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 Codono carpus cotinfolius, claims Gyrostemonaceae to be capparalean. This view is seem ingly endorsed by Goldblatt et al. (1976), although their description of pollen does not suggest a capparalean affinity. The second view may be attributed to Thorne (1976), who finds a sapindalean placement probable for Gyrostemonaceae. Although he does not detail the reasons for this treatment, Thorne’s placement is based on such features as floral morphology, ovule and seed morphology, and presence of stipules. The issue at hand, then, is whether the single chemical character cited as indica tive of relationship of Gyrostemonaceae to Capparales is reinforced by other lines of evidence, or whether Thorne’s summation based on morphology is supported by additional information. There is, of course, also a possibility that affinities other than capparalean and sapindalean will be advanced by some authors, or that, in seeking comparisons, one ought to cast a much wider net. Siylobasiaceae.—The history of the placement of Stylobasium, sole genus of its family, has been detailed neatly by Prance (1965). Prance shows that, although Stylobasium was referred to the tribe Chrysobalanoideae of Rosaceae (Chrysobal anaceae of some authors), a wide spectrum of evidence for regarding the genus as sapindalean exists. Prance does recognize Stylobasium as constituting a mono- generic family, whereas Thorne (1976) regarded Stylobasium as forming a subfamily of Sapindaceae. The difference between those two viewpoints is obviously minor.

MATERIALS AND METHODS The wood sample of Batis maritima was collected by me at Hanauma Bay, Oahu, on November 24, 1972. Batis maritima is common in a number of localities in the Hawaiian Islands, although it was originally introduced in Honolulu harbor (Hille brand, 1888). The stem studied was 35 mm. in diameter, which is rather large for this species. Material of B. argillicola was not studied because van Royen’s (1956) description of wood anatomy reveals no appreciable differences between B. argilli cola and B. maritima. Woods of Gyrostemonaceae were collected in Western Australia in 1967 and 300 ALLERTONIA 1:5

TABLE I. FEATUREs OF WOOD ANATOMY OF BATAcEAF,

Species I 2 3 4 5 6 Bataceae Bat is mariti,na L. 1972 S 40 126 109 388 s.n. Gyrostemonaceae Codonocarpus coun/1ius 5150 S 107 14 169 874 (Desf.) F. Muell. B 74 60 156 766 531/ S 99 IS 149 899 B 50 95 220 736 Gtrosternon racemigerus 5210 5 50 75 220 606 H. Walt. C. ramu/osus Desf. 2989 S 52 115 321 773 3/53 S 53 95 156 717 515! S 103 19 158 659 5183 S 87 39 170 743 5467 S 64 36 148 642 C. sheathii W.V. Fitzg. 6010 S 102 32 172 862 C. subnudus (Nees) Diels 55/4 5 51 90 147 646 B 49 106 167 588 C. tepperi F. Muell. S 54 94 222 818 Tersonia brei’ipes Moq. 5385 S 75 30 175 689 R 107 31 212 707 Stylobasiaceae St,!obasium australe 5227 S 40 75 289 749 (Hook.) Prance 5434 5 41 119 263 620 S. spathulatum Desf. 5453 S 55 74 266 631

Explanation of columns 1-14. I. S. Carlquist collection number. 2. Portion: S = stem; B = branch; R r root. 3. Vessel diameter, mean, pm. 4. Vessels per sq. mm. of transection, mean. 5. Vessel element length, mean. pm. 6. lmperforate element length, mean, pm. 7. Imperforate element wall thickness, mean, pm. 8. Ray histology: u upright; s square; p procumbent; the predominant type or types in-

1974. Although all Western Australian species could be said to occur in sandplain, habitats differ somewhat. Gyrostemon ramulosus is widespread in the southwestern corner of the state and extends well into interior desert regions. Gyrostemon race migera is a local species from an area west of Coorow. Gyrostemon sheathii is endemic to sandhills along the coast near Esperance, and shows a very marked de gree of succulence probably correlated with this maritime habitat. Gyrostemon tepperi is known only from scattered localities in the desert interior of Australia. Codonocarpus cotinfo1ius is widespread from moderately dry areas, as at Merredin (Carlquist 5311) to desert areas far into the interior (Carlquist 5150). Tersonia brevi pes tends to grow in sandy areas paralleling the southwestern coast to the north and south of Perth, in a belt between the immediate coast and inland dry areas. Citation of localities for the various species of Gyrostemonaceae may be found in Walter (1909). The genera Didymotheca and Cypselocarpus were deliberately omitted from the present study because they accumulate very little secondary xylem and thus would not be truly comparable with the woodier species. Tersonia brevipes can be called a prostrate shrub; the root as well as older stems becomes woody. The two species of Sivlobasiurn also tend to occur in sandy areas of Australia. My material of S. sparhulatum came from a coastal sand area near Northampton. WA., whereas material studied by Prance (1965) was collected in an interior area, 1978 CARLQUIST: WOOD ANATOMY 301

GyRosrEM0NAcEAE, AND STYL0BAsIAcEAE

7 8 9 10 II 12 13 14

2.4 USP 1382 — 1.6 3.56 0.32 35

2.6 usP 392 107 2.1 5.17 7.41 1252 2.5 usP 392 107 2.1 4.91 1.24 193 3.0 usP 442 69 2.2 6.03 5.55 827 2.5 usP 549 87 4.! 3.35 0.53 1l7 3.5 usP 736 84 2.6 2.75 0.67 147

3.3 usP 1290 195 3.3 3.50 0.45 99 3.8 usP 795 109 2.7 4.60 0.55 86 3.0 usP 693 96 2.6 4.17 5.4! 855 3.1 usP 52! 90 2.4 4.37 2.22 377 2.9 usP 508 123 2.6 4.34 1.78 263 4.5 usP 629 120 2.3 5.0! 3.19 547 3. I usP 998 97 6.9 4.39 0.56 82 3.8 usP 843 127 5.6 3.52 0.45 75 2.8 usP 408 92 4.8 3.68 0.57 126 3.3 usP 1041 24 1.9 3.94 2.54 445 3.5 usP 1538 129 1.8 3.35 3.45 730

