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Home , Allocasuarina, Allocasuarina nana, Casuarina, Casuarinaceae, Gymnostoma

Some Morphological Features for Generic Characterization Among the Author(s): John G. Torrey and R. Howard Berg Source: American Journal of Botany, Vol. 75, No. 6 (Jun., 1988), pp. 864-874 Published by: Botanical Society of America Stable URL: http://www.jstor.org/stable/2444006 . Accessed: 23/08/2011 15:52

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http://www.jstor.org Amer. J. Bot. 75(6): 864-874. 1988.

SOME MORPHOLOGICAL FEATURES FOR GENERIC CHARACTERIZATION AMONG THE CASUARINACEAE'

JOHN G. TORREY AND R. HOWARD BERG Harvard Forest, Petersham, Massachusetts 01366; and Biology Department, Memphis State University, Memphis, Tennessee 38152

ABSTRACT The family Casuarinaceae has been divided recently into four genera on the basis of differences in chromosome numbers, morphological characters including stem morphology, structures of the male and female and , their modem biogeography, and record. All members of the family are characterized by highly reduced , photosynthetic deciduous branchlets, and adaptations to xeric or hot humid environments. The is characterized by multiple rows of naked stomata along the central vertical faces of the stem internodes. and develop vertically aligned rows of stomata in deep furrows that alternate with stem ridges. Epidermal hairs, formed within the furrows, occur in different species in conformity with other xeric characters. The characters described in this paper conform to the view that the morphological structures represent reduction and special- ization rather than primitiveness.

THE FAMILY Casuarinaceae is comprised of four 1981). All members of the family show highly genera: Allocasuarina L. Johnson, Casuarina reduced vegetative morphology with reduced Adans., Gymnostoma L. Johnson and a fourth scale-like leaves and needle-like branchlets that genus, , still to be described serve as photosynthetic organs. The reduction (Johnson and Wilson, 1981; Johnson, 1982, or simplification of the vegetative axis char- in press). The family is a large group native to acterizes the xeromorphy of the shoot in this , Malaysia and Polynesia. Members family whose growth habits range from low- of the genus Casuarina sensu stricto have been growing woody a meter or so in height disseminated by man to most parts of the trop- (e.g., ), to tall in excess ical and subtropical world where they serve as of 35 meters in height as in Casuarina cun- important sources of fuel wood (National ninghamiana or C. grandis (Doran and Hall, Academy of Sciences, 1980), and multi-use 1981). According to Moseley (1948), members trees for windbreaks, dune stabilization and of the Casuarinaceae were derived in brackish for soil improvement by virtue of their sym- areas, sand dunes or dry, marginal, low-nu- biotic association with the -nodulating ni- trient sites that were either physiologically or trogen-fixing actinomycete (Midgley, climatically arid environments. After study of Turnbull, and Johnston, 1983; National Acad- structures within the family, Moseley conclud- emy of Sciences, 1984). ed that their morphological characteristics rep- Taken as a whole, the family probably in- resent reduction rather than primitiveness. cludes some 90 species distributed among the Of special interest in the group is the de- four genera in the following ratio: Allocasua- velopment of cuticular modifications accom- rina 54 spp., Casuarina 16 spp., Gymnostoma panying their xeromorphic characters, the 18 spp. and Ceuthostoma, 2 species (Johnson, variation in distribution of stomata on the 1982, and personal communication). Separa- branchlets and reduced scale leaves, and the tion of the four genera is based on a series of occurrence and distribution of epidermal hairs characters including chromosome numbers, (Withers, 1978). Five species among three of morphology of the vegetative shoot, including the four genera in the Casuarinaceae have been the distribution of stomata on the branches, selected as representative of the morphological structures of the male and female inflores- characteristics among the major groups. Some cences and fruits together with their biogeog- of their distinctive features based on scanning raphy and fossil record (Johnson and Wilson, electron microscopy (SEM) are presented here.

