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Heidemarie Halbritter · Silvia Ulrich
Friðgeir Grímsson · Martina Weber · Reinhard Zetter
Michael Hesse · Ralf Buchner · Matthias Svojtka
Andrea Frosch-Radivo
Illustrated
Pollen
Terminology
Second Edition
Illustrated Pollen Terminology
Heidemarie Halbritter • Silvia Ulrich
Friðgeir Grímsson • Martina Weber
Reinhard Zetter • Michael Hesse
Ralf Buchner • Matthias Svojtka
Andrea Frosch-Radivo
Illustrated Pollen
Terminology
Second Edition
Heidemarie Halbritter Silvia Ulrich
Division of Structural and Functional Botany Division of Structural and Functional Botany
Department of Botany and Biodiversity Department of Botany and Biodiversity
Research Research
University of Vienna University of Vienna
Vienna Vienna
Austria Austria
© The Editor(s) (if applicable) and The Author(s) 2009, 2018. This book is an open access publication.
Open Access This book is licensed under the terms of the Creative Commons Attribution 4.0
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Preface to the First Edition
There are more things in heaven and earth, than are dreamt of in our philosophy.
Shakespeare, Hamlet to Horatio
The principal aim in compiling this book was to provide the reader with first-
hand information about the structure and outlook of the extremely manifold
pollen in seed plants. This book should not be seen as a mere collection of
striking and/or informative light and electron micrographs. Each of the micro-
graphs is intended to convey a specific message related to properties and
functions of the pollen grains shown. The authors hope that the book will be
useful for experienced researchers as well as for beginners in palynology, but
also for medicine, biochemistry, or even for lawyers and artists as an aid and
guide for the evaluation and interpretation of pollen features.
V
Preface to the Second Edition
The first edition of this book Pollen Terminology: An illustrated handbook was
extremely successful and well received by the worldwide palynological com-
munity. As with the first edition, our main intention when compiling the second
edition was to provide both scientists and the public with an easily under-
standable and primarily illustrative access to the hidden beautiful world of pol-
len and the fascinating subject of palynology. This new edition titled Illustrated
Pollen Terminology allowed us to improve many aspects of our book and to
illustrate in more detail the important concepts and various preparation tech-
niques applied in (paleo)palynology. It is our hope that this edition will become
the guidance tool for all students of palynology, as well as reference work and
illustrated encyclopedia for the more advanced scientists.
VII
Contents
IX
X Contents
Aperture��������������������������������������������������������������������������������������������������������������� 207
angulaperturate ����������������������������������������������������������������������������������������� 208
annulus/annulate���������������������������������������������������������������������������������������� 209
aperture/aperturate����������������������������������������������������������������������������������� 211
aperture membrane����������������������������������������������������������������������������������� 214
atrium, oncus����������������������������������������������������������������������������������������������� 220
brevicolpus/brevicolpate��������������������������������������������������������������������������� 221
brevicolporus/brevicolporate ������������������������������������������������������������������� 222
bridge����������������������������������������������������������������������������������������������������������� 223
colpus/colpate, dicolpate������������������������������������������������������������������������� 225
colpus/colpate, tricolpate������������������������������������������������������������������������� 226
colpus/colpate, tetracolpate, pentacolpate����������������������������������������� 229
colpus/colpate, hexacolpate������������������������������������������������������������������� 230
colpus/colpate, stephanocolpate����������������������������������������������������������� 231
colpus/colpate, pantocolpate����������������������������������������������������������������� 232
colporus/colporate, dicolporate��������������������������������������������������������������� 233
colporus/colporate, tricolporate��������������������������������������������������������������� 234
colporus/colporate, tetracolporate��������������������������������������������������������� 237
colporus/colporate, pentacolporate, stephanocolporate������������������� 238
colporus/colporate, stephanocolporate������������������������������������������������� 239
ektoaperture, endoaperture, lalongate, lolongate ������������������������������� 240
heteroaperturate����������������������������������������������������������������������������������������� 242
inaperturate������������������������������������������������������������������������������������������������� 244
leptoma ������������������������������������������������������������������������������������������������������� 247
margo����������������������������������������������������������������������������������������������������������� 248
operculum/operculate������������������������������������������������������������������������������� 251
pantoaperturate, pantocolpate��������������������������������������������������������������� 255
pantoaperturate, pantocolporate, pantoporate����������������������������������� 256
pantoaperturate, pantoporate����������������������������������������������������������������� 257
papilla����������������������������������������������������������������������������������������������������������� 259
planaperturate��������������������������������������������������������������������������������������������� 260
pontoperculum/pontoperculate��������������������������������������������������������������� 262
porus/porate, diporate������������������������������������������������������������������������������� 263
porus/porate, triporate������������������������������������������������������������������������������� 264
porus/porate, tetraporate ������������������������������������������������������������������������� 266
porus/porate, pentaporate����������������������������������������������������������������������� 267
porus/porate, stephanoporate����������������������������������������������������������������� 268
porus/porate, pantoporate����������������������������������������������������������������������� 269
poroid/poroidate����������������������������������������������������������������������������������������� 272
pseudocolpus����������������������������������������������������������������������������������������������� 273
ring-like aperture����������������������������������������������������������������������������������������� 274