6.1 USp 282 155 2.8 2.59 0.53 153 5.1 USp 318 235 5.! 2.36 0.34 89 6.3 USp 247 267 3.9 2.37 0.74 146 dicated by upper case. 9. Height multiseriate rays, mean. pm. 10. Height uniseriate rays, mean, pm. II. Vessels per group, mean. 12. Ratio of imperforate element length to vessel element length. 13. “Vulner ability” ratio: vessels per sq. mm. transection divided by vessel diameter. 14. “Mesomorphy” ratio: “Vu! nerability” ratio multiplied by vessel element length. near Alice Springs, Northern Territory. My material of S. lineare was collected in southwestern Australia, near Morawa (5227) and in the Gardner Reserve near Merredin (5434). Bataceae, Gyrostemonaceae, and Stylobasiaceae could all be said to be weedy in nature and found on disturbed or rapidly changing habitats. The littoral sands and mud flats where Batis occurs obviously qualify for this description. Gyrostem onaceae often occur on recently burned areas and are not characteristic of stable scrub. The interior sands on which Gyrostemonaceae occur can be regarded as un stable, since the sands are sparsely vegetated and winds tend to shift them. Stylo basium often occurs in weedy areas at roadsides. All species of Bataceae, Gyro stemonaceae, and Stylobasiaceae seem to form shrubs or trees of relatively brief duration, and none has the aspect of a truly woody plant. This may be correlated with the disturbed habitats in which they occur. For all species concerned in this study, wood samples of maximal diameter within a population were collected except for the branch wood samples noted in Table 1. Voucher specimens are located in the herbarium of the Rancho Santa Ana Botanic Garden; duplicates have been distributed to other herbaria. Wood samples were prepared by drying. Wood samples were sectioned and macerated according to the usual techniques. 302 ALLERTONIA 1:5

For vessel element and libriform fiber dimensions, means were derived from fifty measurements. For other quantitative features, means were derived from ten mea surements.

WOOD ANATOMY Bataceae.—Although Soldereder (1908), Metcalfe and Chalk (1950), and McLaughlin (1959) have described the wood of Batis maritima, and van Royen (1956) has contributed information on that of B. argillicola, a summary of wood fea tures seems in order. Quantitative features are given for B. maritima in TABLE 1. The data and photomicrographs of van Royen (1956) for B. argillicola show no marked differences, with exceptions as noted below. Growth rings in Batis maritima are absent (FIGuRE 1). Vessels solitary or grouped (figures given by McLaughlin), most commonly in radial chains, although tangential bands (as shown by McLaughlin) occasionally present. Perforation plates simple. Intervascular pitting alternate on lateral walls of vessels, alternate with occasional opposite pits on vessel-ray contacts. Thin-walled tyloses reported in some vessels by McLaughlin (1959). All imperforate tracheary elements are fiber tracheids, with narrow borders present on pits (FIGuRE 5). Axial parenchyma vasi centric; banded apotracheal parenchyma also present. The apotracheal paren chyma bands may be the result of fiber dimorphism, as discussed below. Axial parenchyma cells not subdivided or in strands of two cells (the latter true of vasi centric parenchyma). Rays predominantly multiseriate (FIGURE 2), uniseriate rays extremely few. Height of rays given in TABLE I; dimensions in terms of cells given by McLaughlin (1959). Perforated ray cells present (FIGuRE 3). Multiseriate rays heterocellular, with procumbent, square, and erect ray cells about equally abundant in my material. All ray cells are procumbent in B. argillicola according to van Royen (1956), but his figures suggest that a few upright ray cells may be present also. Pits on ray cells simple or bordered (FIGURE 6). Amorphous crystalline material in ray cells reported by McLaughlin, but not present in my material or in B. argillicola according to van Royen. Dark-staining deposits in the form of droplets or small masses present in ray cells and in some vessels (FIGURES 1—3, 6). Wood clearly storied only with respect to axial parenchyma bands (which may be alternatively interpreted as short fibers), as shown in FIGuRE 2. One may assume that the cam bium is storied, and that elongation of fiber-tracheids during their maturation ob scures the basically storied pattern (as noted in TABLE 1, fiber-tracheids are 3.56 times as long as vessel elements in B. maritima). Gyrostemonaceae.—Some details on wood anatomy of Codonocarpus and Gyrostemon (species not cited) are offered by Metcalfe and Chalk (1950). 1 have presented data on seven species belonging to three genera. Cypselocarpus halora goides F. Muell. (FIGURES 57, 58) was collected, but this species has too little wood to be useful in analysis of secondary growth. Tersonia subvolubilis Benth. is prob ably synonymous with C. haloragoides, as claimed by Prijanto (1970a). Didymo theca was likewise omitted from this study because it has very little secondary xylem. Of the remaining Gyrostemonaceae listed by Walter (1909), only Codonocar pus attenuatus (Hook. f.) Walter, C. pyramidalis F. Muell., Gyrostemon australa sicus (Moq.) Heimerl, and G. brownii S. Moore are not represented here. Walter’s Gyrostemon spinoso-stipulatus seems clearly a synonym of G. sheathii, a name 1978 CARLQUIST: WOOD ANATOMY 303

Walter does not seem to have used. Walter evidently neglected the distinctive spe cies G. tepperi (FIGuREs 55, 56). Blackall and Grieve (1954) evidently neglected G. racemigerus, a species which seems worthy of recognition. Wood anatomy for Gyro stemonaceae as a whole can be summarized as follows. Growth rings absent or very few and inconspicuous (the banded parenchyma is not produced annually and therefore does not constitute a form of growth ring ac tivity). Vessels occasionally solitary, but more commonly united in tangential group ings (FIGuREs 8, 10, 12, 18); groupings large in G. subnudus(FIGuRE 14). Intervascu lar and vessel-ray pitting basically alternate (FIGuRE 17, left) but some scalariform or scalariform-like pitting present. The latter pitting probably represents lateral elongation of pits in an alternate pattern rather than in a true scalariform pattern. Perforation plates simple. Tyloses not observed. Imperforate tracheary elements all tracheids, with pit apertures not longer than the diameter of the pit cavity. Tra cheid walls moderately thin (FIGuREs 8, 14, 18) to relatively thick (FIGUREs 10, 12). Axial parenchyma cells in tangential bands, chiefly paratracheal. Diffuse parenchy ma also present between bands in all species (FIGuRES 8, 10, 12, 14, 18). By virtue of abundance of the diffuse parenchyma cells, diffuse-in-aggregates are necessarily present. Axial parenchyma cells not subdivided or more commonly in strands of two to four cells, two being most common (FIGuRES 13, 15). Diffuse axial parenchy ma strands of roughly the same length as the tracheids they accompany, whereas paratracheal parenchyma strands approximate the length of the vessel elements they accompany. Multiseriate rays more abundant than uniseriate rays, although uniseriates are present in all species. Multiseriate rays wide in my material of all species, although Metcalfe and Chalk (1950) state that rays do not exceed 3 or 4 cells in width in Codonocarpus. Although upright and square cells are present as sheath ing cells (generally a single or partial layer at edges of multiseriate rays: FIGURES 15, 19), the vast majority of ray cells are markedly procumbent (FIGuRES 7, 16, 21) in all genera. Banded axial parenchyma cells in bands clearly to vaguely storied (FIGUREs 11, 13, 15, 19, 20). Shorter imperforate elements (which may be termed axial parenchyma of the banded sort) storied (FIGuRE 20). The cambium is therefore basically storied, with elongation of the tracheids obliterating the storied pattern. This can be readily understood, as in Batis, if one views the figures in TAHLE 1, column 12, for ratio between imperforate tracheary element (tracheids in the case of Gyrostemonaceae) length to vessel element length. These relatively high ratios, compared with primitive or even specialized dicotyledonous woods (Cariquist, 1975: 141) inevitably mean that a storied pattern cannot prevail with such great (and, in a cell population, varied) elongation of the imperforate tracheary elements. Rays do not conform to the storied pattern, with the exception of a very few short uniseriate rays. Dark-staining amorphous deposits not noted in any wood cells. Crystals apparently absent. One can point to distinctive characteristics in the species studied. Codonocarpus cotinfolius is notable by virtue of the sparsity and great width of vessels (FIGURE 8) and by the small size of ray cells (FIGURE 9). Gyrostemon ramulosus has thick-walled tracheids in prominent bands, a mode of structure which may be related to the arboreal tendency of this species compared with the smaller stature of other species, such as G. sheathii (FIGURES 12, 13) and G. subnudus (FIGURES 14, 15). Ray cell size and number of vessels per group are notably higher in G. subnudus than in the other