MATERIALSAND METHODS-For the most ' Received for publication 16 February 1987; revision part, material for SEM was taken from living accepted 26 August 1987. This research was supported in part by the Maria Moors started from and grown in the Cabot Foundation for Botanical Research of Harvard Uni- greenhouse at the Harvard Forest in Peter- versity and by USDA Grant 8 3-CRCR- 1-1 315 to RHB. sham, Massachusetts.

864 June 1988] TORREY AND BERG-CHARACTERIZATION AMONG CASUARINACEAE 865

Sources of seed were as follows: (1950), Flores (1977, 1978, 1980), Barlow Miq. From col- (1983), and Woodall and Geary (198 5). lection by Division of Forest Research, CSI- The stem comprises two types of structures: RO, Canberra, Australia. indeterminate persistent branches which undergo secondary thickening and form the Forst. & Forst. Col- lected from coastal sand dunes at Indialan- permanent aboveground body and de- tic, . Figures 3-6 were photographs terminate deciduous branchlets that have ra- of shoots from tissue cultures derived from dial, jointed stems, possessing green photo- axenically-grown seedlings. synthetic tissues and widely distributed Allocasuarina lehmanniana (Miq.). L. John- stomata. The deciduous branchlets are the son. made available from collections photosynthetic organs of the plant, rather than of the King's Park Botanical Garden, Perth, the scale leaves. They are characterized by Western Australia. ribbed and furrowed longitudinal internodes Allocasuarina decaisneana (F. Muell.) L. with nodes defined by whorls of scale leaves. Johnson. Collected from trees near Alice In Gymnostoma the stems are always quad- Springs, , Australia. rangulate but such 4-sided branchlets occur Gymnostoma papuanum (S. Moore) L. also in some species of Allocasuarina. Johnson. Collected from trees at The Waiak- Leaves are reduced lateral appendages, white ea Arboretum, U.S. Forest Service, Institute or brown and scale-like. The symmetrical lam- of Pacific Islands Forestry, near Hilo, Ha- ina is linear or lanceolate with 1 or 3 veins. waii. The leaves in whorls are fused laterally at their bases, and alternate at successive nodes with For scanning electron microscopy shoots those below. were fixed in 2.5% redistilled glutaraldehyde Stomata occur in characteristic fashion in in 0. 1 M Na cacodylate buffer at pH 7.2. After the different genera, typically oriented trans- primary fixation for 1.5-2.5 hr, the tissue was versely with respect to the long axis of the rinsed 30 min in 3 changes of buffer and post- internode and either within deep vertical fur- fixed in several changes of buffered 2% osmium rows or grooves running longitudinally along tetroxide at 25 C for 48 hr. Specimens were the internodes or slightly sunken on the flat critical-point dried after dehydration to 100% face of the branchlet stem surface. Epidermal ethanol. After being coated with gold, samples hairs are typically uniseriate either branched were examined in a Hitachi S-450 SEM, using or unbranched, and associated with stomatal an accelerating voltage of 20 kv. distributions in furrows or on adaxial faces or leaf edges. The epidermis of the stem typ- OBSERVATIONS-In Table 1 are summarized ically shows elaborate, waxy surface incrus- the major features of these five species brought tations with heavy cuticle and warty surface together from available sources and from the modifications. SEM observations made during this study. The The number of scale leaves per whorl at each chromosome numbers of the genera are listed node is not a particularly useful taxonomic according to Barlow (1983) and reflect the dis- character within the Casuarinaceae (Doran and tinctness of the three different genera. Accord- Hall, 1981). In Casuarina and Allocasuarina ing to Barlow, Gymnostoma may be the least the number of scale leaves and leaf ridges may specialized with Allocasuarina the most spe- range from 4 to about 16; in contrast, in Gym- cialized. The geographical occurrence of these nostoma the branchlets are consistently 4-sid- species and the climatic conditions are taken ed, the number of leaves 4 and the stomata are from Doran and Hall (1981) and from Johnson located on the four faces of the branchlets as (1982). The sites and climates epitomize the will be described below. In Table 1 are given differences in the distributions of these genera the number of scale leaves per leaf whorl found and further emphasize the distinctions among on the specimens described in this study. the genera. Two characters of particular interest in de- There exist much discussion and controver- fining these genera, i.e., the occurrence of sto- sy over the appropriate descriptive terminol- mata and epidermal hairs which are described ogy for the vegetative features of these genera. in this paper, are listed in summary form in The following general description of the veg- Table 1 and are illustrated in the SEM's pre- etative body utilizes terms which seem most sented in the figures. Emphasis will be placed acceptable based on existing evidence. For ex- on stomatal distribution on the stem or branch- tended discussions of the features of the veg- let surfaces and on the adaxial surfaces of the etative axis one is referred to the published leaf scales. The occurrence of epidermal hairs, work of Moseley (1948), Metcalfe and Chalk especially in relation to the stomata, is also 4RO~~~~~~~~~EcIm~~~~~~~~I

