viviparum 14d, e

KK. Perprolate, exine of varying thickness, pore elongated

equatorially. Rare Umbelliferae 14f-h

GG. Striate or reticulate
T. Striate

11

U. Striae very pronounced and parallel with meridional orienta-
tion, transverse colpus absent. Rare. Gentianaceae

Menyanthes 1 4i, J

UU. Striae finer, pore indistinct, transverse colpus present or

absent. Includes among others Potentilla and Prunus

Rosaceae p.p. 1 4k, l

TT. Reticulate

V. Oblate, endexine conspicuously thickened around pore, colpi

short and indistinct, ca. 36 ixm. Tiliaceae Tilia 6l; 14m

VV. Spherical to prolate, endexine not conspicuously thickened

around pore, colpi longer and distinct

W. Pore large (about Vi diameter of spheroidal grain), colpus

with an operculum, grain 20-22 \xm. Rare. Leguminosae

Amorpha 1 4n, o

WW. Pore smaller

X. Colpus margin psilate, reticulum fine, prolate, pore not

pronounced. Salicaceae Salix p.p. 14p, q

XX. No distinct colpus margin

Y. Pore an equatorial constriction of the colpus, grain
15-20 fim. Rare. Caprifoliaceae Sambucus 15a
YY. Pore circular and annulate

Z. Pore wider than colpus, micro-reticulate. Rare.

Vitaceae Vitis 1 5b, c

ZZ. Pore not as wide as colpus, reticulum coarser.

Rare to locally abundant. Rubiaceae

Cephalanthus 1 5d, e

16. Stephanocolporate

A. With c. 15 colpi, pores fused together around the equator. Rare.

Lentibulariaceae Utricularia 1 5f, G

AA. With 6 colpi that form 3 pairs, each pair of colpi within an equa-

torially elongated pore. Rare. Rosaceae

Sanguisorba canadensis 1 5h, I

12

Key to Spores

1 . Alete Perine wrinkled and grain often broken. Rare to locally abundant.
Equisetaceae Equisetum 5d; 1 6a

2. Monolete

A. Perine absent. Polypodiaceae

B. Psilate. Includes among others Asplenium trichomanes, Cystopteris

fragilis, Dryopteris spinulosa and Onoclea sensibilis

Polypodiaceae p.p. 16b, c

BB. Verrucate

C. Grain < 42 tun, verrucae small and scattered. Rare

Athyrium filix-femina 16d

CC. Grain > 44 /urn, verrucae larger and dense. Rare

Poly podium virginianum 1 6e

AA. Perine present

D. Perine psilate-scabrate

E. Perine ragged in the region of the laesura. Rare to locally
abundant. Isoetaceae Isoetes 16f

EE. Not ragged in the region of the laesura. Rare. Polypodiaceae

Pteretis pensylvanica 1 6g

DD. Perine variously sculptured, not psilate-scabrate

F. Perine not echinate

G. Perine reticulate and loose fitting. Rare. Polypodiaceae

Woodsia ilvensis 1 6h

GG. Perine variously ridged and folded. Includes among others
Athryiwn thelypteroides, Dryopteris thelypteris and D.