aliLl I In—N 306 ALLERTONIA 1:5

21. Axial parenchyma in paratracheal forms such as aliform or aliform-confluent: some genera, such as Toulicia (FIGURE 42), Hemigyrosa, etc. These cells are shorter than diffuse parenchyma cells (FIGUREs 40, 45). 22. Axial parenchyma bands tending to be apotracheal: Placodiscus, Pometia, etc. Some bands perhaps related to growth ring phenomena but others clearly not. 23. Apotracheal bançls of short parenchyma-like fibers (perhaps the result of fiber dimorphism) present, fibers not subdivided into strands or in strands of two cells or so: Paranephelium, Sapindus (Metcalfe and Chalk, 1950); storied in Aphania (Solereder, 1908). 24. Vascular rays uniseriate exclusively: some genera, such as Toulicia (FIGuRE 44). 25. Vascular rays uniseriate and biseriate: some genera. 26. Vascular rays uniseriate and multiseriate: some genera, such as Dodonaea (FIGURE 33) and Thouinidium (FIGURE 38). 27. Vascular rays heterocellular, but with procumbent cells predominant: some genera (listed by Metcalfe and Chalk, 1950). 28. Vascular rays homocellular or nearly so, nearly all cells procumbent: most genera, such as Dodonaea (FIGuRE 34), Thouinidium (FIGURE 40), and Toulicia (FIGURE 45). 29. Ray cell walls thick, pits often bordered: Dodonaea (FIGURE 35), but probably other genera; this character has been studied insufficiently by wood anatomists, who apparently do not examine pits in rays in radial sections frequently. 30. Ray cell walls thin and with simple pits: presumably many genera. 31. Gummy amorphous dark-staining deposits present in ray cells, vessels, and other cells: Dodonaea (FIGUREs 32-36). Toulicia, and other genera. 32. Gummy amorphous dark-staining deposits absent: some genera.

Notable in the above is the tendency for rays to be composed of procumbent cells mostly or exclusively. Absence of numerous upright cells is unusual for a large and diverse family such as Sapindaceae. Also significant is the diversity in axial paren chyma distribution. Most genera have more than a single type of axial parenchyma, as the genera for which photomicrographs are presented illustrate. These two fea tures seem of key importance in relating Sapindaceae to other families. Ray his tology was stressed by Heimsch (1942) in claiming relationship between Sapinda ceae and Aceraceae or Sapindaceae and Hippocastanaceae. The features presented for Sapindaceae above in serial form are all represented in Bataceae, Gyrostemonaceae, and Stylobasiaceae with some exceptions, which will be noted. Bataceae have the following features: 1, 2, 3, 5, 7,8,9, 11, 13, 19, 23, 27, 29, 30, 31. The only feature of Bataceae unlike Sapindaceae is the relative abundance of multiseriate rays, with very few uniseriate rays (FIGURE 2). The short imperforate elements which occur in bands and may be termed axial parenchyma occur in a storied pattern; storying of libriform fibers is reported in Sapindaceae only in Aphania thus far. Presumably these fibers are rather like the parenchyma in Sapin dus which Metcalfe and Chalk (1950) interpret as fiber dimorphism, apparently: “many of the species with little parenchyma have vasicentric sheaths of wide septate 1978 CARLQU1ST: WOOD ANATOMY 307

