Kapp, R. O., O. K. Davis, and J. E. King, 2000. Illustrated by R. C. Hall.

Reviewed by Alwynne B. Beaudoin
Provincial Museum of Alberta, Edmonton, Alberta

It is over a quarter of a century since I first sat down at a microscope to try and do pollen analysis. It was a sunny September afternoon. I well remember the feeling of bewilderment verging on panic as I peered through the eyepieces and saw the array of peculiar objects on the microscope slide. What was all this stuff? Which bits were the pollen grains? How on earth was I ever going to learn how to identify these things? In these moments of alarm, when I could see my prospects of completing a thesis disappearing into the mists of impossibility, Kapp's "Little Red Book" became a constant companion. Indeed, it might almost have had those comforting words, "Don't Panic", inscribed on the cover. In authoritative terms, with clear line drawings and easy-to-follow keys, it imposed order on a complex subject. By diligent study and hundreds of hours at the microscope, I gradually came to recognize a few of the palynomorphs that I saw, and greet the appearance of yet another pine grain as an old friend.

I still have my original copy of Kapp's book, entitled How to Know Pollen and Spores. Now it is rather battered, the pages are yellowed and extensively annotated, but the cover remains intact and I still consult it regularly. It has probably been one of the books I have used most in my career. I remain amazed at the amount of work that went into the book's development, especially because it was produced long before the era of desktop computers with word processing and database software. Through this book, Kapp, who died in 1990, made an outstanding and continuing contribution to the discipline. Originally published in 1969, it has been out of print for years. Every time I teach, I recommend it to students. But, other than chance finds in second-hand bookstores, it has not been readily available to new generations of novice palynologists. So I was pleased to learn that AASP was publishing a new edition. The preparation of the second edition was undertaken by two senior palynologists with excellent credentials for the task - James E. King, retired Director of the Cleveland Museum of Natural History, and Owen K. Davis, Department of Geosciences, University of Arizona, and past President of the International Federation of Palynological Societies.

The new edition retains many of the features of the old version. It has the same size and layout, with a similar card cover and coil binding so it lies flat on the laboratory bench. The organization of the book is the same, with taxa arranged according to major morphotypes (e.g., tricolpate, periporate, etc.), as they are encountered under the microscope. The book covers the same geographic area (temperate and boreal North America). Because it focusses primarily on "pollen and spore types routinely encountered by pollen analytical workers" (p. 5), it features mainly types that are wind-transported. Many common tree and shrub taxa are included. Temporally, it focusses on Quaternary palynology, that is, pollen and spores that can be extracted from non-lithified sediment or collected by air sampling and that can be referred to extant plant taxa. The original illustrations have been included. So what are the differences between this and the earlier edition? Overall, differences are comparatively minor, mainly relating to some rearrangement or expansion of the keys, with a few new illustrations and descriptions, and some updating.

The introduction and opening sections (including discussions of pollen formation and function, structure, preparation and sampling techniques) have been revised and updated, reflecting some of the research and including references to some regional pollen and spore guides that have appeared since 1969. Of the 82 references in this edition, only 12 have carried over from the first. These sections remain, however, a "once over lightly" view of their subjects. The sample preparation and sediment sampling sections are particularly scant. The book is focussed on identification of palynomorphs from unlithified sediments, yet the extraction of palynomorphs from coal is still outlined, although other more relevant techniques are not. I would have liked to see a broader reference list here, for example, inclusion of Cushing and Wright (1965), with specifications for the Livingstone sampler, the useful review article by Wright (1991) on coring, and the extensive discussion of preparation techniques for Quaternary sediments by Moore et al. (1991), a widely-used textbook. Undoubtedly, each of these sections could have formed the basis for an entire chapter! And this is not a "What is Pollen" or "How to do Palynology" manual. But I was disappointed that Davis and King did not take the opportunity to revise these sections more comprehensively.

The heart of the book remains, however, the drawings and accompanying descriptions. The drawings are deceptively simple in appearance, yet remain the most useful ones of pollen that you are likely to find. In many instances, the illustrations are reproduced somewhat larger than in the first edition. My impression too is that the font is slightly larger. In some cases the reproduction of the drawings is crisp and clear. Overall, however, the reproduction is so light that parts of drawings have disappeared. This has especially affected those drawings that have extensive shading. The reproduction is often so poor that the critical characteristics cannot be distinguished. These cases include Trifolium pratense (p. 110) where the reticulate surface cannot be seen, the equatorial view of Glandularia bipinnatifida (p. 115) where almost all features of the complex colpus are not visible, the image of Prunus virginiana pollen (p. 122) on which the striae and colpus can barely be distinguished, and Ptelea trifoliata (p. 155) on which colpus and pore details cannot be seen. The generally poor reproduction of many drawings is by far the least satisfactory aspect of this book, and must be disheartening for Davis and King.