goldiana Polypodiaceae p.p. 16i, J

FF. Perine echinate. Polypodiaceae

H. Echinae projecting from a tight-fitting perine

I. Echinae sharp and up to 6 /xm long. Rare

Cystopteris fragilis 16l

II. Echinae blunt and 2-4 /un long. Rare

Cystopteris bulbijera 1 6k

HH. Perine loose-fitting, echinae on ridges

J. Echinae c. 1 /xm long. Rare Dryopteris spinulosa 1 7 a

JJ. Echinae 2-4 /un long and very irregular. Rare

Camptosorus or Polystichum lonchitis 17b, c

3. Trilete

A. With perine

B. Perine 70-85 /xm with folds and thickenings forming an imperfect

reticulum. Rare. Polypodiaceae Cheilanthes 17d, h

BB. Perine 30-55 tun

C. Perine large (40-55 /un) loose-fitting and densely verrucate.

Rare to locally abundant. Selaginellaceae

Selaginella rupestris 17e

13

CC. Perine smaller (30-55 fim) tight-fitting and densely verrucate.

Polypodiaceae Pteridium aquilinum 1 7f, g

AA. Without perine

D. With negative sculptural elements or only psilate-scabrate
E. Psilate or scabrate. Polypodiaceae

F. Spore > 45 /xm, psilate. Rare Cheilanthes 17d, h

FF. Spore < 45 (xm, scabrate

G. Laesurae with lips, exine > 1.5 fxm thick. Rare

Adiantum or Dennstaedtia 1 8a, b

GG. Laesurae simple, exine < 1 .5 fim thick

Pteridium aquilinum 1 8c, D

EE. Foveolate or fossulate
H. Foveolate. Lycopodiaceae

I. Spore < 25 /xm, sides of spore concave in polar view. Rare
Lycopodium lucidulum 18e, f

II. Spore > 30 /urn and nearly triangular in polar view. Rare
Lycopodium selago 1 8g, h

HH. Fossulate. Rare. Ophioglossaceae

Botrychium multifidum or B. dissectum 1 8l, J-L

DD. With positive sculptural elements, not scabrate

J. Sculptural elements radially elongated (baculate or echinate)
K. Baculae of varying size, spore spheroidal. Osmundaceae
L. Baculae about twice as long as broad and not fused. Rare

Osmunda cinnamomea 1 9a

LL. Paculae broader and some fused to form short ridges.

Rare Osmunda regalis or O. clay toniana 19b-d

KK. Echinate with spines pointed or truncate and 5-8 yu,m long,

spores often in tetrads. Rare to locally abundant. Selaginel-

laceae Selaginella selaginoides 2c, 19e

JJ. Sculptural elements elongated parallel to surface
M. Rugulate or reticulate

N. Rugulate, distal face with zig-zag ridges and proximal face

verrucate. Rare. Lycopodiaceae

Lycopodium inundatum 1 9f, g

NN. Reticulate (at least on distal surface)

O. Muri with a broad base (tapering in cross-section) . Rare.

Ophioglossaceae Ophioglossum 20a, b

OO. Muri with vertical walls. Lycopodiaceae

P. Reticulum extending well onto proximal surface

Q. Nodes at angles of muri. Rare

Lycopodium clavatum 20c, D

QQ. Without nodes. Rare

Lycopodium

complanatum or L. tristachyum 20e-h
PP. Reticulum not extending well onto proximal surface
R. Laesurae about two-thirds spore radius, lumina

small (up to 7 pum). Rare

Lycopodium obscurum 20l, J

14

RR. Laesurae reaching equator, lumina larger (up to

1 0 pm ) . Rare

Lycopodium annotinum 21a, b

MM. Verrucate

S. Laesurae without lips, verrucae up to 6 ftm broad, spore

30-36 )U,m. Rare. Ophioglossaceae

Botrychium virginianum 21 c, D

SS. Laesurae with lips

T. Spore < 35 yu,m. Sphagnaceae Sphagnum 21e, f
TT. Spore > 35 pan. Rare. Includes among others Botry-
chium lunaria and B. lanceolatum. Ophioglossaceae
Botrychium simplex type 21G-K

Glossary of Morphological Terms

Terms are mostly defined in their singular noun form; adjectival forms are indi-
cated with the suffix "ate".

AMB: outline of a pollen grain or spore in polar view (Betula Fig. 6g-i; Adi-

antum Fig. 18a).
ANNULUS: area surrounding a pore characterized by a thickening of the pollen

wall (Caryophyllaceae Fig. 8c, d; Gramineae Fig. 5k-m).
APERTURE: a thinning or break in the pollen or spore wall that functions as a

site for the emergence of the pollen tube or gametophyte.
ARCUS: elongate, narrow, curved thickening (arc) of the exine not parallel to

the edges of the apertures (Alnus Fig. 7e, f).
BACULUM : sculptural element with radial projection isodiametric, at least one

dimension > 1 /mi, height > width, sides parallel and top not pointed

(Nymphaea Fig. 8 J, k).
CAP: the thickened proximal surface of the body of vesiculate grains (Picea

Fig. 3a, c).
CLAVA: sculptural element with radial projection isodiametric, at least one

dimension > 1 /mi, height > width, base constricted and top rounded (Nemo-
pant hus Fig. 10p).
COLPUS: an aperture, arbitrarily defined as one with a length-breadth ratio

^ 2.0, the ends more or less acute (Quercus Fig. 10k).
COLUMELLA: radial rod of the exine separating a basal layer from the outer

tectum (Artemisia Fig. 13e).
DISTAL: that part of a pollen or spore which faces outward in its tetrad, repre-
senting for example, in monoporate and monocolpate pollen, the area around

the aperture.
ECHINUS: sculptural element with radial projection isodiametric, at least one

dimension > 1 /tm, height > width, and pointed (Agoseris Fig. 12a-c: Helian-

thus Fig. 12d).
EKTEXINE: the outer part of the exine, consisting of a basal layer, radial

columellae and an outer tectum.

