ALLERTONIA a Series of Occasional Papers
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Vol. 1 No.5 ALLERTONIA A Series of Occasional Papers Wood Anatomy and Relationships of Bataceae, Gyrostemonaceae, and Stylobasiaceae by Sherwin Cariquist Lawai, Kauai, Hawaii February, 1978 F ALLERTONIA is a series of occasional papers intended to serve for publication of longer studies presenting results of original botanical or horticultural research undertaken by members of the staff of the Pacific Tropical Botanical Garden or in collaboration with the Garden and its programs. The title commemorates the late Mr. Robert Allerton (1873—1964). His gener osity and guidance, as one of its first Trustees, were instrumental in the establish ment of the Pacific Tropical Botanical Garden. EDITORIAL COMMITTEE A. C. Smith, Editor H. Kamemoto J. L. Brewbaker C. H. Lamoureux S. Carlquist Y. Sagawa L. Constance W. L. Theobald, ex officio Numbers of ALLERTONIA are priced individually. Standing orders may be placed by writing:. Publications Secretary Pacific Tropical Botanical Garden P.O. Box 340 Lawai, Kauai, Hawaii 96765 Volume I, to date, consists of: No. 1. Rare and Endangered Species of Hawaiian Vascular Plants. By F.R. Fos berg and Derral Herbst Price: $6.00 No. 2. The Pacific Species of Pittosporum Banks ex Gaertn. (Pittosporaceae). By Judith E. Haas Price: $8.00 No. 3. The Family Thelypteridaceae in the Pacific and Australasia. By R. E. Holt turn Price: $5.50 No. 4. Revision of Perymenium (Asteraceae-Heliantheae) in Mexico and Central America. By John J. Fay Price: $5.50 No. 5. Wood Anatomy and Relationships of Bataceae, Gyrostemonaceae, and Stylo basiaceae. By Sherwin Carlquist Price: $3.50 © 1978, by Pacific Tropical Botanical Garden WOOD ANATOMY AND RELATIONSHIPS OF BATACEAE, GYROSTEMONACEAE, AND STYLOBASIACEAE’ SHERWIN CARLQuIsT2 INTRODUCTION One of the problems encountered in application of evidence from anatomy or other disciplines to phylogenetic decisions is the tendency for a worker to support concepts already in existence, rather than to cast widely for relationships that may, at first glance, seem unlikely. A second tendency is for an author to view a genus or family as more isolated on the basis of new evidence than it had hitherto been re garded. These perils are, to a large degree, a function of limited knowledge avail able or, in some instances the lack of acquaintanceship of the author with knowledge already in existence. The present study does not pretend to escape from these diffi culties. To the extent they are overcome, credit is due to the reassessment of centro spermous families and the exclusion of Bataceae and Gyrostemonaceae from that grouping. In particular, I have relied upon my esteemed colleague, Dr. Robert F. Thorne, whose (1976) system comes close to the conclusions tentatively reached here. Thanks to my field work in Western Australia in 1974 and to the availability of other materials, 1 do have a wood collection nearly definitive for the three fam ilies concerned. Wood anatomy is only one line of evidence: in some instances, it merely reveals parallelisms among families; in a few cases, decisive features are revealed. In view of this, data from as many fields of inquiry as possible are cited here. 1 would like to pay credit to the tendency of Thorne (1968, 1976) to utilize phytogeographical plausibility as one of his criteria in grouping families into orders. Ultimately, groups with a common ancestor must have radiated from a particular region. HISTORICAL REVIEW Baiaceae.—Clearly Batis has troubled phylogenists and systematists much more greatly than any but a handful of angiospermous genera. As McLaughlin (1959) notes, Batis has been claimed to have affinities with such remarkably disparate families as Buxaceae, Empetraceae, Fagaceae, Hamamelidaceae, Juglandaceae, J ulianiaceae, Plantaginaceae, Podostemaceae, Polygonaceae, Salicaceae, Thelygon aceae, and Vaticaceae, as well as with the families of the classical Englerian “Cen trospermae” (Chenopodiales of some authors). As a matter of convenience rather than for any especially compelling reason, most authors have settled on what Eichler (1876) termed Centrospermae as a repository for Batis. Eckardt (1976) has reviewed ‘This paper is based on research aided by grants from the National Science Foundation, GB 38901 and B MS 73-07055A I. I wish to thank Dr. Larry DeBuhr, for his aid in the accumulation of data, and my colleague Dr. Robert F. Thorne, for his many helpful comments. 