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Fig. 1-4. Casuarina equisetifolia stem and leaves of deciduousbranchlet. 1. Nodal regionof stem showingalternation of leaves in successive nodes. Epidermalhairs (EH) erupt from intemodal grooves (IG). x 70. 2. Transverseface of stem showing 7 ribs and furrowswith stomata lining grooves. x 100. 3. Rows of stomata (S) in grooves (IG) extending to scale leaf (SL)bases. Note lack of epidermalhairs. x 80. 4. Adaxial surfaceof scale leaf (SL) showing2 double rows of stomata (S) along either side of midrib. x 250. June 1988] TORREY AND BERG -CHARACTERIZATION AMONG CASUARINACEAE 867

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Fig. 5, 6. Casuarina equisetifolia leaf epidermis showing stomata. 5. Epidermalcells, including guard cells (GC) lackingwaxy deposits. x 1,200. 6. Epidermalcells surroundingstomata (S) encrustedwith waxy deposit. x 1,350. described for each species. Finally, comments hidden by epidermal hairs (Fig. 8, 9). Scale are made on the nature of epidermal modifi- leaves typically exhibit epidermal hairs along cations and appearanceof the abaxial surfaces the distal edges of the leaves (Fig. 7) continuing of scale leaves and branchlet stems. up from the stem grooves. The adaxial face of the scale leaves shows two parallel uniseriate Casuarina species-In Fig. 1-6 are shown rows of stomata unadorned by hairs (Fig. 7). distinctive featuresof the stem and scale leaves Beading of the stem epidermal cells and the of C. equisetifolia.Seven scale leaves arisefrom midrib cells is usual. These two Casuarina the stem formingseven ridges, visible in cross- species have in common, therefore, the dis- section of the stem (Fig. 2), with deep grooves tribution of stomata in deep grooves in the between the ridges, each groove arrangedwith stem surface and limited occurrence of epi- two double or triple vertical rows of stomata dermal hairs in relation to the groove. facing inward along the grooves (Fig. 3). Epi- dermal hairs which are absent on the scale Allocasuarina species-Both A. Iehman- leaves (Fig. 4) occur along the lower third of niana, which occurs in western Australia,and the grooves between internodes (Fig. 1) but are A. decaisneanain centralAustralia show mor- lackingalong the upperportions ofthe grooves. phological characterspresumed to be adapta- The adaxial face of each scale leaf shows two tions to the hot, seasonally dry sites in which parallelrows of stomata arrangedtypically in they may be found. Both show elongate epi- two rows each of alternatingstomata (Fig. 4). dermal hairs clustered around and along deep Thus along much of the length of the green grooves in which stomata occur. In A. leh- photosynthetic stem, the stomata face into a manniana the vertical grooves of the stem are partiallyenclosed groove, allowing for gas ex- lined with stomata in scattered vertical rows change but probably reducing moisture loss. (Fig. 12) and are partiallycovered by branched The adaxial surfaces (Fig. 5) show less epicu- epidermal hairs (Fig. 12). Elongate epidermal ticular wax deposition than abaxial surfaces hairs edge the scale leaves and the adaxial sur- around stomata (Fig. 6). faces of the scale leaves over and around the In C. cunninghamiana vertical grooves be- two rows of stomata (Fig. 13). The stem ridge tween stem ribs are less evident (Fig. 9, 10) midribsare prominentand the surfacesbeaded than in C. equisetifolia.Stomata occur in ver- (Fig. 11, 14). tical rows within these grooves almost totally Allocasuarinadecaisneana, which grows in June 1988] TORREY AND BERG -CHARACTERIZATION AMONG CASUARINACEAE 869