15

ENDEXINE: the innermost part of the exine, a relatively homogeneous zone

below the ektexine.
EQUATOR: the imaginary line midway between the two poles that divides the

grain into two polar hemispheres.
EXINE: the highly resistant wall surrounding the cytoplasm of pollen and

spores, divisible into two main layers, the inner endexine and the outer

ektexine.
FOSSULATE: sculptural type with surface of exine grooved (Botrychium multi-

fidum (Fig. 18i, J).
FOVEOLATE: sculptural type with surface of exine pitted > 1 ^m diameter

{Lycopodium lucidulum Fig. 18e).
GEMMA: sculptural element with radial projection isodiametric, at least one

dimension > 1 ^m, height ^ width, base constricted and top rounded (Juni-

perus Fig. 5b).
HETEROPOLAR: apertures not distributed with equal density over the distal

and proximal hemispheres (Juglans Fig. 7n-p).
INTECTATE: pollen in which the ektexine elements, if present, are free and

isolated (Ilex Fig. lOo), or form an open pattern (Fraximts nigra Fig. 10e, f).
LACUNA: depressed area in the ektexine, few in number and regularly spaced

in relation to the symmetry of the pollen grain (Agoseris Fig. 12a-c); c.f.

lumen.
LAESURA: aperture of spores functioning as a dehiscence fissure.
LIP: a swelling or thickening along the margin of a laesura (Adiantum Fig. 18a);

c.f. annulus.
LUMEN: the space bounded by the muri (walls) of a reticulum, not regularly

spaced (Impatiens Fig. 1 lG, h) ; c.f. lacuna.
MEMBRANE: exine that forms the floor of an aperture (Saxifraga oppositifolia

Fig. 9c, d).
MURUS: wall of the network that together with the lumina forms a reticulum

(Impatiens Fig. 1 1g, h) .
OBLATE: shape class with ratio between the polar and equatorial axes less than

0.75 (Ulmus Fig. In).
OPERCULUM: an isolated part of the ektexine that is contained within a colpus

or pore (Amorpha canescens Fig. 14n, o).
PERINE: loose-fitting sac enclosing certain spore taxa (Pteretis pensyhanica

Fig. 16g).
PERPROLATE: shape class with ratio between polar and equatorial axes ^ 2.0

(Umbelliferae Fig. 14f-h).
POLAR AREA INDEX: greatest distance between the ends of two colpi ex-
pressed as a ratio of the greatest breadth of the grain.
POLE: that part of a pollen grain or spore turned either inward or outward in

its tetrad, characterized for example in the tricolpate class by the area between

the ends of the colpi.
PORE: an aperture, arbitrarily defined as one with a length breadth ratio < 2.0

(Juglans Fig. 7n-p) .
POROID: pore-like, a poorly-defined pore (Cyperaceae Fig. 7l, m).
p.p.: pro porta, in part.
PROLATE: shape class with ratio between polar and equatorial axes 1.33 - 2.0

(Shepherdia canadensis Fig. 13n, o).
PROXIMAL: that part of a pollen grain or spore which faces inward in its tetrad,

representing for example in trilete spores the area around the triradiate

laesurae.

16

PSILATE: sculptural type with elements absent, exine smooth (Larix Fig. 4d).
RETICULUM: net-like sculpturing with diameter of lumina > breadth of muri

(Lycopodium obscurum Fig. 20j).
RHOMBOHEDRAL: shape of a tetrad that approximates a rhombus (Typha

lati folia Fig. 2a).
RUGULA: sculptural element tangentially elongated and with an irregular

distribution (Lycopodium inundatum (Fig. 19f, g).
SCABRA: sculptural element with radial projection isodiametric and no dimen-
sion ^ 1 /nm (Cyperaceae Fig. 7l, m).
STRIA: sculptural element tangentially elongated and more or less parallel to

another stria {Acer rubrum Fig. 9a, b).
SUBOBLATE: shape class with ratio between polar and equatorial axes 0.88 -

0.75.
SUBPROLATE: shape class with ratio between polar and equatorial axes 1.33 -

1.14 (Salix herbacea Fig. 9o, r).
TECTATE: exine structure where tectum covers most of the surface of the grain.
TECTUM: outer layer (roof) of the exine that is most easily observed when

separated from the endexine by a cavity (Artemisia Fig. 13e, f).
TETRAD: the four pollen grains or spores that are the product of a mother cell;

although pollen or spores of most species separate and are dispersed as monads,

some disperse as tetrads in a tetrahedral arrangement (Ericaceae Fig. 2e, j;

SelagineUa selaginoides Fig. 2c), or a rhombohedral arrangement (Typha

latifolia Fig. 2a) .
VERRUCA: sculptural element with radial projection more or less isodiametric,

at least one dimension > 1 fim, height ^ width, base not constricted, and top

not pointed (Plantago major Fig. 8i).