spiral aperture/spiraperturate ������������������������������������������������������������������� 277
stephanoaperturate����������������������������������������������������������������������������������� 279
sulcus/sulcate����������������������������������������������������������������������������������������������� 281
sulcus/sulcate, disulcate����������������������������������������������������������������������������� 287
sulcus/sulcate, trichotomosulcate������������������������������������������������������������� 288
sulcus/sulcate, polychotomosulcate ������������������������������������������������������� 289
synaperturate, syncolpate, syncolporate ����������������������������������������������� 290
synaperturate, syncolporate ��������������������������������������������������������������������� 291
ulcus/ulcerate ��������������������������������������������������������������������������������������������� 292
Ornamentation ��������������������������������������������������������������������������������������������������� 295
areola/areolate������������������������������������������������������������������������������������������� 296
baculum/baculate������������������������������������������������������������������������������������� 298
bireticulate��������������������������������������������������������������������������������������������������� 299
Contents XIII
elastoviscin��������������������������������������������������������������������������������������������������� 416
viscin thread������������������������������������������������������������������������������������������������� 418
Ubisch body (Ubisch bodies)��������������������������������������������������������������������� 421
Pollen Class��������������������������������������������������������������������������������������������������������� 429
pollen class��������������������������������������������������������������������������������������������������� 430
Part IV Annex
DMP 2,2-Dimethoxypropane
KMnO4 Potassium permanganate
LM Light microscope
PA Periodic acid
PA+TCH+SP Thiéry test
PA+TCH+SP (short) Modified Thiéry test
TCH+SP Lipid test
Pb Lead citrate
SEM Scanning electron microscope
SP Silver proteinate
TCH Thiocarbohydrazide
TEM Transmission electron microscope
U Uranyl acetate
Prefixes
a- Prefix meaning absent
bi- Prefix for two
brevi- Prefix meaning short
di- Prefix meaning two
eu- Prefix meaning true
hetero- Prefix meaning different
hexa- Prefix meaning six
homo- Prefix meaning equal
in- Prefix meaning absent
infra- Prefix meaning beneath
inter- Prefix for in between
intra- Prefix for within
iso- Prefix meaning identical
meso- Prefix meaning middle
micro- Prefix for small; features between 1 and 0.5 μm
mono- Prefix meaning one
nano- Prefix for very small; features between 0.5 and 0.1 μm
panto- Prefix for global
penta- Prefix meaning five
XV
XVI Abbreviations
XVII
XVIII Introduction
all terms are listed along with their definition in the “Glossary of Palynological
Terms”. Numbers following terms refer to the respective page(s) where the
terms are discussed. Numbers in bold relate to illustrations in the chapters of
the “Illustrated Pollen Terms.” The fourth part “Annex” comprises the “Picture
Copyrights” and the “Index” listing all plant names occouring in this book.
1 I
General Chapters
Contents
Palynology: History and Systematic Aspects – 3
Pollen Development – 23
Misinterpretations in Palynology – 67
Methods in Palynology – 97
Palynology: History
and Systematic Aspects
The History of Palynology – 4
Future Perspective – 16
References – 18
3
PALYNOLOGY: HISTORY AND SYSTEMATIC ASPECTS
Palynology is the science of palynomorphs, a gen- pound microscope was developed by Hooke
eral term for all entities found in palynological prep- (1665). This was an important contribution to the
arations (e.g., pollen, spores, cysts, diatoms). A study of pollen morphology. Malpighi in his
dominating object of the palynomorph spectrum is “Anatomia Plantarum” was the first to describe pol-
the pollen grain. The term palynology was coined len grains as having germination furrows while Grew
by Hyde and Williams (1955; Fig. 1). It is a combina- noted in his famous work “The Anatomy of Plants”
tion of the Greek verb paluno (пαλύνω, “I strew or the constancy of pollen characters within the same
sprinkle”), palunein (пαλύνειν, “to strew or sprinkle”), species (Fig. 2; Grew 1682; Malpighi 1901). They are
the Greek noun pale (пαλƞ, in the sense of “dust, both considered the founders of pollen morphol-
fine meal,” and very close to the Latin word pollen, ogy. Camerarius described several pollination
meaning “fine flour, dust”), and the Greek noun experiments and communicated the results in his
logos (λογος, “word, speech”). letters about plant sexuality to Valentini (Camerarius
1694). He stated that male “seed dust” is necessary
for seed development. Von Linné (also known
The History of Palynology before his ennoblement as Carl Nilsson Linnæus) first
used the term pollen (in 1750). In the 18th and the
A
ssyrians are said to have known the principles early nineteenth centuries, there was considerable
of pollination (they practiced hand pollination progress in pollen research and the understanding
of date palms), but it is unclear if they recog- of pollination. In 1749, Gleditsch demonstrated in a
nized the nature of pollen itself. The invention of the spectacular experiment (Experimentum
first microscopes and especially the compound Berolinense) the central role of pollen in double fer-
microscope in the late sixteenth century represents tilization. He organized the transport of an inflores-
the starting point of a new fascinating era. Some of cence from a male fan palm in Leipzig to a hitherto
the most important findings and scientists within the “sterile” female fan palm growing in a greenhouse
long tradition of light microscopy are mentioned in Berlin. After pollination, the female flowers pro-
here. For a more comprehensive overview, see duced fertile seeds for the first time in the palms life-
Wodehouse (1935) and Ducker and Knox (1985). time (Gleditsch 1751, 1765). Placing male
Following the invention of the simple microscope inflorescences within groups of female date palms
by J. Janssen and Z. Janssen in 1590, the first com- for pollination was according to Theophrast already
Fig. 1 The right word. Excerpt from Hyde and Williams (1955). Pollen Analysis Circular no. 8, p. 6
4 GENERAL CHAPTERS
PALYNOLOGY: HISTORY AND SYSTEMATIC ASPECTS
practiced by the Assyrians and Egyptians. Kölreuter, in the outer wall of both pollen grains and spores
together with Sprengel, the founder of research on (Zetzsche and Huggler 1928; Zetzsche and Vicari
flower biology, perceived the importance of insects 1931; Zetzsche et al. 1931). Campbell reported pol-
in flower pollination and discovered for the first time len of seagrasses (Naias and Zannichellia) to be thin
that pollen plays an important role in determining walled, without exospore (exine), and two-celled.