Fig. 7-10. Casuarinacunninghamiana stem and leaves of deciduous branchiet.7. Adaxial surfaceof scale leaves (SL), showing2 single rows of stomata (S) parallelingmidrib and numerousepidermal hairs along leaf edge. x 160. 8. Enlargedview of internodalgroove showing stomatal rows within groove and emergingepidermal hairs (EH). x 250. 9. Transverseview of stem showing 10 shallow grooves alternatingwith stem ribs. Note epidermal hairs emerging from grooves. x 90. 10. Scale leaves with epidermal hairs merge into the internodalribs separatedby grooves (IG). Grooves show emergentepidermal hairs. Scale leaves and internodalribs have a midrib (M). x 90. 870 AMERICAN JOURNAL OF BOTANY [Vol. 75

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Fig. 11-14. Allocasuarinakehmanniana stem and leaves of deciduous branchiet. 11. Node and portions of two intemnodesof branchietshowing prominent ribbing and deep grooves.Leaves and ribsalternate at each node. Epidermal hairs emergefrom the scale leaves. x 70. 12. Enlargedview looking into intemnodalgroove (IG) showing multiplerows of stomata (S) with branchedepidermal hairs (EH) overarching.x 340. 13. Adaxial surfaceof scale leaves showing 2 rows of spaced single stomata hidden by epidermalhairs (EH). Small epidermalhairs are at edges of leaves. x 140. 14. Transverseview of stem showing deep grooves (IG) alternatingwith prominentlyribbed internodal surfaces of stem. x 90. June 1988] TORREY AND BERG-CHARACTERIZATION AMONG CASUARINACEAE 871

Fig. 15-18. Allocasuarinadecaisneana stem and leaves of deciduousbranchiets. 15. Node of stem with prominent scale leaves (SL)and alternatingintemnodal tissues above. Note abundantepidermal hairs (EH)exerted from internodal grooves and on adaxialface of scale leaves. x 50. 16. Enlargedview of internodalgroove showingepidermal hairs and rows of stomatawithin groove. x 460. 17. View into internodalgroove (IG) showingmultiple verticalrows of stomata (S) and overarchingepidermal hairs (EH). x 260. 18. Adaxial view of scale leaf covered with epidermal hairs (EH) arisingfrom cells around verticallyoriented rows of stomata (S). x 190. 872 AMERICAN JOURNAL OF BOTANY [Vol. 75