Going through the book section by section, some changes are evident. The first section, showing objects that might be confused with pollen, has been considerably enhanced and expanded. It now describes 20 types, including algae, dinoflagellates, and phytoliths, as opposed to 13 in the first edition. Here, however, it would have been useful to have a reference to work of Bas van Geel (see, for example, 1978) who has produced very helpful illustrated documentation of the "other stuff" that may be found in pollen preparations.

This is followed by a "Key to Major Spore and Pollen Groups", which divides palynomorphs into categories based on readily observable morphological features (number of colpi, number of pores, etc.). These categories form the basis for the remaining sections of the book, with subtleties of surface features and wall structure used to distinguish the various types. The use of morphotypes as an initial organizational method provides a sound and usable framework. Within each section, dichotomous keys guide the user to the most likely pollen types. The keys focus on features that can usually be seen even in poorly preserved material.

The Dicolpate (3 types), Stephanocolpate (8 types), Pericolpate (3 types), Heterocolpate (1 type), Diporate (4 types), Triporate (27 types), Stephanoporate (12 types), Periporate (23 types), Dyad (3 types), and Lophate and Fenestrate (5 types) sections cover the same taxa as before. The Stephanocolporate section (4 types) has been expanded with an entry for Sanguisorba canadensis. In the Tetrad section (14 types), the illustration for Asmina triloba (p. 217) is a duplicate of one of the drawings of Berberis fremontii (p.102). The discussion of the complexities of ericaceous pollen would have been enhanced by reference to Warner and Chinappa (1986) and Cwynar (1982). An entry for Sporormiella has been included in the Polyad section (15 types), though the name is misspelt in the caption and description. The accompanying key has been rearranged accordingly.

The Vesiculate section (11 types) has been expanded slightly, with a distinction made between Diploxylon and Haploxylon pine pollen and the addition of an entry for Tsuga mertensiana. These revisions will be helpful to people working with material from western Canada. The Polyplicate section (5 types) includes one new illustration and description (Ephedra torreyana). The Trilete section (24 types) covers the same taxa as before. Most choices in the keys are based on measurable or observable characteristics, things that you can actually see under the microscope. However, in the Trilete section, the distinction between Cyathidites minor and Sphagnum is still made on the basis of age (Cretaceous vs. Recent). This always puzzled me in the first edition and I am still unclear as to how you are supposed to know the age just from looking at the spore! The discussion of Selaginella (p. 53), focusses on S. kraussiana, an introduced species, but it does now mention S. selaginoides, an indigenous species. The latter is not uncommon in some pollen preparations from suitable areas, such as the Rockies in Alberta.

The Inaperturate section (46 types) includes many fungal spores, which can be quite common in some preparations. Here, the Cupressaceae category has been subdivided on the basis of size and the arrangement of verrucae and gemmae. These descriptions have also been expanded. Some descriptions (e.g., for Taxodium) in the Monoporate section (15 types) have been expanded. In the Monocolpate/Monolete section (36 types), there is a re-arrangement of categories at the initial level, with addition of Isoëtes muricata, and, towards the end of the key, re-arrangement and addition of Brasenia schreiberi.

Students struggling to identify pollen grains will spend much time perusing the Tricolpate (52 types) and Tricolporate (94 types) sections. Together, these sections dominate the book, in line with the frequency of these pollen types in many pollen preparations. Often, especially when the material is poorly preserved, just figuring out whether a "mystery grain" is tricolpate or tricolporate can be a difficult task! Indeed, some pollen types are keyed out in both categories.

The Tricolpate section is where I noticed most changes. Initial parts of the key (dealing with Salix pollen) have been re-arranged. In the course of this, the illustration and description of Catalpa pollen appear to have been dropped. However, there is still an index entry for Catalpa, which gives its page assignment as p. 97, where it appeared in the first edition. The section includes a new entry for Pedicularis (p. 114). In the Tricolporate section (94 types), the illustrations for Philadelphus inodorum and Mitchella repens appear to be reversed (pp. 150-151) in comparison with the first edition, i.e., the Philadelphus entry has the Mitchella drawing and vice versa.