fibers that resemble parenchyma strands.” Gyrostemonaceae have the following features cited for Sapindaceae: 1, 2, 3, 4, 6, 7, 9, 10, 13, 15, 18, 22, 23, 26, 27, 28, 29, 30, 32. The tendency in some species for ves sels to form large clusters is somewhat different from aggregations of vessels in Sapindaceae. The presence of borders on pits of imperforate tracheary elements, which are therefore tracheids, is distinct from the condition in Sapindaceae, which have at best vestigial borders on pits of imperforate tracheary elements. The banded axial parenchyma of Gyrostemonaceae, as the accompanying photomicrographs show, differs somewhat from that of Bataceae and Stylobasiaceae in that the cells are most frequently in strands of two cells, rather than non-septate. The relatively wide multiseriate rays are unlike those of the majority of Sapindaceae, but a few Sapindaceae have been reported to have wider multiseriate rays (Metcalfe and Chalk, 1950). Stylobasiaceae have the following features cited for Sapindaceae: 1, 2, 3, 4, 5, 8, 10, 13, 14, 15, 19, 23 (storied), 25, 27, 29, 31. Erect ray cells are somewhat more abundant than in the majority of Sapindaceae. This could be ascribed to the more herbaceous habit of Stylobasium, if my (1962) hypothesis is applicable here. Gelat inous libriform fibers, characteristic of Stylobasium, are not characteristic of Bata ceae or Gyrostemonaceae, and gelatinous libriform fibers have not yet been reported for Sapindaceae, although they may occur. Bataceae, Gyrostemonaceae, and Stylobasiaceae are alike in having axial paren chyma cells like short fibers, and in a storied pattern. Since this condition has been reported for Sapindaceae, as noted above, this fiber dimorphism may be an interest ing clue to relationship. Predominance of uniseriate rays, with some biseriate rays, presence of radial chains of vessels, presence of libriform fibers rather than tracheids or fiber-tracheids, presence of tyloses, and accumulation of amorphous dark-staining deposits are characters in which Stylobasiaceae agree with Sapindaceae. The oc currence of tracheids in Gyrostemonaceae should probably be interpreted as indica tive that Gyrostemonaceae diverged from ancestors which possessed true tracheids. Thus, Gyrostemonaceae could not be derived from modern Sapindaceae or any other family with libriform fibers, but they could have been derived from a pre sapindaceous (or other) stock in which tracheids were present. The presence of wide multiseriate rays is a feature in which Bataceae resemble Gyrostemonaceae. The herbaceous tendencies of Gyrostemonaceae and Bataceae might be thought to be related to wider rays, which would represent a departure from the rather narrow multiseriate rays of most Sapindaceae. However, in both Bataceae and Gyrostemo naceae, the rays are composed mostly of procumbent cells, which is certainly not true of herbs in general (Carlquist, 1962), but may well indicate relationship be tween Sapindaceae and both Bataceae and Gyrostemonaceae. The presence of dif fuse parenchyma in Gyrostemonaceae is paralleled in Sapindaceae, but Bataceae and Stylobasiaceae cannot be said to have true diffuse parenchyma. If diffuse paren chyma is primitive, as often alleged, its presence in Gyrostemonaceae, along with presence of tracheids rather than libriform fibers or fiber-tracheids, would indicate a derivation from a level at least as primitive as that represented in Sapindaceae. Evidence from wood anatomy thus seems to show resemblance between Stylobasi urn and Sapindaceae, although the familial distinction of Stylobasiaceae seems warranted. Wood of Bataceae and Gyrostemonaceae shows some key resemblances

if to xeromorphic be to expected be ought halophytes of anatomy Wood halophytes.

of about wood more much know should clearly 1977). We (Cariquist, shrubs desert

or epiparasites either of range the in low, is M value The xylem. of columns water

the in tension high with but water, of availability constant conditions: similar under

exists Bails that indicate may 1977). This (Carlquist, epiparasite an of that like

more rather is shrubs. It desert values for V the as low as not is 13) column I, (TABLE

maritima Bails for value V 1969). The (Carlquist, xeromorphy do indicate not shrub,

strand a taccada, Scaevola of tissue conductive on figures Comparable however.

anatomy, of wood terms in be xeromorphic not may halophytes All xeromorphy.

of be to indicative these tend All of maritima). Batis im in 4 averages thickness

(wall thick-walled relatively mm. and sq. per numerous relatively also are Vessels

and narrow. short relatively Batis are of elements vessel The halophyte. true a not is

but it of Australia, areas and alkaline in coastal grow can Dodonaea that argue can

here. One studied groups other the of any unlike halophytes, are clearly Bataceae

1977). 1975, (Carlquist,

earlier been developed have for this Criteria regard. in this consideration of worthy

and are anatomy, wood in reflected are Stylobasiaceae and Gyrostemonaceae, ceae,

Bata of habitats distinctive anatomy.—The wood of interpretation Ecological

alone. wood basis of on the parales

Cap with relationship rule out to able be not would but one Sapindaceae, of wood

does than and Stylobasiaceae Gyrostemonaceae, Bataceae, to resemblances fewer

shows of Capparales features wood of The totality Sapindaceae. and Stylobasiaceae,

Gyrostemonaceae, Bataceae, in been reported not have pits vestured whereas 1950),

Chalk, and (Metcalfe and Brassicaceae Capparaceae in occur pits Vestured ceae.

in Bata condition the resembles Brassicaceae in rays uniseriate of rarity The 1975).

(Carlquist, Asteraceae in described was first phenomenon this that fact the from

be discerned as may families, relate necessarily not does dimorphism Fiber 1971).

(Carlquist, Sinapidendron as such Brassicaceae, in found be may dimorphism fiber

of product a represent which cells fibriform storied containing parenchyma cheal

apotra banded However, above. considered families the of to those compared cells

procumbent few relatively have but heterogeneous, are Rays cambia. successive its

of by virtue distinct quite is which (Capparaceae), Forchammeria in except chyma

paren diffuse no with scanty vasicentric, is Resedaceae and Moringaceae, paraceae,

Cap Brassicaceae, in parenchyma 1950). Axial ChaLk, and (Metcalfe Moringaceae

of those for pits and simple Capparaceae of those for borders vestigial but ceae,

Brassica in elements tracheary imperforate for claimed are pits bordered simple;

are plates perforation specialized: highly is rather wood Capparalean 1976). et al.,

Goldblatt 1975; (Dahlgren, been suggested has Gyrostemonaceae and Bataceae

for placement capparalean because here be examined should Capparales of Wood

Dodonaea. of that in but not Diplopeltis of

my material in observed were strands parenchyma axial crystalliferous that except

Dodonaea of that resembles closely Diplopeltis of wood The features. daceous

sapin typical with keeping in is and structure, of modes herbaceous of hint no gives

Diplopeltis of wood However, in Sapindaceae. find can one as shrub a herbaceous

nearly as and also Australian is it because pertinent is Diplopeltis George. S. Alex

by Mr. provided kindly were which of samples wood Diplopeltis, of species three of

wood examined deliberately I regard, In this distinctions. also but Sapindaceae, with

1:5 ALLERTONIA 308 1978 CARLQUIST: WOOD ANATOMY 309

foliar apparatus does not alter the water relations of the plant. Bails is unusual in that one would deem it a succulent on the basis of foliar anatomy, but its quantita tive wood features do not put it in the range of most succulents (Carlquist, 1977). Gyrostemonaceae, on the other hand, exhibit V and M values (TABLE 1, columns 13 and 14) like those of succulents. The family was cited earlier in this regard (Carl quist, 1977). Gyrostemonaceae are among the few plants of a succulent nature in the Australian flora, and foliar morphology as well as thickness of bark suggests this (see FIGuREs 48, 49, 52, 53, 55—58). Codonocarpus leaves are not particulatly succu lent. However, Codonocarpus should be regarded as a short-lived monocarpic plant (see FIGURE 46), growing in periods of water availability and more comparable to an annual than to a tree. There are relatively few groups of succulents in the Australian flora. Gyrostemonaceae are certainly one of the better developed groups of succu lents in the flora, and the wood anatomy does appear to validate Gyrostemonaceae as a succulent group in quantitative terms. For Stylobasiaceae, the V and M values (TABLE 1, columns 13 and 14) are above those of sand-heath and desert shrubs of Western Australia (Carlquist, 1977). This is perhaps surprising, in view of the fact that the two species of Stylobasium grow with shrubs which have lower V and M values. Stylobasium australe, on account of its terete leaves, could perhaps qualify as a succulent, in which case these values would be understandable. Also, Stylobasium is conspicuous to botanists as a shrub invading weedy ditches and other disturbed habitats where more water would be available than in undisturbed open sand hills. Woods of some Sapindaceae, such as Dodonaea (FIGURES 32, 33) have narrow, short vessel-elements suggestive of adaptation to drier habitats. Most Sapindaceae have vessel-elements that are wider, longer, and fewer per sq. mm. than any Bata ceae, Gyrostemonaceae, or Stylobasiaceae. This is not surprising in view of the fact that most Sapindaceae occupy relatively mesic habitats.