the characters of the offspring (Kölreuter 1761-1766; Moreover, Campbell described the mitotic division
Sprengel 1793). Kölreuter (1806, 1811) also discov- of the generative cell into two (sperm) cells
ered that the pollen wall is c onsisting of two distinct (Campbell 1897). Hofmeister and Strasburger pro-
layers and made the first attempt to classify pollen vided ground-breaking insights into the develop-
based on their morphology. Sprengel recognized ment and internal structure of pollen and fertilization
pores and furrows in the pollen wall and demon- (involves the fusion of a single sperm nucleus and
strated the effects of cross pollination, dichogamy, the egg nucleus) and investigated the bi- and tri-
and distinguished between entomo- and ane- cellular pollen condition of many angiosperms
mophily (Candolle and Sprengel 1821). Moreover, (Strasburger 1884; Hofmeister 1849). The role of the
he also realized that every plant species has a char- second sperm nucleus in the pollen tube remained
acteristic pollen type (Sprengel 1804). unexplained until double fertilization was discov-
During the first half of the nineteenth century, ered by Guignard (1891, 1899); Nawaschin (1898).
some fundamental insights into pollen morphology Nägeli studied the ontogeny of pollen grains within
and physiology were achieved. Purkinje made the anthers and was the first to recognize the callose
first attempt for a palynological terminology by wall (Nägeli 1842). Schacht described differences in
classifying pollen based on their morphology exine patterning, exine thickness, and apertures
(Purkinje 1830). Wodehouse (1935) pointed out that covered by an operculum. He also used cytochem-
“Purkinje’s system of nomenclature deserved much ical staining techniques to detect pollen reserves.
more attention than was ever given to it by subse- He was also the first to cut sections of embedded
quent investigators. A system of this kind, had it pollen with razor blades for anatomical studies
been put into use, would have saved much confu- (Schacht 1856/59). Strasburger described the basic
sion.” Brown gave the first description of the origin concepts of pollen wall development already in
and role of the pollen tube (Brown 1828, 1833). He 1889, but major break-through in pollen wall ontog-
credited Bauer as the first observer of the pollen eny was achieved much later by Heslop-Harrison
tube’s nature, of the double wall in Asclepias pol- (1975). The first successful classification of orchida-
len, and for his minute drawings of Asclepias pollen. ceous plants based on pollen features was made
His brother Bauer, a great botanical artist, was the by Lindley (1836). Later, Fischer recognized the
first to recognize compound pollen in Acacia and potential of pollen morphology in aiding the phylo-
orchids. Cavolini described and illustrated the fili- genetic position of angiosperms (Fischer 1890).
form pollen of sea grasses Zostera and Cymodocea Paleopalynology was established at the end of
(Cavolini 1792). the nineteenth century, when P. Reinsch published
Göppert and Ehrenberg were the first to describe the first photomicrographs of fossil pollen and spores
and depict fossil pollen grains and spores (Göppert from Russian coals (Reinsch 1884). He also described
1837, 1848; Ehrenberg 1838). In 1834 von Mohl wrote methods for the extraction of palynomorphs from
his fundamental work entitled “Über den Bau und coal samples with concentrated potassium hydrox-
die Formen von Pollenkörner/On the structure and ide (KOH) and hydrofluoric acid (HF). Von Post pub-
diversity of pollen grains,” which was a major contri- lished the first pollen diagram (profile) using
bution to the knowledge of pollen structure and exclusively arboreal pollen (von Post 1916). Already
descriptive classification. von Mohl and Fritzsche before and especially after the Second World War,
recognized the principal layers of the pollen wall Schopf as well as Potonié published their impressive
and published new surveys on pollen morphology publications devoted to fossil spores and pollen
(von Mohl 1835; Fritzsche 1837). The term pollenin (e.g., Potonié 1956; Schopf 1957, 1964). Schopf
goes back to von Grotthuss (1814), John (1814), established the systematic study of palynomorphs,
Stolze (1816), and Fritzsche (1834). The terms “exine,” while Potonié was one of the first who recognized
“intine,” and “Zwischenkörper” were established by the stratigraphic value of paleopalynology, apply-
Fritzsche and published in his book “Über den ing his “turmal classification” system (Potonié 1934;
Pollen” (Figs. 3 and 4; Fritzsche 1837). He also dem- see also “The Treme System and the NPC-
onstrated that apertures are predetermined in most Classification” below). The rise of stratigraphic paly-
angiosperm pollen while others are inaperturate. nology started shortly before 1950 and played a
Zetzsche first coined the term “sporopollenin” to prominent role in petroleum explorations during the
describe the resistant chemical substance present second half of the twentieth century (Manten 1966).
6 GENERAL CHAPTERS
PALYNOLOGY: HISTORY AND SYSTEMATIC ASPECTS
Fig. 3
Detailed drawings of pollen. Fritzsche (1837) “Über den Pollen”
The key role of palynology in stratigraphy depends The twentieth century up to ca 1960 was domi-
upon the fact that the natural biopolymer sporopol- nated by the skillful use of the LM, with many new
lenin in the spore/pollen walls is extremely resistant; findings; for example, the LO-analysis, a method for
thus, pollen/spores are often abundantly preserved analyzing patterns of exine organization by light
in sedimentary rocks. microscopy, focusing at different levels distinct
GENERAL CHAPTERS 7
PALYNOLOGY: HISTORY AND SYSTEMATIC ASPECTS
Fig. 4 First detailed drawings of pollen. Fritzsche (1837) “Über den Pollen”
features appear bright (L = Lux) or dark in good use (Erdtman 1943, 1952, 1957, 1969; Fægri
(O = Obscuritas). Textbooks by Wodehouse, and Iversen 1950, 1989; Wodehouse 1935). During
Erdtman, or Fægri and Iversen summarized the this time palynology also became more diverse and
knowledge of palynology from that time, but are still applied in numerous fields among others: aeropaly-
8 GENERAL CHAPTERS
PALYNOLOGY: HISTORY AND SYSTEMATIC ASPECTS
F
1955; Bradley 1958; Rowley and Flynn 1966). The or the scientist, categories are essential for classi-
time-consuming and laborious TEM replica proce- fying natural characters in their diversity, defining
dures were an obstacle to extensive surveys of pol- their range and placing them in a systematic
len morphology and later replaced by SEM (Harley order. In addition to the theoretical concept, cate-
and Ferguson 1990). The introduction of SEM in paly- gorization always depends on the manner in which a
nology in the mid of the 1960s was a key innovation feature is perceived: i.e. on the visibility of a feature,
in the study of the fine relief (sculpture) of pollen and/or their specific value. Categorization also
and spore surfaces. Advantages of SEM include the greatly depends on the technical equipment and
relatively simple and rapid preparation methods method(s) used, as well as on the subjective interpre-
and the supreme depth of focus. SEM was consid- tation of character(s) (see “Methods in Palynology”).
