) Madrono, Vol. 47, No. 4, pp. 253-258, 2000 FLORISTIC STUDIES IN CONTEMPORARY BOTANY Theodore M. Barkley Botanical Research Institute of Texas, 509 Pecan Street, Fort Worth, TX 76102-4060 Abstract This paper outlines the traditional procedures for monographic and floristic studies, and points out that floristic studies are a Hnk between the producers and the consumers ofbotanical information. This paper is derived from a talk titled ''System- which have been added many new sources of in- atics. Informatics, and Floristics" that was present- formation and schemes for interpretation. ed at the Jepson 50th Anniversary Celebration and Two matters were particularly important in the Scientific Symposium in June, 2000, at the Univer- first half of the 20th century for the development sity of California, Berkeley. The purpose of this of monography. The first was the intentional incor- paper is to review how botanical information flows poration of evolutionary questions. Beginning from creators to consumers, and the central role of about 1930, a monograph was regarded as incom- floristics in that process. plete if it did not offer some understanding ofevo- Mankind's age-long study ofplants has produced lutionary relationships among the entities under an amazing legacy, which is evident in any schol- consideration. One of the first and clearest mono- arly library (Barkley 1993). Systematists who add graphs to be published in the USA that had evo- to the accumulating knowledge believe that it all lutionary relationships as a chiefgoal was the tieat- has meaning, and presumably it is useful to people ment of the genus Haplopappus by H. M. Hall outside ofthe bounds ofbotany. The diagrams pre- (1928). Therein the author tried mightily to develop sented here show how the information of system- phylogenies as he understood them. It is not im- atic botany is accumulated and how it reaches the portant that HalTs techniques are now seen as in- consumer in useful form. adequate and that many of the monograph's con- Monographs and revisions (i.e., "monography" clusions are no longer tenable; what is important is are the "soul of systematics,'' as pointed out so that he acted upon the assumption that good mon- neatly by Stuessy (1975). Those studies are taken ography must be centered upon evolutionary rela- to be the fundamental syntheses of systematic tionships. knowledge and are the roadmaps for subsequent The second matter was the advent of new labo- studies. The term "monograph" has been used for ratory techniques, which boosted systematic studies the larger and more sumptuous studies, while "re- into an experimental science with garden and vision" has meant studies presented in less detail. greenhouse studies, cytogenetics, comparative cy- The central goals ofboth are similar, however, and, tology, the analysis of secondary metabolites, etc. as with many species, there is no sharp distinction These studies came to be called "biosystematics" between the two. In this paper, all such studies are and have produced imaginative and detailed inon- called "monographs." Traditional monographs fo- ographic treatments. Biosystematics coupled com- cus on some natural group, such as a genus or a fortably with ecological studies such as pollination, section of a genus, and they include, among other seed dispersal, population biology, and geohistory, things, delimitations and descriptions of the enti- thereby further enriching the content of monogra- ties, keys, hard data on ranges and habitats, and an phy. ordered nomenclature. Modern monographs are A new vista in monography was introduced with also expected to include information on the biology the arrival of cladistic theory and new information of the group and on natural relationships. A how- from molecular studies. Cladistic theory supplied a to outline for a monographic study was presented workable tool for showing evolutionary relation- by A. S. Hitchcock, the noted agrostologist, in a ships, resulting in phylogenetic trees that could be remarkable book called Descriptive Systematic Bot- objectively tested. Molecular studies have proved any (Hitchcock 1925). The book is the product of to be particularly coinpatible to cladistic analyses, an earlier era, and even though the author had few and the two have created a vital subset of system- of the techniques that are available to us now, it is atic studies that focuses on evolutionary relation- still well worth