POLLEN Pollen grains of Stylobasium are tricolporate, with a markedly triangular outline (amb), as shown by Prance (1965). Although his details on fine structure of exine are vague, Prance does present a convincing case that pollen grains of Stylobasium fall within the range of structural types found in Sapindaceae, a range the breadth of which has been stressed by Erdtman (1952). Pollen grains of Bataceae resemble those of Gyrostemonaceae closely in that the fine structure of the exine is pertectate (lacking in columellae), as the transmission electron photomicrographs of Prijanto (l970a, 1970b) show clearly. Prijanto does note the similarity. His designation of the apertures of pollen grains as compound in Bataceae but simple in Gyrostemonaceae is probably not as deep-seated a distinc tion as one might think. Rather, it seems to me a descriptive way of stressing that there is somewhat more differentiation in colpi of Bataceae, a differentiation that appears minor in Prijanto’s figures. The pertectate nature of exine ultrastructure and the psilate exine surface of pollen grains in Bataceae and Gyrostemonaceae are features very likely related to anemophily. Conceding that Bataceae and Gyrostemonaceae can be paired on the basis of palynological evidence, are they in turn allied to Sapindaceae or other families on the basis of pollen structure? The answer is complicated by two factors: Bataceae 310 ALLERTONIA 1:5 and Gyrostemonaceae are anemophilous, whereas putatively related families are entomophilous and therefore have different pollen morphology and ultrastructure; and pollen grains of both Sapindaceae and Capparales are, in a generalized way, common types in dicotyledons. The presence of a colpus with a transverse furrow (ora), for example, is too widespread in dicotyledons to be of diagnostic value. At tempting to compare pollen ultrastructure of Bataceae and Gyrostemonaceae to that of entomophilous dicotyledons not only shows marked differences, but it forces comparisons based on vague and misleading resemblances at best.

CHEMICAL STUDIES Because Bataceae and Gyrostemonaceae lack betalains and because they have S-type rather than P-type plastids, they have been justifiably excluded from Cheno podiales (see Behnke, 1976, and the other symposium contributions accompanying that paper). On the other hand, presence of glucosinolates in Bataceae and Gyro stemonaceae has been adduced as evidence for inclusion of these two families in Capparales (Ettlinger and Kjaer, 1968; Dahlgren, 1975; Goldblatt et al., 1976). The question of glucosinolates is a problematic one, and needs further examination. The distribution of glucosinolates within dicotyledons may not yet be completely known. However, the occurrence of these distinctive sulfur compounds was long ago sig naled at the anatomical level by the description of so-called myrosin cells, which are the idioblastic sites for deposition of these compounds. Myrosin cells are known from Brassicaceae, Capparaceae, Moringaceae, and Resedaceae, according to Met calfe and Chalk (1950). These four families are indeed closely allied; for example, they form the Capparales of Thorne (1976). However, myrosin cells have been re ported from Bretschneidera (see Tang, 1935; Heimsch, 1942), a report that has neither been denied nor confirmed. Unfortunately, Bretschneidera is native to re mote areas of China and has not been accessible for study by various workers. If, as Heimsch contends, Bretschneidera wood anatomy places it clearly in Sapindales as a monogeneric family and not in Capparales, and if Bretschneidera does prove to have myrosin cells and therefore glucosinolates, have myrosin cells and glucosino lates arisen more than once in dicotyledons? Myrosin cells were undoubtedly used as guides in exploration for glucosinolates, but Bataceae and Gyrostemonaceae do not have myrosin cells. This may suggest Bataceae and Gyrostemonaceae, despite presence of the compounds, are at some remove from Capparales, but also it may suggest that glucosinolates may yet be uncovered in other families lacking myrosin cells.

CHROMOSOME NUMBERS Goldblatt (1976) notes that thus far x 11 can be claimed for Bataceae and x = 14 for Gyrostemonaceae. Because Bataceae and Gyrostemonaceae are held to be well demarcated from each other even if they are related, these two chromosome numbers do not rule out a relationship. Likewise, they do not prove helpful in finding which dicotyledon families might be most closely related to Bataceae or Gyro stemonaceae.

FLOWER AND FRUIT MORPHOLOGY Bataceae.—If a capparalean position for Bataceae and Gyrostemonaceae is to be entertained, one would expect that floral morphology and fruit anatomy would yield 1978 CARLQUIST: WOOD ANATOMY 311