2Claremont Graduate School, Pomona College, and Rancho Santa Ana Botanic Garden, Claremont, California9l7ll. 298 ALLERTONIA 1:5 the varying concepts of Centrospermae in the century since Eichler’s publication. The reader is, therefore, referred to Eckardt’s review for a summary of the phylo genetic peregrinations of Batis. The congested compound inflorescences of Bails, its anemophilous habit, and its preference for saline habitats doubtless invited com parison to, say, Sa/icornia and other Chenopodiaceae, Amaranthaceae, Halophyta ceae, and Aizoaceae. Exclusion of Batis from the centrospermous families can be traced to Bartling (1830), Rallier (1912), Engler (1964), and Thorne (1976). The phylogenists currently most widely cited, Cronquist (1968) and Takhtajan(1973), have retained the centro spermous (chenopodialean) positioning. McLaughlin’s (1959) study of wood anat omy concluded in favor of a centrospermoid placement for Bails. One must concede that all phylogenetic studies are, to a certain extent, influenced either by the pres sure of tradition or by the selection or availability of materials and taxa that a worker has chosen to study. Depending on the nature of these influences, one can discount phylogenetic conclusions as casual or regard them as compelling. In the case of Batis, the use of relatively few characters and the narrowness of potential relation ships entertained by most workers should encourage one to be more open-minded about the potential affinities. Gyrostemonaceae have traditionally been a part of Centrospermae, often treated within the family Phytolaccaceae, as in Walter’s (1909) account, if one is willing to overlook that tradition for the moment, one can see that a number of features ally Bataceae with Gyrostemonaceae. For example, Takhtajan (1959, 1966, 1973) paired Bataceae and Gyrostemonaceae. Confirming this treatment is the lack of betalains in both Bataceae and Gyrostemonaceae, as well as the presence of S-type plastids (rather than the P-type of Centrospermae) in both families (see the review of Behnke, 1976). Likewise, Prijanto (1970a, l970b) found that ultrastructure of pollen exine and other features supported relationship between Bataceae and Gyro stemonaceae. We have been limited in our viewpoint of Bataceae by the fact that prior to 1956 only Bails maritima was used to examine relationships of Bataceae. With the de scription of B. argillicola by van Royen (1956; see also van Heel, 1958), our horizons are widened. One consideration in this regard is phytogeography. The distribution of B. maritima includes tropical and subtropical coasts of both Atlantic and Pacific shores of the New World (McLaughlin, 1959). Presence of B. maritima in the Ha waiian Islands is very likely a result of human introduction, judging from the com ments of Hillebrand (1888). Bails maritima is undoubtedly distributed by means of oceanic drift. Thus, a center for its origin would be difficult to discern on the basis of the single species. However, B. argillicola, with a constellation of characters more primitive for the family than those of B. maritima, has a much more restricted distribution. Originally B. argillicola was reported from southern New Guinea, where it may also be distributed locally by oceanic drift, in mud flats adjacent to mangroves (van Royen, 1956). Batis argillicola is also now known from stations in Australia. It occurs on the Cape York Peninsula, in such places as Kurumba, Burke Dist. (W. G. Trapnell 193, Q) and lda Point, 4 km. SE of Cape York (L. S. Smith 12522, Q). Because its distribution and features are more primitive than those of B. inaritima, one might expect the closest relatives of Bataceae in the Australian area. Gyrostemonaceae is an exclusively Australian family (Walter, 1909). 1978 CARLQUIST: WOOD ANATOMY 299 Gvrostemonaceae. —G yrostemonaceae remained embedded within the Phyto laccaceae and therefore within Centrospermae (Chenopodiales, Caryophyllales) until very recently. The whorl of numerous carpels in female flowers of most genera was undoubtedly the obvious characteristic on which this treatment was based. The history of this disposition is well documented by Eckardt (1976). The first modern author to recognize Gyrostemonaceae as a family was probably Heimerl (1934). Modern phylogenists, such as Takhtajan (1959, 1966, 1973), Hutchinson (1959), and Cronquist (1968) recognized Gyrostemonaceae as a separate family but never theless grouped it with Phytolaccaceae. Now that presence of betalains nd P-type plastids are deemed prime criteria for delimitation of a centrospermous (Cheno podiales) alliance (Behnke, 1975, 1976; Goldblatt et al., 1976), Gyrostemonaceae does not appear to belong near Phytolaccaceae or other families of Chenopodiales. Moreover, pollen of Gyrostemonaceae is not of a type found in centrospermous families (Skvarla and Nowicke, 1976; Prijanto, 1970a). The removal of Gyrostem onaceae from Centrospermae (Chenopodiales) forces phylogenists to find a suit able recipient group for the family. Thus far, two hypotheses may be said to be current. Dahigren (1975), on the basis of presence of glucosinolates in