revisiting, for the author laid out a ships, rather than on species delimitation. There is clear recipe for the standard procedures of mon- a rich literature on cladistic theory and derived phy- ography. Arguably, contemporary monography logenies. The application of the phylogenetic ap- rests upon the procedures outlined by Hitchcock, to proach based upon cladistics is comfortably treated 254 MADRONO [Vol. 47 in the recent textbook Plant Systematics: A Phylo- and distributed on a CD-ROM that was prepared genetic Approach by Judd et al. (1999). by Kartesz and Meacham (1999). (2) The detailed Contemporary phylogenetic studies have recog- Flora of Florida project centered at the University nized that the traditional Linnaean concept of spe- of South Florida and prepared by Richard P. Wun- cies is imprecise at best and may be no longerjus- derlin and assisted by Bruce F. Hansen (a manual tifiable (i.e., species are indeed specious). From the was published in 1998). (3) The theoretical works early 1990's to the present there has been a shower ofHugh Wilson at Texas A&M University. The ap- of literature on the creation of a new taxonomic plication of informatics technologies to floristic scheme to reflect phylogenetic relationships, and projects is not easy. The Flora of North America indeed there was a symposium on the topic at the made an effort to incorporate informatics theory, XVI International Botanical Congress in St. Louis, which proved to be administratively difficult. MO in 1999 (cf. Cantino 2000, and Cantino et al. 1999, for an introduction into the literature). It is Explanations of the Figures doubtless true that changes are coming in how we conceive of "species," but the proposed phyloge- Figure 1 simply notes the cascade ofinformation netic classifications are yet to be elaborated, and from monographs and revisions through the floras are yet to be taken into the thinking ofthe consum- and on to the consumers. The consumers are a ers ofbotanical information. For the present, a con- mixed lot; here they are called "primary," "indi- servative approach is prudent, and so the treatment rect," and "ultimate." The primary consumers are of floristic botany rests upon the standard, albeit scientific and academic professionals whose exper- flawed, Linnaean notions of species. tise is not in systematic botany but whose experi- Floristic studies account for all of the plants that ence gives them the ability to judge the accuracy occur in a particular region. Usually this is taken of the information. These are the botanists' col- to mean the vascular plants, although the currently leagues. The indirect consumers are a large group active Flora ofNorth America project also includes who use what is in floristic treatments essentially the bryophytes. Hitchcock (1925) also includes a on faith. It is this group for whom the accumulated discussion on the methods of floristics, but without wisdom in the herbaria and libraries is likely to be the notion of floras as encompassing summaries. of greatest interest and least accessible. Floristic The products of floristic studies are floras or man- projects have an awesome opportunity to connect uals. The two are similar and intergradant, but as this group with botanical information. The ultimate with monographs and revisions, the former are consumer is simply the person who needs infor- more sumptuous, often in several volumes, while mation about a plant, e.g., the person who asks, "Is the latter are stripped-down forconvenient use. Flo- this crabgrass in my lawn? What do I do about it?" ristic botanists derive their information from mono- In many states, the Cooperative Extension Services graphs and revisions, but when no monographic are geared to accumulating information from pri- studies have been done, they must prepare nonce- mary consumers and delivering it to the ultimate treatments with the information at hand. If a flo- consumer. ristic program required that all groups be treated at Figure 2 summarizes the preparation of a mono- equal levels ofsophistication, the flora would never graph, starting with the definition of the problem be written. and the early survey work. Items 3 and 4 are crit- Floristics are best done by botanists with field- ical, for here the monographer's experience (or ifa familiarity in their region who also have good her- graduate student, the experience of the student's barium and library resources. The techniques for mentor) calls for