Historical Matters

Wodehouse (1935) presents a marvelous history of the study of extant pollen grains and spores, and this book should be consulted by those who need more information about the early history of palynology-proper = "pollen analysis." A more modern summary of the history of pollen studies is presented by Ducker and Knox (1985). The development of pollen research follows the history of plant anatomy and morphology in general and is dependent to a large degree on developmental stages of the microscope. Nehemiah Grew first observed pollen microscopically in Britain about 1640. Malpighi noted differences in size and color of pollen about the same time. Various people later studied the biology of pollen and spores, especially with reference to the function of pollen in fertilization of ovules, in the 18th century. Camerarius usually gets credit for proving the maleness and fertilizing function of pollen in the late 17th century. Curiously, ancient peoples knew what pollen was for, and such aboriginal people as American Indians have understood the maleness, and the precise function, of pollen, apparently for thousands of years: pollen played a prominent role in some Indian puberty ceremonies. Southwestern Indians seem also to have understood the dietary advantage of eating pollen, long before health-food stores began promoting it (see Fig. 1.4). The Indians perhaps noticed that various insects, especially beetles and hymenopterids, use pollen as a major food staple.

In the 19th century, with the coming of much improved microscopes, the anatomy of pollen and spores was carefully studied and catalogued by German scientists, e.g., von Mohl (d. 1872), Fritsche (d. 1871) and Fischer (pollen work published in 1889). Robert Brown noted in 1809 that pollen could be used to advantage for systematic studies of seed plants, and Brown's illustrator, F. Bauer, described 175 species of pollen for this purpose (see Graham and Barker, 1981). Pollen morphologists of today continue the work of Brown, Fritsche and Fischer, employing better optical microscopes, and especially scanning and transmission electron microscopes (SEM, TEM). Ultramicrotome techniques, in conjunction with transmission electron microscopes (TEM) have played a critical role in recent years in elucidating the internal structure of the exine. The study of spore/pollen morphology has had impact on paleopalynology, of course, but the history of paleopalynology is in practice a separate matter.

The first person to describe fossil spores/pollen, and figure them in line drawing, was apparently Goeppert in Germany in 1838. Alnus-like and Betula-like pollen are easily recognizable in Goeppert's plates. Ehrenberg (Fig. 1.5a),

Western Apache Writing Symbol:

"He who is decorated with and enriched by pollen"

Western Apache Writing Symbol:

"He who is decorated with and enriched by pollen"

Figure 1.4 The vital, dynamic nature of pollen has long been recognized by humans in many cultures. The symbol shown above, from Silas John's Western Apache writing system (Basso and Anderson, 1973) bespeaks the ceremonial use of pollen by American Indians, symbolizing fertility, among other things. The pollen used by Silas John (a shaman) was apparently Typha, although the symbol would seem to represent a Zea tassel, the pollen of which was and is also used ceremonially by Indians. The jars illustrate the widespread modern use of pollen as a dietary supplement. Such pollen is sometimes harvested by human vacuum cleaners, sometimes taken from domesticated bees, which use pollen for hive nutrition. The human use is based on the vitamin, mineral, and nutritive (lipids, carbohydrates, amino acids) content of pollen. Apiaries sell untreated bee leg loads stripped from worker bees by a device at the hive entrance-hence the variously colored blobs in the jar, each being one leg-load. "Vibrant Health Bee Pollen" is compressed into tablets, for a considerable elevation in price per gram.

who pioneered almost everything in micropaleontology, certainly also saw them, and described what we later were to discover are acritarchs and dinoflagellate cysts in the 1830s. By 1867, Schenck was illustrating with good line drawings the in situ fossil spores he removed from fossil fern compressions. Reinsch in 1884 published the first photomicrograph of a fossil spore. It was of Carboniferous age, and the genus to which it belonged was long afterwards named, in his honor, Reinschospora, by Schopf et al. (1944). Bennie and Kidston published in 1886 descriptions of megaspores from the Carboniferous of Scotland. Actually, that is about where the paleopalynological matter ended for many decades, with somewhat peripheral exceptions, such as studies in the early 1900s by Thiessen of spores seen in Pennsylvanian coal thin-sections, and unpublished studies by Wodehouse of about the same time of pollen and spores in thin-sections of the Paleogene Green River Oil Shales.