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AP$ June 1988] TORREY AND BERG-CHARACTERIZATION AMONG CASUARINACEAE 873 one of the driest habitats in central Australia, the presence of numerous deciduous, needle- shows elaborated epidermal hairs along the like branchlets with highly reduced leaves, stem grooves (Fig. 15-17) surrounding and members of this family have a deceptively co- overarching the stomatal rows that run lon- nifer-like habit. This impression is augmented gitudinally in the sunken grooves of the stem. by the occurrence of cone-like infructescences. Epidermal hairs cover the lower half of the On careful examination one observes its basic adaxial face of the scale leaves (Fig. 15), nearly angiospermous traits. The simplified and re- obscuring the two rows of stomata (Fig. 17, duced male and female flowers are unisexual, 18). The paired stomatal rows extend nearly borne on plants that are monoecious or dioe- to the tip of the scale leaves where epidermal cious. These plants are adapted to wind pol- hairs are lacking (Fig. 15, 18). Scale leaves are lination. The family has been considered prim- rounded abaxially showing no midrib but epi- itive within angiosperms or in more recent years dermal cells are covered with beaded cuticular as moderately specialized and reduced. (For incrustation. the review of these arguments, consult the pa- pers by Moseley, 1948; Flores, 1977, 1978, Gymnostoma species-Although less famil- 1980; Flores and Moseley, 1982; and Barlow, iar because, unlike Casuarina spp., Gymno- 1983.) stoma has not been distributed worldwide by Of special interest have been the morpho- man, it is a large genus comprised of about logical and physiological adaptations to a great eighteen species (Barlow, 1983). Gymnostoma range of soils and climates. One can name is restricted to the hot, humid tropics on the species within the family, many of them en- islands of the South Pacific and Malaysia. The demics, adapted to every one of the fifteen genus, as the name given by Johnson (1980) described ecosystems on continental Australia adapted from a term used by Poisson (1871) (Specht, Roe, and Boughton, 1974) and still indicates, is distinctive in having naked sto- others on the South Pacific islands and in Ma- mata located in vertical rows along each face laysia. In part because of their capacity to form of the quadrangular stem. Each row consists dinitrogen-fixing root nodules in response to of two to five axial series of stomata (Fig. 19, infection by the soil actinomycete, Frankia 20, 22). In Gymnostoma papuanum sections (Torrey, 1982), these plants survive in a wide of the stem (Fig. 20, 23) show that the stomata range of nutrient-poor soils. open internally into large intercellular cham- Particularly striking are the morphological bers that facilitate gas exchanges between the characters involving presumed adaptations to atmosphere and the green photosynthetic tis- xeric or moist environments. Justification for sues of the stem, an adaptation lacking in the the generic segregation on morphological and more xeromorphic genera of the family. Sto- other grounds is summarized by Barlow (1983) mata also occur almost at random in vertical and much further detailed information prom- arrays on the adaxial surfaces ofthe scale leaves ises to be forthcoming from the studies of L. (Fig. 21). Epidermal hairs are reduced and re- A. S. Johnson and his associates. The remark- stricted to the distal edges of the scale leaves able reduction of the leaf and the question of (Fig. 19, 21). The prominent "midrib" is interpreting the internodal tissue as axial or as rounded, smooth and shows little epidermal related to leaf bases early led to coining the cuticular development. The morphology re- term "phyllichnium" (Loew, 1865) to desig- flects adaptations to a moist environment. nate the single leaf base-like components of internodes. More recently, references to the DISCUSSION-Genera of the Casuarinaceae photosynthetic branchlets as "cladodes" have pose intriguing problems for the morphologist, been criticized (Rao, 1972; Flores, 1977) and systematist, physiologist and biologist inter- discarded by them. Arguments over termi- ested in biogeography and evolution. Due to nology in this group reflect the uniqueness of