evidence for this proposed alliance. The flowers of Batis maritima have been figured in a number of publications. They are well figured, together with flowers of B. argillicola, by van Royen (1956), and therefore are not reproduced here. Both species have unisexual flowers (a con dition perhaps related to anemophily), although B. maritima is dioecious whereas B. argillicola is monoécious. Van Royen (1956) convincingly interprets the spathella surrounding Bails flowers as a tube composed of two united bracts conforming to the decussate pattern of foliage. Thus the male flower of Bails consists of four tepals with four stamens alternate with them. Tetramery is also suggested in female flow ers. The gynoecium of Bails is interpreted as consisting of two united carpels, each with two locules each containing an ovule at the locule base. The ovules mature into nutlets. The fruit is indehiscent. The nutlets and the parenchymatous fruit are un doubtedly adapted to dispersal by means of flotation in seawater. Therefore, one cannot make comparisons readily between fruit histology of Bails and that of other genera which have quite different dispersal mechanisms. Absence of arils from seeds, for example, would be expected in the case of an indehiscent nutlet such as Bails. Likewise, the number of floral parts may not be significant, although one is tempted to align the tetramery of flowers to the decussate nature of foliage. The morphology of the four-loculate bicarpellate gynoecium appears more nearly con gruent with Sapindales than with Capparales. However, too little is known at pres ent to draw a reliable conclusion. Interpretation may be aided by developmental studies, which might elucidate the nature of the false septum in carpels. Further more, study of ovules, embryo sac development, and embryos is essential before a satisfying assessment of Bataceae can be achieved. Gyrostemonaceae. —Gyrostemonaceae offer features that lend themselves to interpretation more readily. Plants are dioecious. In female flowers, carpel number ranges from one in Gyrostemon tepperi (FIGuRE 56) and Cypselocarpus haloragoides (FIGuRE 58) to two in the genus Didymotheca to five in some species of Gyrostemon (e.g., sheathii, G. FIGuRE 49) to numerous, as in G. ramulosus (FIGuREs 53, 54) and the genera Codonocarpus (FIGuRE 47) and Tersonia (FIGURES 60, 62). The majority of Gyrostemonaceae thereby have more than five carpels. If Phytolaccaceae were re lated to Gyrostemonaceae, one might hypothesize a polycarpic ancestry for Gyro stemonaceae. However, now that there is reasonable certainty that Gyrostemona ceae are not related to Phytolaccaceae or other Chenopodiales (Behnke, 1976), we may reasonably assume that Gyrostemonaceae with numerous carpels represent secondary increase in carpel number. Likewise, we may reasonably assume the monocarpellate Gyrostemonaceae to represent reductions. However, there is no clear evidence regarding the probable original carpel number of Gyrostemonaceae. There are four fruit types in Gyrostemonaceae. Cypselocarpus has achenes. Didymotheca has capsules dehiscing both loculicidally and septicidally, according to Walter (1909). Codonocarpus (FIGURE 47) and Gyrostemon (FIGURE 54) have samaroid follicles (or mericarps) which shatter from a central axis at maturity and probably are dispersed by wind without release of the seeds they contain. Tersonia (FIGUREs 61, 62) bears indehiscent syncarps, dry at maturity and covered with hook- like enations that suggest dispersal in animal fur. The variety in fruit type seems significant because seeds of all genera have the same form: semicircular in outline (the contained embryo is curved), with an aril (even in the seeds of Tersonia, in 312 ALLERTONIA 1:5 which seeds are not released from the syncarp), and with a ridged seed coat. These features are strongly reminiscent of Sapindaceae. Male flowers of Gyrostemonaceae are basically whorls composed of an indefi nite number of stamens (FIGURES 55, 57, 59). Some species of Gyrostemon have more than one whorl of stamens (FIGuRES 48, 51, 52), very likely a condition derived from the single-whorled pattern. Neither the number of carpellodes (which are rare ly evident in any case) or stamens or tepals give any useful hint as to the number of stamens basic to the family. Study of initiation of primordia might be helpful in this regard. One may hypothesize increase in stamen number concomitant with evolu tion of wind pollination. The perianth of Gyrostemonaceae consists of imbricate tepals (FIGUREs 49, 51, 53, 60). Although tepals of Bataceae and Gyrostemonaceae differ, one may note that both families have only a single perianth whorl. This condition is also reminis cent of the perianth of Stylobasium. The tepal number varies widely within Gyro stemonaceae, and is not constant even within an individual plant. Stylobasiaceae.—The polygamo-dioecious nature of Stylobasium was clarified by Prance (1965), although details of sexual conditions need investigation on the basis of field studies. The gynoecium consists of a single carpel with the micropyle of the ovule facing the basifixed style. There is no aril on the seed (FIGuRE 66). How ever, absence of an aril is to be expected because the fruit is an indehiscent drupe (FIGURE 65) and thus an aril would not be a functional part of a dispersal system. Prance (1965) notes that the stigma and numerous stamens are presumptive reasons for suspecting anemophily, but states that there is no field evidence. I can at this time furnish simple field observations on flowers of Stylobasium australe (FIGURE 63) which show anemophily inescapably. At anthesis, the filaments are very elongate and limp, so that anthers are pendent. Thus, the anthers are positioned as far away from the style and stigma as possible. An entomophilous flower must re tain a definite positioning of anthers with respect to a stigma, so that an insect con tacting the anthers will, upon visiting a flower, be presented with the same spatial separation of pollen-bearing and pollen-collecting surfaces. This is not possible with Stylobasium, where anthers, by virtue of the limp filaments to which they are at tached, cannot possibly retain a constant position with relation to the stigma and are not positioned so that any insect visiting the flower could possibly be expected to carry away pollen. The flower of S. spathulatum is undoubtedly similar; the flower shown in FIGuRE 64 is malformed. The flower in question was the only one found on a shrub which was otherwise in fruit. Very likely hormonal events leading to fruit formation resulted in thickening of filaments, which contained chlorenchyma, and their persistence despite the fact that the flower shown did not contain a fertiLe seed. There is no evidence of corolla or nectary development in Stylobasium. The synsepalous perianth, very likely calycine in nature, has lobes that open in imbricate fashion. The seed of Stylobasium corresponds very closely to seeds of Sapindaceae.

STIPULES A character of considerable potential significance occurs in Bataceae, Gyro stemonaceae, and Stylobasiaceae: presence of stipules. These stipules are never foliate, but are of the most vestigial sort, caducous or otherwise easily overlooked. Van Royen (1956) figures the stipules of Bails well. A typical gyrostemonaceous 1978 CARLQUIST: WOOD ANATOMY 313

stipule is shown here in FIGURE 50. Minute subulate stipules of the same sort are cited for Stylobasium by Prance (1965). They are more prominent on the seedling than on the adult plant, judging from his figures of S. spathulatum. Stipules are a pervasive feature of Sapindaceae; in that family also, stipules are minute and tooth- like. Undue emphasis should not be placed upon a single feature. However, one may note that families other than Sapindaceae proposed to have affinity with Bataceae and Gyrostemonaceae either have no stipules or have stipules of quite a different sort.