However, the investigation of possible use of fossil (as some would say, "subfossil") spores/pollen in investigation of "post-glacial" or "Holocene" ("present interglacial" would perhaps be a preferable term for the last approximately 10,000 years) sediments went off rather independently, more or less unheeded by paleobotanists, about 1900. Details of this story are related by Erdtman (1954). A Swedish botanist, Lagerheim, realized that the pollen in, e.g., peats of Sweden, told the story of the vegetation in the vicinity of the

Figure 1.5 Important founding figures of paleopalynology: (a) Christian Gottfried Ehrenberg, 1795-1876; (b) E.J. Lennart Von Post, 1884-195l; (c) left, Gunnar Erdtman, 1897-1973; right, William S. Hoffmeister, 1901-80. From Saxony in Germany, Ehrenberg was originally a mycologist, but in 1837 he presented a paper to the Berlin Academy of Science, in which most of the major categories of what we now call palynomorphs

Figure 1.5 Important founding figures of paleopalynology: (a) Christian Gottfried Ehrenberg, 1795-1876; (b) E.J. Lennart Von Post, 1884-195l; (c) left, Gunnar Erdtman, 1897-1973; right, William S. Hoffmeister, 1901-80. From Saxony in Germany, Ehrenberg was originally a mycologist, but in 1837 he presented a paper to the Berlin Academy of Science, in which most of the major categories of what we now call palynomorphs peat deposition, and he published some brief notes on preliminary studies based on rather primitive pollen spectra. Lagerheim himself depended on previous work based on geological and paleobotanical studies of Blytt, Sernander, and others, showing that vegetational changes marked the climatic history of the latest Neogene (Faegri 1974, 1981). It remained, however, for a Swedish assistant and student of Sernander, a protégé of Lagerheim, Lennart Von Post (see Fig. 1.5b), to put the subject on a sound footing with thorough studies of a number of cored sequences of Holocene peat in Sweden. The publication of Von Post's dissertation work in 1916 is usually accepted as the beginning of pollen-analysis or pollen-statistics, as such studies came to be known. Von Post was apparently not as versatile in languages as some Scandinavians (he published only in Swedish and German), and did not widely popularize the new subject outside of Scandinavia. That task fell to the one man who has probably most influenced the subject, Von Post's doctoral student, Gunnar Erdtman (Figs. 1.5c and 1.6a). A very gifted musician (flutist) and surrealist artist (see frontispiece "self-portrait" sketch in Fig. 1.6a of pollen analyst with flute!), Erdtman was urbane and very skilled at languages. (He was, in fact, so good at English and so confident of his talents that he would argue nuances of the language even with well educated, native speakers of English such as this writer! He was often, but not always, right!) He traveled widely, indeed he loved to travel, and wherever he went he practiced and "sold" palynology, e.g., trapping pollen grains with a vacuum cleaner on a transatlantic cruise. He was responsible for a great expansion of pollen-analytical/statistical studies in many parts of the world in the 1920s, 1930s and 1940s. The terminology he developed for pollen morphology came to be dominant, partly because of the pre-existing void, partly because of his talent for coining new terminology, sometimes even in anticipation of the discovery of features not yet found! Also, <-

Figure 1.5 were described. The science of palynology is usually reckoned as commencing in 1916 with the introduction by Von Post of analytical pollen diagrams for post-glacial sediments. Von Post, a Swede, was developing ideas actually pioneered earlier by Gustav Lagerheim and others in Scandinavia. Von Post did not publicize pollen analysis much outside of Scandinavia, partly for linguistic reasons.

Erdtman, a younger one-time associate of Von Post's in Sweden, was urbane, fluent in various languages; he traveled very widely, evangelizing for the new scientific method. To Erdtman should go most of the credit for putting fossil spores/pollen studies on the map, over much of the world. He was very conscious of his contribution and would not have liked appearing beneath Von Post here, as Erdtman felt Von Post never sufficiently recognized his work. Hoffmeister, a paleontologist for Esso (Exxon), more than any other one person in industry was responsible for recognition that palynomorphs were a critically important group of microfossils that could be used for practical correlation, where other microfossils failed or were less satisfactory. (a) reproduced from Sarjeant, 1978, courtesy of AASP Foundation; (b) reprinted by permission from Erdtman, 1954; (c) courtesy of W. R. Evitt.

