Reinvestigations of the Wood Anatomy of Duabanga and Sonneratia with Particular Reference to Their Systematic Position

IAWA Bulletin n.s., Vol. 8 (4),1987 337

REINVESTIGATIONS OF THE WOOD ANATOMY OF DUABANGA AND SONNERATIA WITH PARTICULAR REFERENCE TO THEIR SYSTEMATIC POSITION

R. Vijendra Rao, Babulal Sharma, Laxmi Chauhan and R. Dayal Wood Anatomy Branch, Forest Research Institute & Colleges, Dehra Dun 248006, India

Summary The wood anatomy of Duabanga and Sonne swarlu and Rao (1964) suggested that these ratia is described in detail. Based on the present two genera be treated as a subfamily within the investigation as well as on the well recognised Lythraceae while Van Vliet and Baas (1984) differences between these two genera, their sys- treated them as a family. tematic position is discussed. The study endorses These two genera along with other families the suggestion made by Dahlgren and Thorne form the 'nucleus' or 'core' of the Myrtales, all (1984) to raise the status of these two genera characterised by two important anatomical fea- to two subfamilies within the Lythraceae. The tures: intraxy lary phloem and vestured inter- ecological differences between these two genera vascular pits. in relation to their xylem anatomy is also dis- So far as the closely related families, Lythra- cussed. ceae, Punicaceae and Sonneratiaceae of the Key words: Sonneratiaceae, systematic wood Myrtales are concerned, the vessels have been anatomy, ecological trends. shown to have only simple perforations (Ven- kateswarlu & Rao, 1964; Baas & Zweypfenning, Introduction 1979; Van Vliet & Baas, 1984). Even in the en- Sonneratiaceae Engl. are a small family of tire myrtalean complex, presence of scalari- two genera and seven species distributed in form and reticulate type of perforations has tropical East Africa and Asia to Australia and been reported only in a few genera of the Myr- the West Pacific (Airy Shaw, 1966). taceae (Schmid & Baas, 1984). However, Baas In recent times no other order in dicotyle- and Zweypfenning (1979) found only once a dons has been subjected to such an extensive vestigial bar in the first formed rings of the wood anatomical study as the Myrtales (Venka- secondary xylem of Cuphea carthagensis, a teswarlu & Rao, 1964; Mujica & Cutler, 1974; member of the Lythraceae. Van Vliet, 1975, 1976, 1979, 1981; Van Vliet The present investigation pertains to a de- & Baas, 1984; Carl quist, 1975a; Carlquist & De tailed anatomical study using many samples of Buhr, 1977; Bridgwater & Baas, 1978; Baas, Duabanga and Sonneratia. On the basis of qua- 1979, 1986; Baas & Zweypfenning, 1979; litative and quantitative anatomical differences Koek-Noorman et aI., 1979; Ter Welle & Koek- found between these two, the systematic posi- Noorman, 1981; Schmid & Baas, 1984). tions of these genera is discussed in the light of The systematic position of Duabanga Buch.- the suggestions made by Dahlgren and Thorne Ham. and Sonneratia L. f. which forms a part (1984) and Van Vliet and Baas (1984). of the Myrtalean complex, has been the subject of discussion. Earlier, Bentham and Hooker (1862) have placed these two genera in Lythreae Materials and Methods of the Lythraceae. However, Thorne (1968, The following are the wood samples avail- 1976, 1981) and Dahlgren (1975) kept these able at the Forest Research Institute Xylarium two genera in the Lythraceae, but in a separate used for detailed wood anatomical studies: subfamily Sonneratioideae. Dahlgren & Thorne 1. Duabanga grandif70ra (Roxb. ex DC.) Walp. (1984) later suggested an independent subfamily (syn. D. sonneratioides Buch.-Ham.): DDw status, Duabangoideae and Sonneratioideae but 1995, Andamans; DDw 8237 BH* /627/ within the Lythraceae. Contrary to these, many 151124, Arunachal Pradesh; DDw 950, DDw systematists accorded a family status, Sonnera- 1230, DDw 1285, DDw 1436, DDw 5650, tiaceae to both these genera (Engler & Gilg, as DDw 7286 BHj73227, DDw 7554, DDw 8011, quoted by Emberger, 1960; Melchior, 1964; DDw 8012, DDw BH (herbarium sheet lost Soo', 1967, 1975; Cronquist, 1968; Takhtajan, 1969; Hutchinson, 1973; Dahlgren, 1980). On * BH = herbarium-backed specimen, followed the basis of wood anatomical studies Venkate- by collection number of herbarium voucher.