SUMMARY OF RELATIONSHIPS One may concede that Stylobasiaceae are, on the basis of all lines of evidence presently available, close to Sapindaceae, as Prance (1965) claims. If we include Stylobasiaceae in Sapindales, it adds some new dimensions, such as the herbaceous tendency and a clear instance of fiber dimorphism combined with storying of the shorter fibers, which must be termed axial parenchyma cells rather than libriform fibers. If one selects Sapindaceae as a family for comparison, one sees that Bataceae and Gyrostemonaceae show similarity in the following features: sexual condition (polygamo-dioecious and dioecious conditions occur in Sapindaceae such as Dodonaea, which also is anemophilous as are Bataceae and Gyrostemonaceae), presence of minute toothlike stipules, non-vestured pits in vessels, preponderance of procumbent cells in rays, diffuse axial parenchyma (present in Gyrostemonaceae but not in Bataceae), carpel morphology (one ovule, axile placentation in Gyro stemonaceae; Bataceae more distant in its pair of basal ovules in each of the two carpels, separated by a false septum), curved arillate seeds (arils apparently lacking in Bataceae), and curved embryos (embryo straight in Bataceae; Gyrostemonaceae has endosperm, Bataceae a little endosperm, and Sapindaceae no endosperm). If one searches for features that would show affinity with Capparales, one can list only a tendency toward herbaceousness and presence of glucosinolates. Since Stylobasiaceae can be added to Sapindales, herbaceousness can be said to be pres ent in Sapindales, however. Glucosinolates indubitably do occur in Capparales, in myrosin cells in that order, but not in myrosin cells in Bataceae and Gyrostemona ceae. If reinvestigation of Bretschneidera, now regarded as a monogeneric family of Sapindales, proves this genus to have myrosin cells, as has been claimed, glucosino lates may be present in Sapindales as well. A number of features cited could point to either Capparales or Sapindales equally as orders showing a greatest number of affinities with Bataceae and Gyro stemonaceae. Imperforate tracheary elements in Bataceae are fiber-tracheids; in Gyrostemonaceae, tracheids. Sapindaceae have fiber-tracheids or libriform fibers. In Capparales, Moringaceae have libriform fibers, Capparaceae fiber-tracheids or libriform fibers, whereas in Brassicaceae I reported libriform fibers (1971) although Metcalfe and Chalk (1950) claim “fibres. . . with minute bordered pits,” in which case tracheids would be present. The fiber dimorphism in Bataceae is very similar to that in either Stylobasium, such Capparales as Sinapidendron of the Brassica ceae, or, for that matter, other families (legumes, composites). The banded paren chyma of Gyrostemonaceae might be an instance of fiber dimorphism, in which case its storied nature is like that of Bataceae. If the parenchyma bands are not a 314 ALLERTONIA 1:5

FIGuREs 1—4. Wood sections of Bails maritima, from Carlquist in 1972, sn. 1. Transection of mature wood; banded apotracheal parenchyma present. 2. Tangential section; storied structure evident in axial parenchyma, vessel elements. 3. Perforated ray cell from radial section. 4. Portion of transection; rays at right and left, a group of apotracheal parenchyma cells visible. Magnifications shown by photographs of stage micrometer enlarged at same scale as photomicrographs to which they pertain. Scale for FIGURES I and 2, above FIGURE I (finest divisions = 10 pm). Scale for FIGuRE 3 above FIGURE 3 (divisions 10 Mm). Scale for FIGuRE 4 above FIGuRE 4 (divisions 10 pm). 1978 CARLQUIST: WOOD ANATOMY 315

1_I :rI

• i. -i ‘‘ :

FIGuRES 5-9. Wood sections of Boris and Codonocarpus. 5 and 6. Bails maritima, from Cariquist in 1972, sn., portions of radial sections. 5. Air-filled fiber-tracheid, showing vestigial borders on pits. 6. Air-filled ray cell, showing bordered nature of pits. 7-9. Codonocarpus cotinUblius, from Car/quist 5150. 7. Radial section, showing procumbent ray cells. 8. Transection; earlywood vessels above, vessels sparse in earlywood below. 9. Tangential section; rays notably wide. FIGuREs 5 and 6. magnification scale above r FIGURE 5 (divisions 10 urn). FIGUREs 7—9, scale above FIGURE 1. 316 ALLERTONIA 1:5

FIGURES 10-13. Wood sections of Gyrostemon. 10 and II. Gyrostemon ramulosus, from Car/quEst 3153. 10. Transection, showing parenchyma bands. 11. Tangential section; storied cells are axial paren chyma. 12 and 13. Gyrostemon sheathii, from Carlquist 6010. 12. Transection; diffuse as well as para tracheal parenchyma cells abundant. 13. Tangential section; storied cells (left) are axial parenchyma, mostly in strands of two cells. FIGUREs 10—13, magnification scale above FIGURE I. 1978 CARLQUIST: WOOD ANATOMY 317

FIGURES 14—17. Wood sections of Gyrosremon subnudus, from Carlquist 5518. 14. Transection, show ing large groups of vessels. 15 Tangential section; both fibers and storied axial parenchyma evident. 16. Radial section; almost ray all cells are procumbent except for tip and some sheathing cells (compare with FIGURE 15). 17. Portion of tangential section; alternate intervascular pitting at left; scalariform-like vessel-parenchyma pitting (slightly out of focus), right. FIGURES 14—16, magnification scale above FIGURE I. FIGURE 17, scale above FIGURE 3. 318 ALLERTONIA 1:5

FIGUREs 18-21. Wood sections of Tersonia brevipes, from Carlquist 5385. 18. Transection of root; axial parenchyma, both paratracheal and diffuse, abundant. 19. Tangential section of root; axial paren chyma in strands of two cells. 20. Tangential section of stem; all imperforate axial elements shown are storied. 21. Radial section of stem; all ray cells shown are procumbent. FIGuREs 18—21, magnification scale above FIGuRE 1. 1978 CARLQUIST: WOOD ANATOMY 319

FIGUREs 22-27. Wood sections of Stj’lobasium australe, from Carlquist 5434. 22. Transection; radial files of vessels evident. 23. Tangential section; rays are narrow, numerous. 24. Radial section; heterocel lular histology evident. 25. Portion of transection; inner walls of libriform fibers are gelatinous. 26. Por tiOn of radial section; deposits of dark-staining materials outline bordered nature of pits in ray cells. 27. Portion of radial section; numerous small tyloses are visible in the two vessels shown. FIGURES 22—24, magnification scale above FIGURE 1. Scale for FIGUREs 25—27 above FIGuRE 3.