Throughout, the descriptions have been re-arranged slightly, with the taxa identified first by their Linnean rather than their common names. Family names have also been included for most entries. Parts of the nomenclature have been updated. Some taxa have been re-named between the first and second editions. For example, Yucca mohavensis is now Y. shidera (p. 93), and Verbena ciliata is now Glandularia bipinnatifica (p. 115) although the illustrated specimens are the same as in the first edition. Trillium flexipes and Polygonatum canaliculatum have been reassigned from the Liliaceae to the Trilliaceae and Convallariaceae respectively. On the other hand, Davis and King have mostly kept the same family names as in the first edition (Gramineae, Leguminosae, Compositae, etc.), despite changes in botanical nomenclature (Poaceae, Fabaceae, Asteraceae, respectively). This is a thorny issue. Sticking to superceded nomenclature lowers the credibility of palynology in the eyes of botanical purists. Yet, these names are well-entrenched in the literature. Still, it would be helpful to have included equivalencies for families in the taxonomic listing (pp. 235 - 256) since this is how they can be found in the current botanical or ecological literature. The source followed for the plant taxonomy should also have been cited.

A geological time chart - a useful new addition - follows the descriptions, references, and taxonomic lists. However, there is no source given for this chart. The glossary (pp. 258 - 269) is enhanced and is now separate from the index, making it a valuable stand-alone feature. For detail on some of these entries, however, Punt et al. (1994) will be found a useful supplement.

Regrettably, the text contains distracting spelling and typographical errors. There are some inconsistencies between names in headings, descriptions, and captions. For example, the genus Clarkia has been changed in the heading (p. 190), though the description refers to the same genus as the first edition (Godetia); Shrankia occidentalis has been renamed Mimosa quadrivalvis in the header and text but not the figure caption (p. 221); Rhus glabra appears as Rush glabra in the figure caption (p. 154). One amusing textual infelicity has been carried over from the first edition: "From the earliest days of pollen analysis, there has been great interest in boring [sic] peat bogs and lakes" (p. 23). A rather revealing slip perhaps!

Davis and King are to be commended for undertaking a difficult task, to take a classic and prepare it for a new edition. Clearly, one of the decisions they had to make was how much to revise and update it. As they point out in their introduction, the pollen types haven't changed over thirty years, and the heart of the book has worn well. Although I would have liked to see more extensive revision of some of the ancillary information, this does not detract from the usefulness of the book as a whole. It remains a valuable resource and I have no hesitation in recommending it. Indeed, I am delighted to see it back in print. With its modest price, it is within reach of students as a textbook. It should become a fixture next to their microscopes. I do urge the AASP Foundation, however, to undertake a second printing and try to improve the quality of the images and correct the text. This single step would turn a good book into a great one.

References

Cushing, E. J., and H. E. Wright Jr. (1965) Hand-operated Piston Corers for Lake Sediments. Ecology 46:380-384.

Cwynar, L. C. (1982) A Late-Quaternary Vegetation History from Hanging Lake, Northern Yukon. Ecological Monographs 52:1-24

Moore, P. D., J. A. Webb, and M. E. Collinson (1991) Pollen Analysis. 2nd edition. Blackwell Scientific Publications, Oxford, England, U.K. 216 pages

Punt, W., S. Blackmore, S. Nilsson, and A. Le Thomas (1994) Glossary of Pollen and Spore Terminology. LPP Contributions Series No. 1. LPP Foundation, Utrecht.

Van Geel, B. (1978) A Palaeoecological Study of Holocene Bog Sections in Germany and the Netherlands, Based on the Analysis of Pollen, Spores, and Macro-microscopic remains of Fungi, Algae, Comophytes and Animals. Review of Palaeobotany and Palynology 25:1-120.

Warner, B. G., and C. C. Chinnappa (1986) Taxonomic Implications and Evolutionary Trends in Pollen of Canadian Ericales. Canadian Journal of Botany 64:3113-3126

Wright Jr, H. E. (1991) Coring Tips. Journal of Paleolimnology 6: 37-49

To Order

To order a copy of this book, please contact V. M. Bryant, Anthropology Department, Texas A&M University, College Station, Texas 77843, USA, E-mail: vbryant@tamu.edu