Jan Muller Palynologist

Figure 1.6 A few guiding spirits, and one poltergeist, of paleopalynology: (a) Gunnar Erdtman's (1897-1973) self-portrait as a wood-gnome. One must not deduce from this bit of whimsy that Erdtman was jovial and light-hearted. He could excoriate a younger palynologist for such a self-defined infraction as publishing a trilete spore photo with no radius of the laesura pointing up; (b) Knut Faegri (1909-2001), a keen student of pollination mechanisms, was the leading Norwegian Pleistocene-Holocene palynologist for decades. Though not primarily a paleopalynologist, he especially contributed to paleopalynology by his justly famed Textbook of Pollen Analysis (the first edition co-authored by J. Iversen);

Figure 1.6 A few guiding spirits, and one poltergeist, of paleopalynology: (a) Gunnar Erdtman's (1897-1973) self-portrait as a wood-gnome. One must not deduce from this bit of whimsy that Erdtman was jovial and light-hearted. He could excoriate a younger palynologist for such a self-defined infraction as publishing a trilete spore photo with no radius of the laesura pointing up; (b) Knut Faegri (1909-2001), a keen student of pollination mechanisms, was the leading Norwegian Pleistocene-Holocene palynologist for decades. Though not primarily a paleopalynologist, he especially contributed to paleopalynology by his justly famed Textbook of Pollen Analysis (the first edition co-authored by J. Iversen);

his prolific production of publications served to popularize and establish his ideas. In his later years he became rather intolerant of what he regarded as deviant, i.e., non-erdtmanian, practices in spore/pollen work and frequently wrote letters to errant authors, dressing them down for their sorry ways. (I have such a letter scolding me for orienting spores/pollen upside down: the laesura ray of a trilete laesura must point up at 90', the long axis of bisaccate pollen must be parallel to the bottom of the page with the distal side up, etc.) Nevertheless, it must be acknowledged that Erdtman's impact on palynology is unequaled, and we all owe his memory a great debt.

After Von Post and Erdtman, came many other Holocene pollen analysts in Scandinavia such as Iversen, Jessen, and Faegri (see Fig. 1.6b) but also Godwin in Britain, Sears and Potzger in America, Neustadt in the (former) USSR, Firbas in Germany, and a long list of others. But from the time of Erdtman and Von Post, this subject, linked to plant geography, ecology generally, paleoclimatology, and archeology-anthropology, tended to go its own way, as it still does today. Most of the practitioners are botanically rather than geologically oriented, and the field and laboratory techniques for taking cores of the relatively shallow, usually unindurated post-glacial sediments and processing them to obtain the usually well preserved palynofloras are more or less special to Holocene palynologists. The pollen/spore types studied are 100% extant species.

Unfortunately, because of all these factors, and other, more personal ones, there tends to be little contact between pollen-analysts and paleopalynologists. This was not always the case, and some pioneer paleopalynologists, such as A. Raistrick in Britain, L. R. Wilson (Fig. 1.6c) in the USA, and others, worked in both Pleistocene/ Holocene pollen analysis and with palynomorphs as old as Paleozoic.

As already noted, coal petrologists studied spores/pollen seen in coal thin-sections long ago, and the American coal petrologist, Thiessen, early in the 20th century even suggested the use of spores for coal-bed stratigraphy. Raistrick, <-

Figure 1.6 (c) Leonard R. Wilson (1906-1998), one of the first American palynologists (usually called "Dick" or "Doc") published papers on what we would now call paleopaly-nology soon after Erdtman's visits to North America in the 1920s; (d) James M. Schopf (1911-78), American paleobotanist, coal petrologist and paleopalynologist, made very important early contributions to establishing the systematic study of palynomorphs on a sound basis. For example, Schopf, et al. (1944), largely his work, became a model for such studies; (e) Robert Potonie (1899-1974), son of the well-known German paleobotanist, Henri Potonie, was one of the first people to recognize and apply the stratigraphic possibilities of paleopalynology, especially in the German coal fields and, through students and associates, elsewhere. After World War II, Potonie's encyclopedic studies of fossil spores/pollen, including introduction of his suprageneric "turmal" system of classification, had great importance in emphasizing the potential for systematic work in the field. (a) reprinted by permission from Erdtman, 1954.