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due to damage), DDw 8369 BH/151149, have been studied in the present investigation. Assam; DDw 713, Bangia Desh; DDw 6283 Structure of the wood: Growth rings distinct BH/40582, DDw 6625, Burma; DDw 652, in D. grandiflora and distinct to indistinct in D. DDw 2380, DDw 5797 BH/26345, DDw molluccana delimited predominantly by radial- 7320, DDw 7394, DDw 7528, DDw 7532, ly flattened fibres (in 70 % of the samples of DDw 8259, West Bengal. the genus). Wood diffuse-porous. Vessels 2-8 2. D. molluccana Blume: DDwF 189, DDwF (-5)/mm 2, predominantly solitary and also in 8627, Manila, Philippines. radial multiples of 2-3, round, average tangen- 3. Sonneratia alba J. Smith: DDw 6223 BH/ tial diameter medium to large (174-250 Mm), 41145, DDw6236,Andamans;DDw 6167 BH average vessel member length medium (753- 39512 and herbarium specimen no 20672, 780 Mm). Perforations exclusively simple with Burma. oblique to horizontal end walls. Inter-vessel pits 4. S. apetala Buch.-Ham.: DDw 6815, Burma; crowded to sparse, alternate, round to poly- DDw 7300 BH/73387, DDw 7303 .BH/73388, gonal, 10-14 Mm in diameter, vestured type B, DDw 7306 BH/73389, DDw 7362 BH/75835 Form 2, with most of the basal trunks thick and herbarium specimen no 36572, West and branching elaborately into the pit chamber; Bengal. warts surrounding pits, also present on the per- 5. S. caseolaris (L.) Engl. (syn. S. acida L. 0: foration rims. Vessel-ray and vessel-parenchyma DDw 6312 BH/395 1 1, DDw 6193 BH/395 1 1, pits abundant, elongate, gash-like (maximum Burma; DDw 8095, West Bengal. size 30 Mm), sometimes unilaterally compound, 6. S. griffithii Kurz: herbarium specimen no vestured; tyloses frequent to infrequent. Paren 16749, Andamans; DDw 6164 BH/39501, chyma exclusively paratracheal, vasicentric. DDw 6247 BH/39501, DDw 6581 and her- Strands of 2-4 cells, crystals solitary and vari- barium specimen no 2966/1 083, Burma. able in size and shape (Chattaway, 1956) in both the species and rarely also as styloids. For anatomical studies, cross, tangential and Rays 11-1 O/mm, heterogeneous, uni- to triseri- radial sections of 20 Mm were cut and perma- ate, predominantly uniseriate, rarely up to 4- nent slides were prepared following standard seriate composed of erect and square cells. The laboratory techniques. Fibre length and vessel height of the uniseriate rays 5-30 cells (281- member length (including tails) were measured 1120 Mm) and of the multiseriate rays 12-30 from macerations. Other quantitative characters cells (30-1120 Mm). Crystals solitary, rhom- were studied from permanent slides. Forproper boidal to variable in size and shape (Chattaway, observation of crystals, radial sections were 1956) in erect cells rarely as styloids. Rays of- bleached in a sodium hypochlorite solution. ten turn black due to fungal attack. Fibres thin- Perforation plates and vestured pit morphology walled, angular arranged in regular radial files were studied with the help of a Cambridge S600 in cross section, libriform, non-septate with scanning electron microscope (SEM). The type minute pits with reduced borders (max. diam- of perforation described is according to the eter 2.9 Mm), pits only on radial walls. Average terminology used by Gray and De Zeeuw fibre length medium (1291-1526 Mm). Fibre! (1974) and Van Vliet's classification (1978) is vessel member length ratio 1.66-2.03 (average adopted in describing vestured pit morphology. 1.85) (Figs. 1-6). The details of qualitative and quantitative characters are given in Table I. Sonneratia L. f. The genus is represented by five littoral spe- Observations cies of trees, growing in the mangrove swamps of East Africa, Southeast Asia and warmer re- Duabanga Buch.-Ham. gions of Australia. Four species, S. alba, S. ape The genus is represented by 3 species of tala, S. caseolaris and S. griffUhU have been trees growing inland in Indo-Malesian regions. studied in the present investigation. Two species, D. grandiflora and D. molluccana Structure of the wood: Growth rings distinct

Fig. I. Duabanga grandiflora. CS, growth rings delimited by radially flattened fibres. Note vasicentric parenchyma. x 43. - Fig. 2. D. grandiflora. TLS, nonseptate fibres and rays. x 42. - Fig. 3. D. grandiflora. TLS, inter-vessel pits vestured. x 380. - Fig. 4. D. molluccana. TLS, nonseptate fibres, uniseriate rays and intervessel pits. x 190. - Fig. 5. D. grandiflora. RLS, vessel-ray pitting, x 380.- Fig. 6. D. grandiflora. Crystals in rays and parenchyma. Note crystal sand and variable crystals in both. x 380.