ered from the very first moment as the quantum Thus, categorization of features is difficult to
leap in EM (Hay and Sandberg 1967). The first SEM standardize. An example is the category pollen size:
micrographs of pollen grains were published by there is not just a natural size variation within a single
Thornhill et al. (1965) and Erdtman and Dunbar anther/flower/taxon, dimensions may also vary
(1966). Since then palynologists have been pro- depending on the preparation method(s) used, and
vided with a plethora of beautiful micrographs. Like the observer’s evaluation. Moreover, sometimes the
Blackmore noted “The scanning electron micro- size of a pollen grain is just at the boundary between
scope has provided a greater impetus to palynol- two adjacent pollen size categories (for size catego-
ogy than any other technical development during ries: see “Pollen Morphology and Ultrastructure”).
the history of the subject.” Blackmore (1992). The LM When describing and categorizing pollen, two
with basic and advanced equipment, such as the basic groupings are known from the literature: pol-
fluorescent super-resolution microscopy, is over- len type and pollen class. Pollen type is a general
coming the Abbe limit of LM resolution (especially term categorizing pollen grains by a distinct combi-
STED m icroscopy, Hell 2009). The super-resolution LM nation of characters and is used in connection with
and the two main types of EM form an expedient systematics, affiliating the pollen type with a distinct
GENERAL CHAPTERS 9
PALYNOLOGY: HISTORY AND SYSTEMATIC ASPECTS
A B
C D
Fig. 5 Pollen type vs pollen class. A-B. Polygonum aviculare, Polygonaceae; all Polygonum pollen sharing the
combined features observed here belong to the Polygonum aviculare type. This pollen can also be included in the
pollen class “tricolporate”. C-D. Leontodon saxatilis, Asteraceae; all Asteraceae pollen sharing the combined features
observed here (lophate, tricolporate, echinate) belong to the Leontodon type, characteristic for the “Liguliflorae”
group within the Asteraceae. This pollen can also be included in the pollen class “tricolporate”
taxon/a (e.g., Polygonum aviculare type/Leontodon grain may belong to more than one pollen class;
type, Fig. 5). The term “pollen type” is sometimes in such cases, the more significant feature should
(colloquially) misused: for example, Croton type, be ranked first (e.g., Pistia: plicate-inaperturate,
which is a distinct feature of ornamentation and is Hemigraphis: plicate-colporate, Typha: tetrads-
correctly termed Croton pattern. ulcerate, Rhododendron: tetrads-colporate).
Pollen class is an artificial grouping of pollen Many terms in palynology were coined at a time
grains that share a single or more, distinctive char- when only LM observations were available. Mainly
acters (see “Illustrated Pollen Terms”). Pollen classes for historical reasons, inconsequent nomenclatural
can refer to pollen units (e.g., polyads, tetrads), to applications, enumerations of synonyms, and even
shape (e.g., saccate, polygonal, heteropolar, differing definitions have been found for one and
arcus), to aperture type and location (e.g., inaper- the same term. During the twentieth century,
turate, sulcate, ulcerate, colpate, colporate, porate, questions of terminology became more and more
synaperturate, spiraperturate), or to an extremely problematic. The main reasons were the increasing
distinctive ornamentation character (e.g., lophate, numbers of publications in palynology, dealing with
clypeate). These classes can be useful in identifica- sometimes insufficiently described or “uncommon”
tion keys as they have a good diagnostic, although pollen features, and simultaneously the advent of
mostly no systematic, value. In general, a pollen manifold applied fields of palynology. For various
10 GENERAL CHAPTERS
PALYNOLOGY: HISTORY AND SYSTEMATIC ASPECTS
A B
C D
Fig. 6 Nomenclature in Paleopalynology. A. Oculopollis sp., fossil, Upper Cretaceous, Hungary, polar view. B.
Oculopollis sp., fossil, Upper Cretaceous, Hungary, polar view. C. Trudopollis sp., fossil, Upper Cretaceous, Hungary,
polar view. D. Trudopollis sp., fossil, Upper Cretaceous, Hungary, polar view
reasons, nearly all authors used their own terminol- consistent use of drawings and the critical
ogy. Nonetheless, in the 1950s attempts were made comments on terms.
to restrict the wording and to state the definitions of A complex category issue in (Paleo-) Palynology
terms more precisely (Erdtman 1947; Erdtman and is the nomenclature question. In Paleopalynology,
Vishnu-Mittre 1956). A limited list of pollen morpho- for morphotaxa often form-generic names are
logical terms and definitions was published as early used. The nomenclature of form-genera is either
as 1950 by Iversen and Troels-Smith. Later, Kremp artificial when the relationship is not known at all
(1968), in his famous encyclopedia, provided a (e.g., Oculopollis and Trudopollis from the
monumental enumeration of all known terms. Normapolles group, Fig. 6), or semi-biological, when
Reitsma (1970) took the first resolute step to over- reference to an extant taxon is suspected but not
come the problem of synonyms in palynological proven (e.g., Liliacidites). However, if reference to
terminology, though unfortunately not taking into extant taxa is certain, then a biological nomencla-
account the variation range of palynological fea- ture is possible (e.g., Quercus sp.).
tures. Fægri and Iversen (1989, 4th ed.) restricted
their glossary of terms exclusively used in their book.
Moore et al. (1991, 2nd ed.) provided a glossary of The Turmal System
selected terms used in their pollen and spore keys.