in Britain in the 1920s, was a pioneer in the use of spores for this important task (see Chaloner, 1968a; Marshall, 2005). Unfortunately, coal beds are very difficult to correlate by spores because coal is primarily derived from woody swamp peat. The spore/pollen flora of such a sediment is notoriously local in derivation, e.g., as compared to deltaic silts, which have a rich, fluvially derived pollen flora representing a large area. Also the palynofloras of coals of an area usually represent a persistent biofacies that tends to recur mostly in response to the environment. Thus, within a given time frame, it is not always possible to correlate coal beds by correlating the facies, as almost identical palynofloras may occur in widely separated horizons. Nevertheless, because the "original" palynologists were the Holocene pollen analysts who preferred to work with post-glacial peats, the idea persisted for decades that paleopalynologists should be looking at fossil peats, that is, coals. Even now it is hard to convince some field geologists that it is usually better to collect the associated shales than coals for palynology, and that presence of plant megafossils in a shale does not necessarily correlate with presence of palynomorphs! It should be noted that some palynologists (Kosanke, 1950; Smith and Butterworth, 1967; Peppers, 1996) have successfully correlated coal beds by their spore-content, despite the attendant difficulties.

In the late 1920s and early 1930s, Robert Potonié (Fig. 1.6e), son of the paleobotanist, Henri Potonié, began to study spores/pollen from German coals and associated sediments, especially at first the Cenozoic lignitic coals, but later the Carboniferous coals as well. Potonié and his students and coworkers made a very significant contribution to paleopalynology with systematic and biostratigraphic studies. Just before the outbreak of World War II, in the late 1930s, Potonié was engaged by the Royal Dutch/Shell petroleum interests to investigate the possibility of using palynology as a biostratigraphic tool. The war ended that, but soon after the war Shell remembered Potonié's work and began palyno-logical research in earnest, employing especially Dutch palynologists such as Waterbolk, whose ultimate palynological roots were in Holocene pollen analysis. Other, more geologically oriented persons such as Kuyl were soon involved, however, and even such botanists as Jan Muller were geologically adept enough to assure a sound geological approach. By the early 1950s, Shell had employed paleopalynology very successfully in the Maracaibo Basin in Venezuela, where marine micropaleontology (= study of foraminifera and ostracodes mostly) was not fully satisfactory because of the extensive non-marine sections (see Kuyl et al., 1955). In North America, Esso (Standard Oil of New Jersey = Exxon) also began looking at palynology as a biostratigraphic tool quite early. By the late 1940s, they also had a laboratory in Venezuela under R. H. Tschudy, but the center of their palynological operation soon shifted to Oklahoma, where W. S. Hoffmeister (Fig. 1.5c), a micropaleontologist working for Esso's research subsidiary, Carter Oil Co., and L. R. Wilson (Fig. 1.6c), an academic palynologist working as a consultant, together developed Esso's palynostratigraphic program. American paleopalynology owes a great debt also to J. M. Schopf (Fig. 1.6d), who started his career with the Illinois Geological Survey and later served with the U. S. Bureau of Mines and the U. S. Geological Survey. He recognized in the early 1930s the great potential of spores/pollen studies for solution of geologic problems. The classic work of Schopf et al. (1944) was one flowering of Schopf's pioneer efforts. By 1955 when I joined Shell Development Company (a research subsidiary of the Royal Dutch/Shell group of companies) as a palynol-ogist, Shell's palynological operations were worldwide, from Nigeria to western Canada. About the same time, nearly all the other oil companies in the world of any size introduced palynology, at least into their research programs. Esso (now Exxon-Mobil) and the Shell group were in the thirties and remain today the giants of the oil industry worldwide, and the early use by them of palynology for stratigraphic and paleoecological purposes was in my opinion pivotal, though others also contributed, such as J. W. Durham, through his connection with oil operations in the 1940s in Colombia (see Langenheim, 1989). The 1950s were the time of greatest expansion of the subject.

A treatment such as this cannot claim to be complete, and I clearly should discuss at length the important and early contributions of the many Soviet (mostly Russian) paleopalynologists: Naumova, Bolkhovitina, Zaklinskaya, and many others. Near the end of the 20th century the Chinese also began to be heard from in large numbers and with many publications. In the English-speaking world, probably no other one institution has had the impact of the University of Sheffield, under L. R. Moore, Charles Downie, and their many students, now practicing the profession all over the world (see Sarjeant, 1984). My treatment of the subject also tends to emphasize the pollen and spore part of the subject more than the "non-spore" aspects. Sarjeant (1998) published an important summary of his extensive contacts with paleopalynological leaders in the study of acritarchs and dinoflagellates, including photographs of many of them.