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delimited mostly by radially flattened fibres (in Van Vliet and Baas (1984). Furthermore, both 59% of the samples). Wood diffuse-porous. Venkateswarlu and Rao (1964) failed to report Vessels 17-45 (average range 21-37)/mm 2 , the occurrence of crystals in the ray cells of both solitary and in radial multiples of 2-4, Duabanga, a character reported earlier by many mostly oval, average tangential diameter small and also observed constantly in all the samples (88-95 }..lm), average vessel member length reinvestigated. Apart from these, none of the short (650-670 }..lm). Perforations simple, sca- workers who studied this family earlier (Pear- lariform, scalariform branched, reticulate irreg- son & Brown, 1932; Reyes, 1938; Metcalfe & ular, reticulate with or without borders and Chalk, 1950; Venkateswarlu & Rao, 1964; Pur- also as combination perforation plates. Inter- kayastha, 1982; Van Vliet & Baas, 1984) re- vessel pits crowded, alternate, round, oval to ported the occurrence of scalariform and reti- polygonal, 5-9}..lm in diameter, vestured type culate perforations in Sonneratiaceae (Sonnera A, with the vestures attached to all parts of the tia) (Vijendra Rao et aI., 1987, this issue). roof of the pit chamber and branching into a Although Duabanga and Sonneratia are treat- compact mass of vestures of equal thickness. ed in the same family, anatomically there are Vessel-ray pits round, oval, more or less gash- more quantitative and qualitative dissimilarities like (maximum diameter 19-20 }..lm), vestured; than similarities between these two genera ty loses and gummy deposits frequent. Paren (Table I). From the observations made, Dua chyma absent. Rays 17-20/mm, homogeneous, banga and Sonneratia show the following simi- predominantly uniseriate and occasionally to larities: diffuse porosity, radially flattened fibres rarely, partly or completely biseriate, composed delimiting growth rings, vestured intervessel pits of mostly procumbent cells, sometimes with in- and minute fibre pits only on the radial walls. tervening upright cells. The height of the rays The dissimilarities between these two genera 19-35 cells (496-800 }..lm). Crystals abundant, are: low and high frequency of vessels, percen- solitary, in cells forming horizontal series. Per- tage grouping of solitary vessels and radial mul- forated ray cells occasionally present in S. grit tiples, tangential diameter of vessels, size of in- fithii. Fibres septate, 2-3 septa per fibre, thin- tervascular pitting, size and shape of vessel-ray walled, arranged in regular radial files in cross pitting, presence and absence of parenchyma, section. Libriform fibres with minute pits hav- uni- to multiseriate heterogeneous rays to al- ing reduced borders (maximum diameter 2.60 most exclusively uniseriate homogeneous rays, }..lm) only on radial walls. Average fibre length variable type of crystals and normal type of medium (1154-1204 }..lm). Fibre/vessel mem- crystals in the rays, absence and presence of ber length ratio 1.77-1.81 (average 1.79) septa in the fibres. Furthermore, there are dif- (Figs. 7-13). ferences in vessel perforation types and morphology of vestured pits between these two Discussion genera. While perforations are exclusively sim- Van Vliet and Baas (1984), who summarised ple in Duabanga, they show a combination of the range of anatomical variation in the struc- simple-scalariform, simple-reticulate and simple ture of Sonneratiaceae, found that the data ob- in Sonneratia. Similarly Duabanga shows type tained by Venkateswarlu and Rao (1964) devi- B form 2 of vestured pit morphology while ate from what they have obtained. Venkate- Sonneratia shows type A (Van Vliet & Baas, swarlu and Rao (1964) studied Duabanga son 1984 and also confirmed in the present study). neratioides and Sonneratia apetala and S. acida From the above it is evident that the differ- in samples obtained from the Forest Research ences between these two genera are real and Institute Xylarium, Dehra Dun. Unfortunately features like absence and presence of paren- they have not quoted the sample numbers. The chyma and the type of vestured pit morphol- data obtained in the present investigation from ogy are to some extent taxonomically impor- the samples available at the F.R.I. Xylarium, tant. The rudimentary presence of scalariform Dehra Dun also confirm the views expressed by and reticulate perforation plates in Sonneratia