Standardization came with the glossary by Punt A quite different classification and nomenclature is
et al. (1994, 2007). The main advance of their Potonié’s turmal system. This is an artificial, informal,
concise and comprehensive terminology is the
neutral suprageneric classification scheme for fossil
GENERAL CHAPTERS 11
PALYNOLOGY: HISTORY AND SYSTEMATIC ASPECTS
(especially Carboniferous or Permian) pollen and ifferent formula, separately. This system does not
d
spores. It is subdivided into a hierarchy of progres- work in, e.g., heteroaperturate or inaperturate (for-
sively finer units (ranks): anteturma, turma, sub- mula 000) pollen, or pollen tetrads. Unfortunately,
turma, infraturma, subinfraturma, and corresponds the NPC-system ignores other pollen characters
mostly to morphological features (for details see including shape and ornamentation that are indis-
Traverse 2007). pensable for a complete description.
A B
Fig. 7 NPC-classification of pollen. A-B. Androsace chamaejasme, Primulaceae, a tricolporate pollen with the
formula N3P4C5
12 GENERAL CHAPTERS
PALYNOLOGY: HISTORY AND SYSTEMATIC ASPECTS
processes and systematics. For taxonomic studies, Cenomanian (Doyle and Endress 2010; Friis et al.
pollen features that have value for the lower and 2011; Doyle 2012). For a detailed overview of struc-
higher taxonomic levels should be obtained by a tural pollen diversification and of the stratigraphic
combined study using LM, SEM, and TEM (Stuessy appearance of major angiosperm pollen types dur-
1979). ing the Cretaceous, see Friis et al. (2011) and
Alternation of generations is a unique feature of Mendes et al. (2014).
plants that occurs in green algae, mosses, ferns, Palynological data may be helpful at all levels of
gymnosperms, and angiosperms. Pollen grains systematics, especially in angiosperms (c.f. Stuessy
develop in anthers as the result of meiosis and mito- 2009). When pollen of a taxon (representing family/
ses (two in angiosperms, three to five in gymno- ies or genus/era) is characteristic and similar among
sperms) and represent an extra generation, the species they are termed stenopalynous (Fig. 8), and
highly reduced male gametophyte. Therefore, pol- occur, for example, in Poaceae, Lamiaceae,
len grains are not simply small parts of a plant like Asclepiadaceae, Brassicaceae, Asteroideae, and
leaves or seeds; they are the complete (hidden) Cichorioideae. On the contrary, eurypalynous
haploid counterpart to the more dominant plant, (Fig. 9) taxa are heterogeneous and pollen can
which represents the diploid generation (Kessler vary among others in size, aperture, and in exine
and Harley 2004). During dispersal, pollen grains are stratification. Examples for eurypalynous groups are
completely separated from the parent plant and Acanthaceae (Sarawichit 2012) and Araceae
perfectly adapted for their role — the transfer of (Harley and Baker 2001; Ulrich et al. 2017).
male genetic material — and are able to resist hos- At the highest taxonomic level (e.g., angiosperms
tile environmental stresses on their way to the vs gymnosperms, dicots vs monocots), a columellate
female flower parts. Usually, pollen does not suffer exine condition occurs exclusively in angiosperms. A
to the same extent from the various and harsh lamellate endexine is typical for gymnosperms,
selective pressures to which the diploid plant is sub- whereas the angiosperm endexine is usually not
jected. Because selective pressures (e.g., tempera- lamellate, except in immature stages (Orobanche
ture, precipitation) upon pollen characters are hederae). But in very few cases there is a continu-
predominantly absent or low, compared to those ously lamellate endexine present, like in Ambrosia
on the diploid plant, pollen features may remain (Furness and Rudall 1999a, b, Weber and Ulrich 2010).
constant for millions of years, meaning pollen fea- In inaperturate pollen of Araceae the endexine is
tures can be conservative and of taxonomic value exceptionally thick and spongy, which may be a
(Wodehouse 1928, 1935; Hao et al. 2001; Grímsson functional benefit and of systematic value. A strong
et al. 2014, 2016, 2017a, b). Therefore, identical and phylogenetic signal comes from the aperture
rare conditions in fossil vs recent pollen probably arrangement: the “tricolpate” condition is a synapo-
belong to only one group and were not invented morphy for eudicots, tricolporate pollen occurs only
independently in distant groups (e.g., fossil in core eudicots, while sulcate pollen is a plesiomor-
Spinizonocolpites pollen and recent Nypa pollen, phic condition in basal angiosperms (Nadot et al.
Arecaceae; Zetter and Hofmann 2001; Gee 2001). 2006). Palynologists have long wondered about the
Selective pressures might concern especially the two fundamental evolutionary shifts occurring at the
pollen aperture number, but also the pollen sculp- base of the eudicot clade, both in aperture position
ture and the mode of pollination ecology (Furness (from distal to equatorial) and number (from one to
and Rudall 2004). Pollen features are, if used for a three or more). Most probably, these changes in pol-
systematic purpose, at least as important as any len morphology have a systematic and simultane-
other morphological character of the diploid gen- ously a functional background. The shift from a distal,
eration. For this reason pollen morphology claims a single aperture to equatorially or globally situated
crucial role in, e.g., systematics and palynostratigra- apertures, increases the number of possible germina-
phy, for example in elucidating the early history of tion sites (Furness and Rudall 2004). Pollen morphol-
angiosperms. Angiosperm pollen from the Early ogy does not support sharp delimitation between
Cretaceous are usually sulcate (typical for basal dicots and monocots, as dicotyledonous pollen char-
angiosperms) with a columellate infratectum
acters also occur in some monocots and conversely.