17

FIGURES

Fig. 2— Class 1, Tetrads, X1000 (p. 5)

A. Typha latifolia, rhombohedral tetrad

b. T. latifolia, irregular tetrad

c. Selaginella selaginoides

D. Drosera rotundifolia

E. Kalmia poli folia

F. K. polifolia

G. Chamaedaphne calyculata
H. Arctostaphylos uva-ursi

I. Andromeda glaucophylla
J. Vaccinium uliginosum

39.6(38-42) /tm

49.8(46-54) Mm
46.6(42-52) /mi
29.2(28-30) ^m

28.1(26-30) Mm
45.2(44-50) ^m
40.3(38-47) /xm
38.2(34-40) Mm

20

FIGURE 2

X

i7 t

>

%

J

1.4-

D

H

2/

Fig. 3— Class 2, Vesiculate, X500 (p. 5)
A. Picea glauca, ev
b. P. glauca, pv, distal
C P. mariana, ev

D. P. mariana, pv, distal

E. Abies balsamea, ev

F. A. balsamea, pv, distal

G. Pinus banksiana, ev

H. P. banksiana, pv, distal

I. P. resinosa, ev

J. P. resinosa, ev

K. P. strobus, pv, distal

68.8(62-74) Mm
56.0(50-60) Mm
69.0(60-80) nm
36.4(34-40) fim
53.0(48-54) ixm
48.6(44-50) fim

22

FIGURE 3

JV

"fV. <i '

•

.

C

• Zfn

a>;£^ |

/

-.

s^V:

V

25

Fig. 4 — Class 3, Polyplicate, XI 000 (p. 5)

a. Ephedra torreyana, ev 45.0(42-50) ^m

B. E. nevadensis, ev 67.4(62-70) ^m

Class 4, Inaperturate, X1000 (p. 5)

c. Ruppia occidentalis 71.0(60-78) /xm

D. Larix laricina 66.8(58-80) ^m

E. Tsuga canadensis 76.0(70-80) /*m

24

FIGURE 4

,<*.''

**•■«

25

Fig. 5 — Class 4, Inaperturate

cont'd.,

X1000 (p. 5)

a. Thuja occidentalis

23.0(22-26) iim

B. Juniperus communis

26.0(24-28) ftm

c. /. communis

D. Equisetum arvense

40.8(36-44) Mm

E. Populus tremuloides

28.8(24-32) Mm

F. Carex pensylvanica

42.0(40^*8) tim

c Cyperus inflexus

22.0(20-24) Mm

H. Potamogeton pectinatus

36.5(32-42) /an

I. P. natans

21.0(20-22) Mm

J. Triglochin maritima

22.4(20-24) jLim

Class 5, Monoporate,

XI 000

(p. 6)

K. Zizania aquatica, ev

34.2(32-36) Mm

L. Andropogon gerardi

39.2(34-44) /tm

m. Zea maj5- pv, distal

82.6(76-88) ^m

26

FIGURE 5

£rt*®

H

>

27

Fig. 6 — Class 5, Monoporate cont'd., XI 000 (p. 6)

A. Sparganium eurycarpum, pv, distal

B. Typha angustifolia, pv, distal
c. Lemna trisulca, ev

Class 6, Diporate, X1000 (p. 6)

D. Morus rubra, ev

Class 7, Triporate, XI 000 (p. 6)

E. Epilobium latifolium, pv

F. Myrica gale, pv

G. Betula papyrifera, pv
H. B. glandulosa, pv

I. B. populifolia, pv
J. Ostrya virginiana, pv
K. Carpinus caroliniana, pv
L. Tilia americana, pv
M. Humuhis lupulus, pv
n. Celtis occidentalis, pv
o. C. occidentalis, ev

26.4(24-30) Mm
21.4(18-22) /im
21.5(19-23) /im

19.0(18-20) /*m

79.0(62-86) nm
27.8(26-30) /am
29.6(28-34) Mm
21.2(18-24) /xm
24.0(20-26) /im
28.0(26-30) fim
31.0(28-32) fim
36.2(34-38) Mm
23.0(20-26) ^m
29.6(28-32) Mm

28

FIGURE 6

29

Fig. 7 — Class 7, Triporate cont'd., X1000 (p. 7)

A. Carya cordiformis, pv

B. Cory Ins cornuta, pv
c. Urtica procera, pv

D. Morus rubra, pv

Class 8, Stephanoporate, XI 000 (p. 7)