Fig. 7. Sonneratia caseolaris. CS, growth rings delimited by radially flattened fibres, vessels in radial multiples. x 43. - Fig. 8. S. caseolaris. TLS, septate fibres, uniseriate rays, intervessel pits. x 42. - Fig. 9. S. apetala. TLS, intervessel pits. x 380. - Fig. 10. S. ape tala. RLS, crystals in horizontal series in rays. x 90. -'- Fig. II. S. caseolaris. RLS, three vessel members showing simple perforations at the bottom and simple, scalariform, and reticulate perforations at the top. x 130. - Fig. 12. S. caseolaris. RLS, reticulate perforation plate and vessel-ray pitting. x 130. - Fig. 13. S. griffithii. RLS, perforated ray cell with simple perforation. x 130.

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Table I. Quantitative and qualitative anatomical characters of Duabanga and Sonneratia.

Characters Duabanga Sonneratia grandiflora molluccana alba apetala caseolaris griffithii

1. Growth rings delimited by radially flattened fibres + + + + + 2. Diffuse porosi ty + + + + + + 3. Vessels a. Average number/vessels/sq.mm 5 5 37 21 21 35 b. Frequency of singles and radial multiples (%) 62-38 55-45 38-62 44-56 50-50 30-70 c. Average tangential diameter (urn) 174 258 95 94 95 88 d. Maximum size of intervascular pitting (J..lm) 13 14 7 7 9 8 e. Type of vestured pit morphology B (2) B (2) A A A A f. Perforation type: Simple (S), Scalarifonn (SC), Reticulate (R) S S S, SC, R S, SC, R S, SC, R S, SC, R g. Average vessel member length (J..lm) 780 753 650 662 650 670 h. Inclusions: Tyloses (T), Gummy deposits (G) T T T,G T,G T,G T,G 4. Parenchyma + + a. Vasicentric type + + b. Inclusions: Crystals solitary (SO), Variable type in size and shape (V) SO,V SO,V 5. Rays a. Homocellular + + + + b. Heterocellular + + c. Number/rays/mm II 10 18 17 20 19 d. Maximum height of uniseriates (cells) 30 25 29 39 29 31 e. Maximum height of multiseriates (cells) 30 rare f. Width of multiseriates (cells) 2-4 2-3 g. Ray-vessel pitting size (J..Lm) 30 30 19 19 20 18 h. Type of ray-vessel pitting: elon- gated gash-like (G+), round (R), oval (0), elliptical more or less gash-like (G), unilateral compound pitting (U) G+, U R,O,G R,O,G R,O,G R,O,G R,O,G i. Inclusions: Crystals solitary (SO), Variable in size and shape (V) SO,V SO,V SO SO SO SO 6. Fibres a. Nonseptate + + b. Septate + + + + c. Wall thickness (J..lm) 2-3 2-3 3-4 4-5 3-5 3-4 d. Average length (J..lm) 1291 1526 1177 1172 1154 1204 7. Fibre-vessel member length ratio 1.66 2.03 1.81 1.77 1.77 1.80 8. Vulnerability value 34.80 51.60 2.57 4.48 4.52 2.51 9. Mesomorphy value 27144 38855 1669 2966 2940 1685