(which is restricted to angiosperms). The first appear- In early-diverging angiosperms the formation of pol-
ance of dispersed tricolpate pollen, typical for eudi- len features appears to be more plastic than in dicots
cots, is not known before the latest Barremian, is (especially in eudicots). Manifold combinations of
rare in the Aptian of Southern Laurasia and Northern pollen features are typical for basal angiosperms and
Gondwana, but is ubiquitous in the Albian of both even for the most basal eudicots, the Ranunculales.
provinces. Tricolporate pollen appears first in the All of them are more or less e
urypalynous. In contrast,
late Albian, and triporate pollen in the middle late-divergent eudicots are often stenopalynous and
GENERAL CHAPTERS 13
PALYNOLOGY: HISTORY AND SYSTEMATIC ASPECTS
A B
C D
E F
Fig. 8 Stenopalynous taxa (family level). Pollen of different Poaceae all look very similar, for example in Alopecurus
(A), Cutandia (B), Dactylis (C), Fargesia (D), Poa (E), Sesleria (F) the pollen is spherical, ulcerate with nano-sized sculpture
elements
14 GENERAL CHAPTERS
Another random document with
no related content on Scribd:
telegrams protesting against the infliction of the death-penalty on a
woman.
One of the reasons which has been urged for the total abolition of
this penalty is the reluctance of juries to convict women of crimes
punishable by death. The number of wives who murder their
husbands, and of girls who murder their lovers, is a menace to
society. Our sympathetic tolerance of these crimes passionnés, the
sensational scenes in court, and the prompt acquittals which follow,
are a menace to law and justice. Better that their perpetrators should
be sent to prison, and suffer a few years of corrective discipline, until
soft-hearted sentimentalists circulate petitions, and secure their
pardon and release.
The right to be judged as men are judged is perhaps the only form
of equality which feminists fail to demand. Their attitude to their own
errata is well expressed in the solemn warning addressed by Mr.
Louis Untermeyer’s Eve to the Almighty,
as to the lenity shown them by men,—a lenity which they stand ever
ready to abuse. We have only to imagine what would have
happened to a group of men who had chosen to air a grievance by
picketing the White House, the speed with which they would have
been arrested, fined, dispersed, and forgotten, to realize the nature
of the tolerance granted to women. For months these female pickets
were unmolested. Money was subscribed to purchase for them
umbrellas and overshoes. The President, whom they were affronting,
sent them out coffee on cold mornings. It was only when their
utterances became treasonable, when they undertook to assure our
Russian visitors that Mr. Wilson and Mr. Root were deceiving Russia,
and to entreat these puzzled foreigners to help them free our nation,
that their sport was suppressed, and they became liable to arrest
and imprisonment.
Much censure was passed upon the unreasonable violence of
these women. The great body of American suffragists repudiated
their action, and the anti-suffragists used them to point stern morals
and adorn vivacious tales. But was it quite fair to permit them in the
beginning a liberty which would not have been accorded to men, and
which led inevitably to licence? Were they not treated as parents
sometimes treat children, allowing them to use bad language
because, “if you pay no attention to them, they will stop it of their
own accord”; and then, when they do not stop it, punishing them for
misbehaving before company? When a sympathetic gentleman
wrote to a not very sympathetic paper to say that the second Liberty
Loan would be more popular if Washington would “call off the dogs
of war on women,” he turned a flashlight upon the fathomless gulf
with which sentimentalism has divided the sexes. No one dreams of
calling policemen and magistrates “dogs of war” because they arrest
and punish men for disturbing the peace. If men claim the privileges
of citizenship, they are permitted to suffer its penalties.
A few years before the war, a rage for compiling useless statistics
swept over Europe and the United States. When it was at its height,
some active minds bethought them that children might be made to
bear their part in the guidance of the human race. Accordingly a
series of questions—some sensible and some foolish—were put to
English, German, and American school-children, and their
enlightening answers were given to the world. One of these
questions read: “Would you rather be a man or a woman, and why?”
Naturally this query was of concern only to little girls. No sane
educator would ask it of a boy. German pedagogues struck it off the
list. They said that to ask a child, “Would you rather be something
you must be, or something you cannot possibly be?” was both
foolish and useless. Interrogations concerning choice were of value
only when the will was a determining factor.
No such logical inference chilled the examiners’ zeal in this
inquisitive land. The question was asked and was answered. We
discovered, as a result, that a great many little American girls (a
minority, to be sure, but a respectable minority) were well content
with their sex; not because it had its duties and dignities, its
pleasures and exemptions; but because they plainly considered that
they were superior to little American boys, and were destined, when
grown up, to be superior to American men. One small New England
maiden wrote that she would rather be a woman because “Women
are always better than men in morals.” Another, because “Women
are of more use in the world.” A third, because “Women learn things
quicker than men, and have more intelligence.” And so on through
varying degrees of self-sufficiency.
These little girls, who had no need to echo the Scotchman’s
prayer, “Lord, gie us a gude conceit o’ ourselves!” were old maids in
the making. They had stamped upon them in their tender childhood
the hall-mark of the American spinster. “The most ordinary cause of
a single life,” says Bacon, “is liberty, especially in certain self-
pleasing and humorous minds.” But it is reserved for the American
woman to remain unmarried because she feels herself too valuable
to be entrusted to a husband’s keeping. Would it be possible in any
country save our own for a lady to write to a periodical, explaining
“Why I am an Old Maid,” and be paid coin of the realm for the
explanation? Would it be possible in any other country to hear such
a question as “Should the Gifted Woman Marry?” seriously asked,
and seriously answered? Would it be possible for any sane and
thoughtful woman who was not an American to consider even the
remote possibility of our spinsters becoming a detached class, who
shall form “the intellectual and economic élite of the sex, leaving
marriage and maternity to the less developed woman”? What has
become of the belief, as old as civilization, that marriage and
maternity are developing processes, forcing into flower a woman’s
latent faculties; and that the less-developed woman is inevitably the
woman who has escaped this keen and powerful stimulus? “Never,”
said Edmond de Goncourt, “has a virgin, young or old, produced a
work of art.” One makes allowance for the Latin point of view. And it
is possible that M. de Goncourt never read “Emma.”
There is a formidable lack of humour in the somewhat
contemptuous attitude of women, whose capabilities have not yet
been tested, toward men who stand responsible for the failures of
the world. It denotes, at home and abroad, a density not far removed
from dulness. In Mr. St. John Ervine’s depressing little drama, “Mixed
Marriage,” which the Dublin actors played in New York some years
ago, an old woman, presumed to be witty and wise, said to her son’s
betrothed: “Sure, I believe the Lord made Eve when He saw that
Adam could not take care of himself”; and the remark reflected
painfully upon the absence of that humorous sense which we used
to think was the birthright of Irishmen. The too obvious retort, which
nobody uttered, but which must have occurred to everybody’s mind,
was that if Eve had been designed as a care-taker, she had made a
shining failure of her job.