the building of hypotheses and synthesis have been largely intuitive, based upon expressing them as testable models. Items 5-8 are ! the botanist's memory and ability to organize great the chief sources of information useful in monog- amounts of detail. But, just as cladistics and mo- raphy; they are not mutually exclusive, and some lecular data added a huge new approach to mon- techniques have elements of two or more of these ography, electronic information management ("in- items. Clearly, comparative morphology is ofgreat formatics") is changing floristics. It is now think- importance because it is easily accessed in the her- able that a floristic project can account for vast barium, there is a lot of it, and the techniques for amounts of information that effectively lie fallow, using morphological information are of long tradi- and that, through floristic programs and their com- tion. Items 6 and 8 include such matters as polli- puter links, this buried information can be brought nation studies, populational studies, introgression, to the surface. To be certain, floristic projects that the role of climate change, etc. The last item has are based on informatic techniques are in their in- become increasingly significant with the advent of fancy, but the future impact is already evident. readily accessible Geographic Information Systems Three notable computer-based programs come to (CIS). Item 9 is legalistic, mechanical, and utterly mind (but there are others, not mentioned here): (1) essential, for it is how the entities are given their The magnificent summary of information on the correct names. Information from items 5-8 are as- North American Flora as compiled by John Kartesz similated and the results are compared to the hy- in his Biota of North America Program (BONAP) potheses and models generated in items 3 and 4. 2000] BARKLEY: CONTEMPORARY BOTANICAL STUDIES 255 TKe Cascade of Systematics Ndonographs and Revisions V 3 Primary Consumers: e.g.. Ecology, Agronomy, Forestry, 1 Conservation Biology, GeograpHy, Pharmacology, History i 'm- \ 4. Indirect Consumers: Popular Works, e.g.. Hooks on Wiklflowers, Niitural History, Trees, Weeds ! The Ultimate Consumers "The Person on the Street' Fig. 1. Explanation in text. During the assimilation stage, phylogenetic (cladis- ters of the project must be understood to account tic) techniques are applied, which yield justified forthe biological complexities ofthe region and the evolutionary trees showing the current understand- expectations of the intended users of the flora. Pri- ing of natural relationships. It is noteworthy that mary information is taken from monographs and many phylogenetic studies that are based on mo- revisions as much as possible, but when no mon- lecular data focus on higher groups, such as genera ographic works are at hand, it is necessary to create or families, and that species-centered phylogenetic treatments as best as possible; this step essentially studies often rest upon large components of mor- incorporates items 3, 4, and 9 of Figure 2. Item 4, phological data. Item 12, integration, is the aligning preparation of the treatments, is demanding and of the information into the customary format for most easily accomplished by botanists with at least monographic studies. Keys, descriptions, specimen some monographic experience. Computer-assisted citations, sources of data, and conclusions drawn techniques are potentially very useful in item 4 are presented in traditional ways, making the mono- (e.g., DELTA) but to date these techniques have graph (item 13) a readily understandable and use- long and steep learning curves. Many floristic bot- able document, whether published as hard copy or anists are not ready to embrace the computer as a on a website. tool to prepare keys, descriptions, and other textual Figure 3 notes the assimilation of information matters. However, computer-based programs for from monographs and other sources into a floristic generating maps are clearly with us. Text matters treatment. Items 1 and 2 are obvious; the parame- generated in item 4 can be entered into a website MADRONO 256 [Vol. 47 Preparation of a MLonograpK or Revision 1. Definition of Problem Hoi-l->i»riiimi, Library, Field Survey ^- Eyeball Materials to Build Hypotheses | 4. TTentiitive Tii.