E. Alnus rugosa, pv

F. A. rugosa, pv

G. A . crispa, pv

h. Ulmus thomasii, ev

I. U. thomasii, pv

J. Myriophyllum alterniflorum, pv

K. M. exalbescens, pv

Class 9, Periporate, XI 000 (p. 7)
L. Eriophorum angustifolium
M. Car ex lacustris
N. Juglans cinerea, pv
o. /. cinerea, pv
P. J. nigra, pv
0. Sagittaria latifolia

44.4(42-46) /mi
22.4(22-24) Km
15.6(14-18) /nil
19.0(18-20) ym

22.0(20-24) fim

18.4(18-20) /im
35.0(32-38) /*m

19.8(18-22) /*m
30.2(30-32) (im

38.8(36-40) fim
45.8(42-50) (im
38.2(36-40) fim

37.4(34-40) ixm
24.0(22-26) fim

30

FIGURE 7

*r

H

)

J

: /

•- i - Q

31

Fig. 8— Class 9, Periporate cont'd., XI 000 (p. 7)

a. Polygonum lapathifolium 34.8(32-38) /*m

B. Liquidambar styraciflua 34.4(32-36) /im

c. Stellaria crassifolia 28.0(24-32) /urn

D. Sarcobatus vermiculatus 25.6(24-28) /*m

E. Plantago lanceolata 24.8(22-26) /im

F. Suaeda depressa 24.6(22-28) /*m

G. Salsola pestifer 26.8(24-30) ^m
H. Thalictrum dasycarpum 22.8(20-26) /im
I. Plantago major 21.8(20-24) /*m

Class 10, Monocolpate, XI 000 (p. 8)

J. Nymphaea sp., ev 25.8(24-28) /urn
K. Nymphaea sp., pv

L. Nuphar variegatum, pv 48.6(46-50) (*m

M. Brasenia schreberi, ev 52.0(48-56) /tm

Class 11, Dicolpate, XI 000 (p. 8)

N. Heteranthera dubia 63.0(50-72) j*m

52

FIGURE 8

€

B

••; a

V

A \:

33

Fig. 9— Class 12, Tricolpate, X1000 (p. 8)

A. Acer rubrum, ev 34.4(32-38) /xm

B. ,4. rubrum, pv

c. Saxifraga oppositifolia, ev 29.8(28-32) /*m

D. 5. oppositifolia, pv

E. /4. saccharinum, ev 37.4(32-42) /*m

F. /I. saccharinum, ev

G. Dryas integrifolia, ev 23.6(20-28) ju.m
H. Z). integrifolia

I. D. integrifolia, pv

J. /Jeer negundo, ev 23.4(22-26) ^m

K. /I. negundo, ev

L. /i. negundo, pv

M. Sa//x discolor, pv 26.8(24-28) ^m

N. 5. discolor, ev

o. 5. Candida, ev 20.8(18-24) ^m

p. 5. interior, ev 15.4( 14-18) /*m

Q. S.herbacea,ev 19.0(18-20) /*m

R. 5. herbacea, ev

s. Petalostemum purpureum, ev 43.6(40-50) /*m

T. /leer saccharum, ev 32.0(28-34) ^m

u. /4. saccharum, pv

54

FIGURE 9

V

/

s

\'

f

u

J

35

Fig. 10— Class 12, Tricolpate cont'd., XI 000 (p. 9)

A. Stachys palustris, pv

B. S. palustris, pv

c. Rorippa island ica, ev

D. Brassica kaber, pv

E. Fraxinus nigra, ev

F. F. nigra, pv

G. F. quadrangulata, pv

h. Platanus occidentalis, pv

I. Quercus rubra, pv

J. Q. rubra, ev

K. Q. macrocarpa, ev

L. Caltha palustris, ev

M. Ambrosia psilostacbya, pv

N. Ranunculus flabellaris, ev

o. //e* verticillata, pv

p. Nemopanthus mucronata, ev

Q. Rubus chamaemorus, pv

R. 7?. chamaemorus, ev

Class 13, Stephanocolpate, XI 000 (p. 10)
s. Galium boreale, ev
T. G. asprellum, pv
u. G. asprellum, pv

25.4(24-28) Mm

19.6(18-22) Mm
27.2(24-32) /xm
23.0(22-24) /xm

22.0(20-24) Mm
18.0(16-20) ,am
24.8(24-28) ^m

28.2(26-30) /mi
24.2(22-26) /xm
23.4(22-24) ^m
37.0(30-40) /xm
28.8(24-32) /xm
24.2(22-26) /xm
33.8(30-36) fim

23.8(22-26) /xm
18.4(16-20) /xm

.?6

FIGURE 10

a

I

<m

D

■"**>»

i

i )

U

37

Fig. 11 — Class 13, Stephanocolpate cont'd., X1000 (p. 10)