As far as I am aware, my 1951 Ph.D. dissertation on Paleogene spores/pollen was the first one dealing exclusively with paleopalynology in North America; it certainly was one of the first, but I never had a course specifically in palynology because there were none at Harvard or Cambridge Universities. However, Sir Harry Godwin's ecology course at Cambridge, England, and E. S. Barghoorn's paleobotany course at Harvard, both of which I took, contained some of what is now called paleopalynology. By the 1960s, Ph.Ds in palynology were common, and by 1981, the American Association of Stratigraphic Palynologists had over 800 members. There were something on the order of 3,000 professional paleopa-lynologists in the world at that time (see Traverse, 1974a, for a history of palynology to 1972). When I was working on my doctoral dissertation in the late 1940s, there was no journal specifically for palynology, and the relatively manageable number of publications in the subject appeared in a wide variety of journals. There was an informally mimeographed newsletter about pollen/spore research in the 1940s called the Pollen Analysis Circular, later the Pollen and Spore Circular, edited and distributed by a Holocene pollen analyst, P. B. Sears

(in the later numbers assisted by L. R. Wilson). It was chiefly intended to inform "friends of pollen" about new developments, but it has achieved lasting recognition, mostly because of the fortuitous circumstance that the word "palynology" was coined by Hyde and Williams in No. 8, October, 1944. (As of mid-2005, the Center for Environmental Science, Northern Arizona University, made it possible to download from the Internet all of the numbers of the PSC. See: www.envsci.nau.edu/faculty/ScottAnderson/Pollen & Spore Circulars.htm. Then click on the desired circular.) Erdtman's Grana Palynologica (now Grana) began in 1954, and the French Pollen et Spores (now moribund) in 1959. There are quite a number of other journals worldwide (see Annotated Bibliography at the end of this chapter). It is now impossible to "stay on top" of the whole palyno-logical literature, even with the aid of the many society newsletters and bibliographies (see abovementioned bibliography). This is dramatized by the explosion in numbers of contributions and pages now being published primarily in Chinese, and requiring those of us who read only languages in greco-Roman alphabets and their derivatives, to wait for translations.

The work of Evitt, Wall, Williams, Norris, Fensome, Edwards, Wood and many others, beginning in the early 1960s, put a new face on paleopalynology by showing that dinoflagellates (apparently almost always their cysts) can be used biostratigraphically. Evitt (Fig. 12.4a) has had an especially important impact on this work. Dinoflagellate cysts are in the same size-range as spores/pollen, and apparently possess a resistant sporopollenin framework in their walls, as do spores/pollen. Most (but by no means all) of the cyst-producing dinoflagellates are marine, and their study has greatly expanded the usefulness of paleopalynology by providing more control for marine rock sequences where spores/pollen may be rare or absent, and because dinoflagellate cysts are often far better chronos-tratigraphic indicators. The pioneer in this field was G. Deflandre (Ehrenberg, as noted above, had seen dinoflagellate cysts in the 1840s), when palynology was in its infancy in the 1930s. However, it was the proof that many of what had been known as "hystrichosphaerids" are in part dinoflagellates that opened up this field. Dinoflagellate palynologists mostly started as spore/pollen palynologists, and perhaps this and the fact that spores/pollen and dinoflagellate cysts occur in the same preparations is responsible for keeping spore/pollen and dinoflagellate people closely allied. The "hystrichosphaerids" which were not transferred to the dinoflagellates were then recognized as acritarchs ("unknown" origin), following a proposal of W. R. Evitt. The study of these, mostly algal cysts or reproductive bodies, was pioneered especially by Alfred Eisenack (1891-1982; see Fig. 12.4d) in Germany, by Charles Downie (1923-1999; see Fig. 6.4a) in England, and by F. H. Cramer and M. d. Carmen R. Diez (Fig. 6.4b), working primarily in Spain and the USA. In recent years, the publications of Fensome, Wood, and Wicander have been especially important.

Pleistocene (including Holocene, as used here) pollen analysis researchers tend to go their own way, now as before. Their orientation has always been ecological, paleoclimatological, and archeological, and still is. Paleopalynologists have been oriented to biostratigraphy and especially to its economic application. Paleopaly-nology owes its origins to present interglacial ("post-glacial" = Holocene) pollen analysis, and even the very name "palynology" to exponents thereof, but very few paleopalynologists have managed to remain closely associated with Pleistocene pollen analysis as well.

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