Royal Ontario Museum Life Sciences Miscellaneous Publication
-<M "CD :CO
■Lf)
O
= co
;°
Key to the Quaternary Pollen and Spores of the Great Lakes Region
John H. McAndrews, Albert A. Berti and Geoffrey Norris
• ■:■■■■.
■
1 ' W&MmI
■
•■■■■->; ':'-'--.-: -
-..•;.'•
SR
•->...-
#i
I I Ivl
\, i ' 1V1
m*3Q
•?;"^ .'.-'."
H8E
iH
2g&g£
■.';-'
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 PUBLICATIONS IN LIFE SCIENCES
The Royal Ontario Museum publishes three series in the Life Sciences:
life sciences contributions, a numbered series of original scientific publi- cations, including monographic works.
life sciences occasional papers, a numbered series of original scientific publications, primarily short and usually of taxonomic significance.
life sciences miscellaneous publications, an unnumbered series of publications of varied subject matter and format.
All manuscripts considered for publication are subject to the scrutiny and editorial policies of the Life Sciences Editorial Board, and to review by persons outside the Museum staff who are authorities in the particular field involved.
LIFE SCIENCES EDITORIAL BOARD
Chairman, r. l. Petersen 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
LIBRARY ROYAL ONTARIO MUSEUM
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
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
LIBRARY 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
LIBRARY ROYAL ONTARIO MUSEUM
m
HM
ISBN 0-88854-149-X