Fig. 19-23. Gymnostoma papuanum stem and leaves of deciduous branchlet. 19. Node of stem showing scale leaves alternating with midribs of adjacent internode. Multiple rows of stomata are distributed along the slightly depressed stem surfaces between prominent ridges of the quadrangulate stem. x 70. 20. Transverse section of 4-sided stem showing rows of naked stomata on the stem surface and elaborate substomatal cavities (SSC) within the stem. x 170. 21. Adaxial surfaces of scale leaves show multirowed stomata distributed parallel to midrib region. Short epidermal hairs occur at the leaf edges. x1 40. 22. Waxy deposits on epidermal surfaces of naked stomata on stem. x 600. 23. Longitudinal section of portion of stem internode showing the substomatal cavities (SSC) beneath stomatal rows. x 140. 874 AMERICAN JOURNAL OF BOTANY [Vol. 75 the morphologies involved. Descriptive terms * In press. Notes on Casuarinaceae III. The genus used here appear to be the simplest that are Ceuthostoma.Telopea. modern students of the , AND K. L. WiLsoN. 1981. Casuarinaceae, then useful and accepted by and now. XIII Int. Bot. Congr. Abstr. 278. family. LOEW,E. 1865. (cited by Flores,1978) De casuarinearum Stomatal occurrence, distribution and as- caulis foliique evolution et structure. Dissertatio in- sociated epidermal modifications, such as epi- auguralis botanica. Berolini. dermal hair formation which are emphasized METCALFE, C. R., AND L. CHALK. 1950. Anatomy of the in this SEM study, all suggest adaptation either . Vol. 2. Oxford University Press, Ox- to humid environments and offer ford. to xeric or MIDGLEY, S. J., J. W. TURNBULL, AND R. D. JOHNSTON one more set of characters which, together with [eds.]. 1983. Casuarina ecology, management and other criteria, present additional bases for utilization. CSIRO, Melbourne, Australia. Johnson's (1982) generic separations and con- MOSELEY,M. F., JR. 1948. Comparative anatomy and form to the view that these structures represent phylogeny of the Casuarinaceae. Bot. Gaz. 110: 231- reduction and specialization rather than prim- 280. NATIONALACADEMYOFSCIENCES. 1980. Firewood crops. itiveness. National Academy Press, Washington, DC. . 1984. : nitrogen-fixing trees for ad- LITERATURE CITED verse sites. National Academy Press, Washington, DC. POISSON, I. 1871. (cited by Flores, 1977) Recherches sur and bio- BARLOW,B. A. 1983. Casuarinas-a taxonomic les Casuarina et en particulier sur ceux de la Nouvelle W. Turnbull, geographic review. In S. J. Midgley, J. Cal6donie. Nouv. Arch. Mus. Hist. Nat. Paris. Ser. I man- and R. D. Johnston [eds.], Casuarina ecology, 10: 59. 10-18. CSIRO, Melbourne, agement and utilization, RAO, A. N. 1972. Anatomical studies on succulent Australia. cladodes in C. equisetifolia L. Proc. Indian Acad. Sci. fifteen Aus- DoRAN, J. C., ANDN. HAnL. 1981. Notes on LXXVI Sec. 86: 262-270. species. Forest Research, CSIRO, tralian Casuarina SPECHT,R. L., E. M. ROE, AND V. H. BOUGHTON[eds.]. Canberra, Australia. 1974. Conservation of major plant communities in in Casuarina FLORES,E. M. 1977. Developmental studies Australia and Papua . Austral. J. Bot. mature (Casuarinaceae). III. The anatomy of the Suppl. Ser. 7. CSIRO, Melbourne, Australia. 25: 65-87. branchlet. Rev. Biol. Trop. TORREY,J. G. 1982. Casuarina: actinorhizal dinitrogen- of Casuarina (Casuari- . 1978. The shoot apex fixing of the tropics. In P. H. Graham and S. C. 26: 247-260. naceae). Rev. Biol. Trop. Harris [eds.], Biological technology and phyllotaxis of . 1980. Shoot vascular system for tropical agriculture, 427-439. CIAT, Cali, Colom- J. Bot. 67: 131- Casuarina (Casuarinaceae). Amer. bia. 140. WITHERS,J. R. 1978. Studies on the status of unburnt JR. 1982. The anatomy of , ANDM. F. MOSELEY, woodland at Ocean Grove, Victoria. III. and of Casuarina the pistillate Comparative water relations of the major tree species. Amer. J. Bot. verticillata Lamarck (Casuarinaceae). Austral. J. Bot. 26: 819-835. 69: 1673-1684. WOODALL,S. L., AND T. F. GEARY. 1985. Identity of on Telo- JOHNSON,L. A. S. 1980. Notes Casuarinaceae. Florida casuarinas. USDA Forest Service SE Forest pea 2: 83-84. Exp. Sta. Res. Note SE-332: 1-10. . 1982. Notes on Casuarinaceae II. J. Adelaide Bot. Gard. 6: 73-87.