That astute Oriental, Sir Rabindranath Tagore, manifested a
wisdom beyond all praise in his recognition of American standards,
when addressing American audiences. As the hour for his departure
drew nigh, he was asked to write, and did write, a “Parting Wish for
the Women of America,” giving graceful expression to the sentiments
he knew he was expected to feel. The skill with which he modified
and popularized an alien point of view revealed the seasoned
lecturer. He told his readers that “God has sent woman to love the
world,” and to build up a “spiritual civilization.” He condoled with
them because they were “passing through great sufferings in this
callous age.” His heart bled for them, seeing that their hearts “are
broken every day, and victims are snatched from their arms to be
thrown under the car of material progress.” The Occidental sentiment
which regards man simply as an offspring, and a fatherless offspring
at that (no woman, says Olive Schreiner, could look upon a battle-
field without thinking, “So many mothers’ sons!”), came as naturally
to Sir Rabindranath as if he had been to the manner born. He was
content to see the passion and pain, the sorrow and heroism of men,
as reflections mirrored in a woman’s soul. The ingenious gentlemen
who dramatize Biblical narratives for the American stage, and who
are hampered at every step by the obtrusive masculinity of the East,
might find a sympathetic supporter in this accomplished and
accommodating Hindu.
The story of Joseph and his Brethren, for example, is perhaps the
best tale ever told the world,—a tale of adventure on a heroic scale,
with conflicting human emotions to give it poignancy and power. It
deals with pastoral simplicities, with the splendours of court, and with
the “high finance” which turned a free landholding people into
tenantry of the crown. It is a story of men, the only lady introduced
being a disedifying dea ex machina, whose popularity in Italian art
has perhaps blinded us to the brevity of her Biblical rôle. But when
this most dramatic narrative was cast into dramatic form, Joseph’s
splendid loyalty to his master, his cold and vigorous chastity, were
nullified by giving him an Egyptian sweetheart. Lawful marriage with
this young lady being his sole solicitude, the advances of Potiphar’s
wife were less of a temptation than an intrusion. The keynote of the
noble old tale was destroyed, to assure to woman her proper place
as the guardian of man’s integrity.
Still more radical was the treatment accorded to the parable of the
“Prodigal Son,” which was expanded into a pageant play, and acted
with a hardy realism permitted only to the strictly ethical drama. The
scriptural setting of the story was preserved, but its patriarchal
character was sacrificed to modern sentiment which refuses to be
interested in the relation of father and son. Therefore we beheld the
prodigal equipped with a mother and a trusting female cousin, who,
between them, put the poor old gentleman out of commission,
reducing him to his proper level of purveyor-in-ordinary to the
household. It was the prodigal’s mother who bade her reluctant
husband give their wilful son his portion. It was the prodigal’s mother
who watched for him from the house-top, and silenced the voice of
censure. It was the prodigal’s mother who welcomed his return, and
persuaded father and brother to receive him into favour. The whole
duty of man in that Syrian household was to obey the impelling word
of woman, and bestow blessings and bags of gold according to her
will.
The expansion of the maternal sentiment until it embraces, or
seeks to embrace, humanity, is the vision of the emotional, as
opposed to the intellectual, feminist. “The Mother State of which we
dream” offers no attraction to many plain and practical workers, and
is a veritable nightmare to others. “Woman,” writes an enthusiast in
the “Forum,” “means to be, not simply the mother of the individual,
but of society, of the State with its man-made institutions, of art and
science, of religion and morals. All life, physical and spiritual,
personal and social, needs to be mothered.”
“Needs to be mothered”! When men proffer this welter of
sentiment in the name of women, how is it possible to say
convincingly that the girl student standing at the gates of knowledge
is as humble-hearted as the boy; that she does not mean to mother
medicine, or architecture, or biology, any more than the girl in the
banker’s office means to mother finance? Her hopes for the future
are founded on the belief that fresh opportunities will meet a sure
response; but she does not, if she be sane, measure her untried
powers by any presumptive scale of valuation. She does not
consider the advantages which will accrue to medicine, biology, or
architecture by her entrance—as a woman—into any one of these
fields. Their need for her maternal ministration concerns her less
than her need for the magnificent heritage they present.
It has been said many times that the craving for material profit is
not instinctive in women. If it is not instinctive, it will be acquired,
because every legitimate incentive has its place in the progress of
the world. The demand that women shall be paid men’s wages for
men’s work may represent a desire for justice rather than a desire for
gain; but money fairly earned is sweet in the hand, and to the heart.
An open field, an even start, no handicap, no favours, and the same
goal for all. This is the worker’s dream of paradise. Women have
long known that lack of citizenship was an obstacle in their path.
Self-love has prompted them to overrate their imposed, and
underrate their inherent, disabilities. “Whenever you see a woman
getting a high salary, make up your mind that she is giving twice the
value received,” writes an irritable correspondent to the “Survey”;
and this pretension paralyzes effort. To be satisfied with ourselves is
to be at the end of our usefulness.
M. Émile Faguet, that most radical and least sentimental of French
feminists, would have opened wide to women every door of which
man holds the key. He would have given them every legal right and
burden which they are physically fitted to enjoy and to bear. He was
as unvexed by doubts as he was uncheered by illusions. He had no
more fear of the downfall of existing institutions than he had hope for
the regeneration of the world. The equality of men and women, as he
saw it, lay, not in their strength, but in their weakness; not in their
intelligence, but in their stupidity; not in their virtues, but in their
perversity. Yet there was no taint of pessimism in his rational refusal
to be deceived. No man saw more clearly, or recognized more justly,
the art with which his countrywomen have cemented and upheld a
social state at once flexible and orderly, enjoyable and inspiriting.