A. Hippnris vulgaris, pv 30.2(26-32) (im

b. Lycopus virginicus, pv 28.6(26-32) /*m

c. Mentha arvensis, ev 32.8(30-36) jum

D. Fraxinus pennsylvanica,pv 22.2(20-24) /xm

E. F. pennsylvanica, ev

f. F. americana, pv 26.2(24-30) /*m

G. lmpatiens capensis, pv 32.4(30-36) /im
H. /. capensis, pv

I. Arceuthobium pusillum,pv 24.0(22-26) //m
J. /I. pusillum, pv

Class 14, Pericolpate, XI 000 (p. 10)

K. Polygonum amphihium 50.2(46-54) ^m

l. P. coccinium 59.8(52-66) ^m

M. Portulaca oleracea 56.8(54-64) ^m

38

FIGURE 11

t" )

4ff-\

t. ■:■

-: >

±L&

)

B

_>^!

<+> C

f " }

.'"'-' ^

■ Si-

[#*;- >*.*5 *-

l5*v~ ' '

'+*/

CH^* .^ —«*•*<» ^

j w

^&&+z£

L

59

Fig. 12— Class 15, Tricolporate, XI 000 (p. 10)

a. Agoseris glauca, e\ 42.4(38-50) /*m

B. A. glauca, pv
c. A . glauca, pv

D. Helianthuslaetiflorus,ev 33.0(30-36) fim

E. H. laetiflorus, pv

F. Eupatorium perfo!iatum,e\ 18.0(16-20) /*m

G. E. perfoliatum, pv

H. Ivaciliata,pv 20.8(20-22) ^m

I. /. ciliata, pv

j. Xanthium pensylvanicum, pv 25.0(24-26) f»m

K. Iva xanthifolia, ev 15.8(14-16) /^m

L. /. xanthifolia, pv

M. Ambrosia artemisiifolia, ev 18.0(16-20) Mm

N. A. artemisiifolia, pv

o. Castanea dentata, ev 13.0(12-14) ^m

p. C. dentata, ev

40

FIGURE 12

•* <• • •- -

■■■x^^,.

• \

N

O

0

4/

Fig. 13— Class 15, Tricolporate cont'd., X1000 (p.

A. Elaeagnus commutata, pv

B. Shepherdia argentea, pv
c. Polygonum aviculare, ev

D. P. cristatum, ev

E. Artemisia ludoviciana, pv

F. A. ludoviciana, ev

G. Nyssa sylvatica, ev
H. N. sylvatica, ev

I. N. sylvatica, ev

J. Fagus grandifolia, ev

K. F. grandifolia, pv

L. Rumex mexicanus, ev

M. R. mexicanus, pv

N. Shepherdia canadensis, ev

o. 5. canadensis, ev

p. Cornus canadensis, ev

Q. C. canadensis, ev

11)

35.8(30-40) nm
34.8(30-40) ^m
28.8(26-32) fim
26.6(26-28) Mm
23.0(22-26) /tm

31.6(30-34) fim

41.2(40-44) ^m
25.4(24-30) (i,m
35.8(34-38) ^m
23.4(22-26) /*m

42

FIGURE 13

43

Fig. 14— Class 15, Tricolporate cont'd., X1000 (p. 11)

a. Cornus stolonifera, ev

b. C. stolonifera, ev

c. C. racemosa, ev

D. Polygonum viviparum, ev

E. P. viviparum, ev

F. Sium suave, ev

G. S. suave, ev

H. Cicuta bulbifera, ev

I. Menyanthes trifoliata, ev

J. M. trifoliata, pv

K. Prunus pensylvanica, ev

L. Potentilla palustris, ev

m. 77/za americana, ev

N. Amorpha canescens, ev

o. /4. canescens, ev

p. Sa//* amygdaloides, ev

Q. 5. amygdaloides, ev

58.2(54-64) Fm

38.5(37^0) jam
47.0(42-54) /urn

27.6(26-30) /xm

26.6(24-28) Mm
30.8(28-34) ftm

29.4(26-32) /*m
20.0(18-22) /iiti
36.2(34-38) cm
20.8(20-22) fim

20.4(20-22) ^m

44

FIGURE 14

**<*>

ft

H

QQ

v

45

ig. 15 — Class 15, Tricolporate cont'd., X1000

A. Sambucus pubens, ev

B. Vitis riparia, ev

(p. 12)

18.6(18-20) ^m
20.6(20-22) fim

c. V. riparia, pv

D. Cephalanthus occidentalis, ev

E. C. occidentalis, ev

17.4(16-20) tim

Class 16, Stephanocolporate, XI 000 (p.
f. Utricularia vulgaris, pv
G. U. vulgaris, ev
H. Sanguisorba canadensis, pv
I. S. canadensis, ev