31. above 31 FIGURE FIGURe

for Scale 4. above 30 FIGURE Scale for FIGuRE 1. scale above FIGuRE magnification 29, and 28 FIGURES

evident. is of rays and the axial parenchyma of pattern storied section; Tangential cells. 31. parenchyma

apotracheal containing xylem fascicular is the center in Transection; 30. parenchyma. axial is band

diagonal pale the uniseriate; mostly are rays section; Tangential 29. light. appear parenchyma bands of

5453. Transection; Carlquist 28. from spathulatum, Stylobasium of Sections 28-31. Wood FIGURES

1:5 ALLERTONIA 320 1978 CARLQUIST: WOOD ANATOMY 321

FIGURES 32-36. Wood sections of Dodonaea burmanniana (Sapindaceae), from Aw-8833. 32. Tran section; vessels tend to be in radial files; axial parenchyma mostly paratracheal. 33. Tangential section; massive deposits of amorphous materials in rays. 34. Radial section; all ray cells are procumbent. 35. Ray cells from radial section; dark-staining deposits demonstrate the bordered nature of pits. 36. Portion of transection; a few axial parenchyma cells may be seen, upper left. FIGUREs 32-34, magnification scale above FIGURE 1. FIGURES 35 and 36, scale above FIGuRE 3. 322 ALLERTONIA 1:5

FIGURES 37-41. Wood sections of Thouinidium decandrum (Sapindaceae), from Yw-1198. 37. Tran section; dark-staining deposits evident in ray cells, vessels, and axial parenchyma cells. 38. Tangential section; paratracheal axial parenchyma cells surround vessels. 39. Portion of transection; large numbers of apotracheal parenchyma cells may be seen in addition to paratracheal parenchyma. 40. Radial sec tion; ray cells exclusively procumbent. 41. Portion of radial section; chambered cells of axial parenchyma strand contain rhomboidal crystals. FIGUREs 37, 38, 40, magnification scale above FIGURE 1. FIGuRE 39, scale above FIGURE 31. FIGURE 41, scale above FIGURE 3. 1978 CARLQUIST: WOOD ANATOMY 323

FIGURES 42-45. Wood sections of Toulicia bullata (Sapindaceae), from Yw-22051. 42. Transection; prominent apotracheal parenchyma bands are present. 43. Portion of transection; diffuse as well as para tracheal and apotracheal banded axial parenchyma cells visible. 44. Tangential section; rays are uniseri ate or biseriate. 45. Radial section; apotracheal parenchyma strands differ in length from paratracheal strands; afl ray cells procumbent. FIGURES 42, 44, 45, magnification scale above FIGURE I. FIGuRE 43, scale above FIGURE 31.

4. X lobes, perianth

imbricate

bases,

decurrent with leaves succulent showing plant, female of Branch A. W. Esperance,

near

6010,

from

Carlquist

from

sheathii, Gyrostemon 49. X 1.5. branches, efoliate on occur which ers,

flow of male

Habit W. A.

Coorow, near 5210, from Carlquist from racemigerus, Gyrostemon 48. 1.5. X

attached,

are

(mericarps) the follicles which to receptacle concave the showing fruits, of Habit 5150.

cotinUolius, Cariquist

from

Codonocarpus 47. W.A. Rawlinna, of north tall; m. 4 trees of pair a of Habit

5150.

Car1quist from

C’odonocarpus 46. cotinf1ius, Gyrostemon. and Codonocarpus 46-49. FIGURES

1:5 ALLERTON1A 324 1978 CARLQUIST: WOOD ANATOMY 325

FiGuRes 50—54. Gyroslemon ramulosus, from Carlquisi 5151, from near Neale Junction, W. A. 50. Node, showing minute stipule at base of axillary branch, X 5.5. 51. Male flower, to show imbricate nature of perianth, X 2.2. 52. Male flower, to show arrangement of anthers, X 2.2. 53. Branch with female flow ers in various stages of development, with stigmas prominent, X 1.5. 54. Branch with fruits; note pore at tip of fruit, X 1.5. 326 ALLERTONIA 1:5

FIGURES 55-58. Gyrosiemon and Cypselocarpus flowers. 55 and 56. Gyrostemon lepperi, from Carl quist 5161, from near Warburton, W. A. 55. Branch with male flowers, showing cycle of anthers and ab sence of carpellodes. 56. Branch with female flowers; perianth vestigial, the stigma attached subterminal ly. 57 and 58. Cvpselocarpus haloragoides, from Carlquist 5758, from near Esperance, W. A. 57. Branch with sessile male flowers and succulent leaves. 58. Branch with monocarpellate female flowers, illus trating the broad oblique stigma. All X 4. 1978 CARLQUIST: WOOD ANATOMY 327

FIGuREs 59—62. Tersonia brevipes, from Cariquisi 5458, from near Medina, W. A. 59. Portion of male inflorescence; filaments and carpellodes are not evident, X 2.2. 60. Female flower in leaf axil; perianth imbricate, X 4. 61. Fruit on branch, showing fimbriate enations, X 1.2. 62. Same fruit in transection; developing seeds visible, X 1.2.

X 4. All illustrated. leaf

of

tip

emarginate seed; curved

funiculate showing fruit, of 66. Longisection perianth. prominent showing

maturity, near

65. Fruit perianth. imbricate note carpel; and anthers of persistence in abnormal ably

prob

male

flower, Presumptively 64. A. W. Northampton, near 5453. from Carlquist from spathulatum,

StyIobasium

64—66. stigma. prominent note anthesis; at male flower Presumptively A. W. Morawa, near

5434, from

Carlquist from

australe, Stylobasium 63. fruits. and flowers Sty/obasium 63-66. FiGuRIs

:

1:5 ALLERTON1A 328 ______•______

1978 CARLQUIST: WOOD ANATOMY 329

product of fiber dimorphism, similarities to axial parenchyma in Sapindaceae might be cited. Imbricate tepals, tending to be united at the base, are a feature of Gyro stemonaceae tending to show resemblance to Sapindaceae; tepals of Bataceae are separate, and thus perhaps like sepals in Capparales. The tendency toward cam pylotropous ovules in Gyrostemonaceae (and perhaps Bataceae) is matched by either Sapindaceae or Capparales. Thus the preponderance of features seems to suggest that Bataceae and Gyro stemonaceae may be sapindalean, but each quite a distinct outlier family within that order. While Bataceae resemble Gyrostemonaceae in a few features (storying of parenchyma bands; abundance of procumbent cells in rays; ultrastructure of pollen grain exine pertectate; glucosinolates present but not in myrosin cells), they are amply distinct from each other. A discussion of genera with some degree of affinity to Sapindaceae would not be complete without mention of Emblingia. Emblingia is a monotypic genus of herbs from Western Australia. Its affinities have been examined in a symposium (Erdtman et al., 1969). This symposium is devoted to floral anatomy and morphology, vegeta tive anatomy, and pollen morphology and ultrastructure. The conclusions voiced by the respective authors of this symposium are remarkably disparate. This may be attributed to incompleteness of material, but perhaps more significantly to the marked modifications this genus must represent in comparison to its closest rela tives. Careful reading of that symposium does not provide a clear decision as to whether the balance of evidence favors polygalalean, sapindalean, or goodeniaceous affinity. If the original description of the species by Mueller is accurate, the arillate seed with curved embryo found in Emblingia (Liens, in Erdtman et al., 1969) is per haps persuasive of sapindalean affinity.

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