That they have been the allies, and not the rulers, of men in building
this fine fabric of civilization was also plain to his mind. Allies and
equals he held them, but nothing more. “La femme est parfaitement
l’égale de l’homme, mais elle n’est que son égale.”
Naturally to such a man the attitude of Americans toward women
was as unsympathetic as was the attitude of Dahomeyans. He did
not condemn it (possibly he did not condemn the Dahomeyans,
seeing that the civic and social ideals of France and Dahomey are in
no wise comparable); but he explained with careful emphasis that
the French woman, unlike her American sister, is not, and does not
desire to be, “un objet sacro-saint.” The reverence for women in the
United States he assumed to be a national trait, a sort of national
institution among a proud and patriotic people. “L’idolâtrie de la
femme est une chose américaine par excellence.”
The superlative complacency of American women is due largely to
the oratorical adulation of American men,—an adulation that has no
more substance than has the foam on beer. I have heard a
candidate for office tell his female audience that men are weak and
women are strong, that men are foolish and women are wise, that
men are shallow and women are deep, that men are submissive
tools whom women, the leaders of the race, must instruct to vote for
him. He did not believe a word that he said, and his hearers did not
believe that he believed it; yet the grossness of his flattery kept pace
with the hypocrisy of his self-depreciation. The few men present
wore an attitude of dejection, not unlike that of the little boy in
“Punch” who has been told that he is made of
The story goes that, after the bloody victory of the Scots under
Kenneth MacAlpine, in 860, only two Picts who knew the secret of
the brew survived the general slaughter. Some say they were father
and son, some say they were master and man. When they were
offered their lives in exchange for the recipe, the older captive said
he dared not reveal it while the younger lived, lest he be slain in
revenge. So the Scots tossed the lad into the sea, and waited
expectantly. Then the last of the Picts cried, “I only know!” and
leaped into the ocean and was drowned. It is a brave tale. One
wonders if a man would die to save the secret of making milk-toast.
From the pages of history the prohibition-bred youth may glean
much off-hand information about the wine which the wide world
made and drank at every stage of civilization and decay. If, after the
fashion of his kind, he eschews history, there are left to him
encyclopædias, with their wealth of detail, and their paucity of
intrinsic realities. Antiquarians also may be trusted to supply a
certain number of papers on “leather drinking-vessels,” and “toasts
of the old Scottish gentry.” But if the youth be one who browses
untethered in the lush fields of English literature, taking prose and
verse, fiction and fact, as he strays merrily along, what will he make
of the hilarious company in which he finds himself? What of Falstaff,
and the rascal, Autolycus, and of Sir Toby Belch, who propounded
the fatal query which has been answered in 1919? What of Herrick’s
“joy-sops,” and “capring wine,” and that simple and sincere
“Thanksgiving” hymn which takes cognizance of all mercies?
time it was, until the gilt began to wear off the gingerbread. But
Evelyn, though he feasted as became a loyal gentleman, and
admitted that canary carried to the West Indies and back for the
good of its health was “incomparably fine,” yet followed Saint
Chrysostom’s counsel. He drank, and compelled his household to
drink, with sobriety. There is real annoyance expressed in the diary
when he visits a hospitable neighbour, and his coachman is so well
entertained in the servants’ hall that he falls drunk from the box, and
cannot pick himself up again.
Poor Mr. Pepys was ill fitted by a churlish fate for the simple
pleasures that he craved. To him, as to many another Englishman,
wine was precious only because it promoted lively conversation. His
“debauches” (it pleased him to use that ominous word) were very
modest ones, for he was at all times prudent in his expenditures. But
claret gave him a headache, and Burgundy gave him the stone, and
late suppers, even of bread and butter and botargo, gave him
indigestion. Therefore he was always renouncing the alleviations of
life, only to be lured back by his incorrigible love of companionship.
There is a serio-comic quality in his story of the two bottles of wine
he sent for to give zest to his cousin Angler’s supper at the Rose
Tavern, and which were speedily emptied by his cousin Angler’s
friends: “And I had not the wit to let them know at table that it was I
who paid for them, and so I lost my thanks.”
If the young prohibitionist be light-hearted enough to read Dickens,
or imaginative enough to read Scott, or sardonic enough to read
Thackeray, he will find everybody engaged in the great business of
eating and drinking. It crowds love-making into a corner, being,
indeed, a pleasure which survives all tender dalliance, and restores
to the human mind sanity and content. I am convinced that if Mr.
Galsworthy’s characters ate and drank more, they would be less
obsessed by sex, and I wish they would try dining as a restorative.
The older novelists recognized this most expressive form of
realism, and knew that, to be accurate, they must project their minds
into the minds of their characters. It is because of their sympathy and
sincerity that we recall old Osborne’s eight-shilling Madeira, and Lord
Steyne’s White Hermitage, which Becky gave to Sir Pitt, and the
brandy-bottle clinking under her bed-clothes, and the runlet of canary
which the Holy Clerk of Copmanhurst found secreted conveniently in
his cell, and the choice purl which Dick Swiveller and the
Marchioness drank in Miss Sally Brass’s kitchen. We hear
Warrington’s great voice calling for beer, we smell the fragrant fumes
of burning rum and lemon-peel when Mr. Micawber brews punch, we
see the foam on the “Genuine Stunning” which the child David calls
for at the public house. No writer except Peacock treats his
characters, high and low, as royally as does Dickens; and Peacock,
although British publishers keep issuing his novels in new and
charming editions, is little read on this side of the sea. Moreover, he
is an advocate of strong drink, which is very reprehensible, and
deprives him of candour as completely as if he had been a
teetotaller. We feel and resent the bias of his mind; and although he
describes with humour that pleasant middle period, “after the
Jacquerie were down, and before the march of mind was up,” yet the
only one of his stories which is innocent of speciousness is “The
Misfortunes of Elphin.”
Now to the logically minded “The Misfortunes of Elphin” is a
temperance tract. The disaster which ruins the countryside is the
result of shameful drunkenness. The reproaches levelled by Prince