12)

36.4(32-40) jam
24.2(22-26) p.m

46

FIGURE 15

^~^

A

\

**% #■

47

Fig. 76— Spore class 1, Alete, XI 000 (p. 13)

A. Equisetum arvense

Spore class 2. Monolete, XI 000 (p. 13)

B. Dryopteris spinulosa, ev

c. Cystopteris fragilis, pv, proximal

d. Athyrium filix-femina, ev

E. Polypodium virginianum, ev

F. Isoetes macrospora, ev

G. Pteretis pensylvanica, ev
H. Woodsia ilvensis, ev

I. Dryopteris thelypteris, ev
J. D. goldiana, ev
K. Cystopteris bulbifera, pv, distal
L. C. fragilis, ev

40.8(36-44) /*m

45.6(40-52) Mm
38.2(34-44) ^m
36.8(34-40) Mm
67.2(56-76) /mi
51.4(46-56) Mm
45.6(38-50) ^m
47.8(44-56) Mm
44.8(42-48) Mm
31.6(28-34) Mm
33.2(30-36) fim
41.4(38-44) Mm

48

FIGURE 16

49

Fig. 17 — Spore class 2, Monolete cont'd., XI 000 (p. 13)

A. Dryopteris spinulosa, ev 45.6(40-52) jum

B. Camptosaurus rhizophyllus, pv, proximal 27.0(24-32) /urn

c. Polystichum lonchitis, pv, distal 35.9(32-38) fitn
Spore class 3, Trilete, XI 000 (p. 13)

d. Cheilanthes feei, pv, proximal 76.8(74-80) ^m

E. Selaginella rupestris, pv, proximal 44.4(38-52) f»m

F. Pteridium aquilinum, pv, proximal 31.2(30-36) yum

G. P. aquilinum, pv, distal

H. Cheilanthes feei, pv, proximal 59.0(52-64) /*m

50

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ROYAL ONTARIO MIIQfriiM

FIGURE 17

H

51

Fig. 18 — Spore class 3, Trilete cont'd., X1000 (p. 14)

a. Adiantum pedatum, pv, proximal 37.0(36-40) /urn

B. Dennstaedtia punctiloba, pv, proximal 31.6(30-34) /um

c. Pteridium aquilinum, pv, proximal 31.2(30-36) iim

D. P. aquilinum, pv, proximal

E. Lycopodium lucidulum, pv, distal 32.0(30-36) /xm

f. L. lucidulum, pv, proximal

g. L. selago, pv, distal 38.4(34-42) ^m
H. L. selago, pv, proximal

I. Botrychium multifidum, pv, distal 35.8(34-40) /xm

J. B. multifidum, pv, proximal

K. B. dissectum, pv, distal 37.2(34-42) ^m

l. B. dissectum, pv, proximal

52

FIGURE 18

—

*A A

■

^ rfl

4

s 7.

V

/?«

K

55

Fig. 19 — Spore class 3, Trilete cont'd., X1000 (p. 14)

a. Osmunda cinnamomea, pv, distal 41.2(38—44) ^m

B. O. regalis, pv, proximal 51.2(48-54) /nm

c. O. regalis, pv, distal

D. O. claytoniana, pv, proximal 42.2(40-46) ^m

E. Selaginella selaginoides, pv, proximal 32.8(30-36) /im
f. Lycopodium inundatum, pv, proximal 48.0(46-50) ^m
G. L. inundatum, pv, distal

54

FIGURE 19

- ~ ^ o ->

ft* D

*

"^

55

Fig. 20 — Spore class 3, Trilete cont'd., X1000 (p. 14)

a. Ophioglossum vulgatum, pv, proximal 44.4(40-50) ^.m

B. O. vulgatum, pv, distal

c. Lycopodium clavatum, pv, proximal 37.6(36-40) jim

d. L. clavatum, pv, distal

E. L. complanatum, pv, proximal 34.6(32-36) ^m

f. L. complanatum, pv, distal

G. L. tristachyum, pv, proximal 32.0(30-36) ^m

H. L. tristachyum, pv, distal

I. L. obscurum, pv, proximal 37.0(34-40) /xm

J. L. obscurum, pv, distal

.56

FIGURE 20

%_

kit

,#

B

M ^

1*

-

rS.