-sLononiv and Fii< a1 Models 5 Comparative Morphology and . Cieography Herbarium) ( Breeding and Behavioral Studies, (5. "Biosystematics" (Field and Ciarden) : -^ -. : "7_ CayntdolMo^goilcecatli,laBrioScthuedmiiecsa(lL,aAbonraattoomriyc)al ,S l^iogeot;rapliV and Cieohistorv (Librarv anil Field) 9. Nomenclati lO. Phylogenetic Analyses I 11. Assimilati* 12. Integration 13. Systematic M^onograph or Revision % with Phylogenetic Interpret;»tion Fig. 2. Explanation in text. to facilitate the following steps and the production not in the minds of the botanists who did the var- of the final flora. Item 6 is also particularly de- ious studies that led to monographs or floras. A manding, for here is where the tentative product is flora has a wealth of information relevant to distri- critically tested and edited for accuracy. Taxonomic butions, variations, phenologies, etc., that may be reviews treat the botanical matters; regional re- coupled with soil types, geohistorical matters, ar- views account for distributions and regional varia- chaeology, medicine, and other areas not yet con- tion. Item 7, amplifications, is the addition of in- ceived. The point is that the hard data of the core formation needed by the intended users ofthe flora, ofsystematic botany are translated foruse by others i.e., the consumers. Item 8 is where the manuscript through floras. is treated for editorial consistency, where the gen- eral keys are created and tested, and where the in- Conclusions troductory essays are prepared and incorporated. The product may be published as hard copy (item The abiding points are simple: There is a huge 10) or posted on a website (item 1 1). A flora that body ofliterature in libraries and specimens in her- is conveniently available on a website is easy for a baria that are the products of botanical enterprise. primary consumer to consult when addressing Monographic studies are done to determine what broad questions (item 12), e.g., questions that were entities exist, how to distinguish among them, how 2000] BARKLEY: CONTEMPORARY BOTANICAL STUDIES 257 Preparation of a Flora or MLarLual 1. Definition of the Project and its Application 2. Field, Herbarium and Library Surveys T 3 Monographs and Revisions . 4_ Primary Treatments; 5 Website Keys, Descriptions, Ecological and . Distributional Data, Nomenclature 6. Taxonomic and Regional Revii Y. Amplifications, e.g.. Horticultural Uses, Legal Status, Toxicity, Weediness, Colloquial Names 8. Integration and General Key ; 11. Flora on Website; Interactive Database IZ. "Broad Questions" Fig. 3. Explanation in text. they are related to each other, and how they behave. Literature Cited Monographic studies have spawned very interesting Barkley, T M. 1993. Synthesis: a historical perspective. derivatives that relate to sophisticated understand- Annals of the Missouri Botanical Garden 80:292- ings ofevolution, but that still fall within the shad- 296. ow of monographic studies. Floristic studies filter Cantino, p. D. 2000. Phylogenetic nomenclature: address- and assimilate the accumulated wisdom of the ing some concerns. Taxon 49:85-93. plants of a region and couple it to those who use , R. G. Olmstead, and S. J. Wagstaff. 1997. A the information. It is not for nothing that we recall comparison of phylogenetic nomenclature with cur- a botanical beatitude attributed to the late Lloyd H. rent systems: a botanical case study. Systematic Bi- Shinners: "Blessed be those who write floras, for ology 46:313-331. they shall discharge the botanists' responsibilities Hall, H. M. 1928. The genus Haplopappus: a phyloge- to the public." netic study in the Compositae. Publ. Carnegie Insti- Note: A review ofthe growth oftaxonomic con- tution (Washington) 389:i-viii, 1-391. cepts over the past half-century was recently pub- Hitcbhoctoacnky,.AJ.ohSn. W19i2l5e.yM&etShoondss,oNfedwescYroirpkt.ivesystematic lished by P. F. Stevens in a series of "Jubilee Pa- JuDD, W. S., C. S. Campbell, E. A. Kellogg, and P. F. pers" in the journal Taxon (Stevens 2000). It ap- Stevens. 1999. Plant systematics: a phylogenetic ap- peared too late to impact the presentation of this proach. Sinauer Associates, Sunderland, MA. paper at the Jepson Symposium in June, 2000. Kartesz, J. T. and C. a. Meacham. 1999. Synthesis of MADRONO 258 [Vol. 47 the North American Flora. CD-ROM. NorthCarolina Stuessy, T. F. 1975. Theimportanceofrevisionarystudies I Botanical Garden, Chapel Hill, NC. in plant systematics. Sida 6:104-113. Stevens, P. F. 2000. Botanical systematics 1950-2000; WuNDERLiN, R. p. 1998. 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