D

'

,f-«r^

V1V

I

.7 \

*-jt*f

Fig. 21 — Spore class 3, Trilete cont'd., X1000 (p. 15)

A. Lycopodium annotinum, pv, proximal

B. L. annotinum, pv, distal

c. Botrychium virginianum, pv, proximal

D. B. virginianum, pv, distal

E. Sphagnum sp., pv, distal

f. Sphagnum sp., pv, proximal

c. Botrychium simplex, pv, proximal

H. B. lunaria, pv, proximal

i. B. lunaria, pv, distal

j. B. lanceolatum, pv, proximal

k. B. lanceolatum, pv, distal

37.8(34-40) Mm

34.0(30-36) fxm

32.0(26-34) ,um

51.2(46-56) /*m
39.2(38-44) Mm

37.4(34-40) fxm

58

FIGURE 21

59

Acknowledgments

We thank Dr. J. C. Ritchie and P. Richard for critically reviewing the manu-
script and supplying helpful suggestions. Miss Margaret Coutinho patiently
typed the difficult key. Mr. Brian O'Donovan's skill in photographic printing
was invaluable. Finally, we sincerely appreciate the comments of the several
students who struggled at the miscroscope with earlier versions of the key.

Support was received through National Research Council operating
grants to J. H. McAndrews (A5699) and to G. Norris (A4246).

Literature Cited

ADAMS, R. J. AND J. K. MORTON

1972 An atlas of pollen of the trees and shrubs of eastern Canada and the
adjacent United States. Part 1. Gymnospermae to Fagaceae. Uni-
versity of Waterloo Biology Series No. 8. Waterloo, Ontario, Uni-
versity of Waterloo. 52 pp.

BASSETT, I. J. AND C. W. CROMPTON

1968 Pollen morphology and chromosome numbers of the family Planta-
ginaceae in North America. Can. J. Bot., vol. 46, no. 4, pp. 349-361 .

BRAUN, E. L.

1950 Deciduous forests of eastern North America. Philadelphia, Blakis-
ton. 596 pp.

ERDTMAN, G.

1957 Pollen and spore morphology/ plant taxonomy. Gymnospermae,
Pteridophyta, Bryophyta. Stockholm, Almqvist and Wiksell. 151 pp.

1965 Pollen and spore morphology/ plant taxonomy. Gymnospermae,
Bryophyta. Stockholm, Almqvist and Wiksell. 191 pp.

1966 Pollen morphology and plant taxonomy. Angiosperms. Reprint ed.
with addendum. New York, Hafner. (Stockholm, Almqvist and
Wiksell, 1962). 553 pp.

ERDTMAN, G. AND P. SORSA

1971 Pollen and spore morphology/ plant taxonomy. Pteridophyta. Stock-

holm, Almqvist and Wiksell. 302 pp.

FAEGRI, K. AND J. IVERSON

1964 Textbook of pollen analysis. 2nd rev. ed. New York, Hafner. 237
pp.

FERNALD, M. L.

1950 Gray's manual of botany; a handbook of the flowering plants and
ferns of the central and northeastern United States and adjacent
Canada. New York, American Book. 1632 pp.

HARRIS, w. F.

1955 A manual of the spores of New Zealand Pteridophyta. Bull. N. Z.
Dep Scient. Ind. Res., no. 116, pp. 1-186.

60

HELMICH, D. E.

1963 Pollen morphology in the maples (Acer L.). Pap. Mich. Acad. Sci.,
vol. 48, pt. 1, pp. 151-164.

KAPP, R. O.

1969 How to know pollen and spores. Dubuque, Iowa, W. C. Brown.
249 pp.

KREMP, G. O. W.

1965 Morphologic encyclopedia of palynology; an international, col-
lection of definitions and illustrations of spores and pollen. Tucson,
University of Arizona Press. 263 pp.

MAHER, L. J., JR.

1964 Ephedra pollen in sediments of the Great Lakes region. Ecology,
vol. 45, no. 2, pp. 391-395.

MCANDREWS, J. H. AND A. R. SWANSON

1967 The pore number of periporate pollen with special references to
Chenopodium. Rev. Palaeobotan. Palynol., vol. 3, pp. 105-117.

RICHARD, P.

1970 Atlas pollinique des arbres et de quelques arbustes indigenes du
Quebec. Le Naturaliste Can., vol. 97, no. 1, pp. 1-34; no. 2, pp.
97-161; no. 3, pp. 241-306.

ROWE, J. S

1959 Forest regions of Canada. Bull. For. Brch. Can., no. 123, pp. 1-71.

SEARS, P. B.

1930 Common fossil pollen of the Erie basin. Botanical Gazette, vol. 89,
no. 1, pp. 95-106.

TSCHUDY, R. H. AND R. A. SCOTT (EDS.)

1969 Aspects of palynology. New York, Wiley-Interscience. 510 pp.

WHITEHEAD, D. R.

1964 Fossil pine pollen and full-glacial vegetation in southeastern North
Carolina. Ecology, vol. 45, no. 4, pp. 767-777.

61

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