Downloaded from Brill.com09/26/2021 10:35:44AM via free access IAWA Bulletin n.s., Vol. 8 (4),1987 343 is only remnant of a primitive feature. This Similarly, Sonneratia and Duabanga show some observation further strengthens the view ex- dissimilarities with Lythraceae. However, the pressed by Van Vliet and Baas (1984) that overall wood anatomical variation is well with- Sonneratia has retained many protomyrtalean in the range of Lythraceae. characters. Baas and Zweypfenning (1979), On the basis of information from different Baas (1986) and Graham et al. (1986) who subdisciplines of botany, Dahlgren and Thorne studied the wood anatomy of Lythraceae, re- (1984) showed differences between Duabanga ported the occurrence of only simple perfora- and Sonneratia on the one hand and Lythraceae tion plates in all the members except Cuphea on the other. While the differences between carthagensis which showed only once vestigial these two genera are distinct, some of the simi- bars in the first formed secondary xylem larities which have been used to justify the (Baas & Zweypfenning, 1979). This observation previous family status are likely to be due to prompted us to check all the slides of Lythra- convergence according to these authors. The ceae available to us. Interestingly we could question arises whether these two genera can find in small late wood vessels of Woodfordia be treated together in the same family. Met- fruticosa Kurz (sample nos. DDw 876andDDw calfe & Chalk (1950) also questioned the re- 4813) perforations with irregular contours or tention of these two genera in the same family. impressions of the foraminate type. In a few Purely from the point of view of wood anat- vessels, scalariform perforations with a single omy, enough anatomical differences exist be- bar have also been seen. The first two types tween these two genera to advocate their treat- mentioned above have also been reported by ment in different families. These differences Schmid and Baas (1984) in Myrtaceae. Perhaps are, however, within the range of Lythraceae. this observation on Woodfordia indicates that Thus the study indicates that these two genera protolythraceous members also might have had cannot be treated together as a family as sug- scalariform and foraminate or reticulate perfo- gested by many or together as a subfamily of rations and thus also qualify for a common an- Lythraceae as suggested by Venkateswarlu and cestry for Lythraceae and Sonneratia. Rao (1964) but should be given subfamily sta- In the introduction it has been shown that tus, Duabangoideae and Sonneratioideae under the systematic status accorded by different the Lythraceae as advocated by Dahlgren and authors to Sonneratia and Duabanga (Sonnera- Thorne (1984). However, such an inclusion, as tiaceae) are varied. Recently Dahlgren and pointed out by Baas and Zweypfenning (1979), Thorne (1984) suggested that these two genera would add somewhat to the heterogeneity of may be given independent subfamily status the family Lythraceae. within Lythraceae. The present study supports the above view and the arguments in favour of Ecological considerations giving such a status is based on the differences An attempt has also been made in the present shown earlier between these two genera. study to correlate the wood anatomical charac- It was Thorne (1976) who reopened the de- ters with ecological factors. Carlquist (1975b) bate of family delimitation of Lythraceae by attempted to analyse factors underlying the incorporating Sonneratiaceae (Sonneratia and selective forces which have guided evolution of Duabanga), Punicaceae (Punica), and Alzatea conductive tissue in vascular plants. According (Crypteroniaceae) in it. Van Vliet (1975), who to him (Carlquist, 1977) short, narrow vessel worked on the wood anatomy of Crypteronia- elements resist high tensions in water columns. ceae sensu lato. showed that Alzatea has struc- To some extent there is an inverse correlation tural similarities with Sonneratia and Lythra- between diameter of vessels and the number of ceae and suggested the inclusion of the former vessels per square millimetre. By dividing mean and Sonneratiaceae in Lythraceae. A view sub- vessel diameter by number of vessels per square sequently endorsed by Baas (1979) and Baas millimetre of transection one gets a value and Zweypfenning (1979). Similarly Bridgwater which may vary. A low value for this ratio indi- and Baas (1979) endorsed the views of Thorne cates a great redundancy of vessels. According (1976) for keeping Punica in Lythraceae. As to him the more numerous the vessels per square regards Sonneratiaceae, Baas and Zweypfenning millimetre the less the chance that disabling of (1979) are of the opinion that inclusion of the a given number of vessels by air embolisms same in Lythraceae would add somewhat to formed under water stress would seriously im- the heterogeneity of the latter from the wood pair water conduction in a plant. A low value anatomical point of view. The present work for this ratio would therefore indicate a capa- also supports the view of Baas (1979) on the city for withstanding water stress or freezing. structural similarity between A lzatea and Son Carlquist termed this ratio as 'vulnerability' neratia although there are some differences. and this ratio when multiplied by mean vessel

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member length gives values which he termed ~ 1977. Ecological factors in wood evolution: 'mesomorphy'. Species showing higher values A floristic approach. Amer. J. Bot. 64: indicate the mesomorphic nature of the plant. 887-896. On the basis of Carlquist's indices the vulner- ~ & L. DeBuhr. 1977. Wood anatomy of Pe- ability and mesomorphy values for both these naeaceae (Myrtales): comparative, phylo- genera were calculated. Sonneratia occurs in genetic and ecological implications. Bot. J. mangrove habitats and is adapted to saline Linn. Soc. 75: 221-227 hence physiologically dry conditions. Accord- Chattaway, M.M. 1956. Crystals in woody tis- ingly it shows smaller vessels with higher fre- sues. II. Trop. Woods 104: 100-124. quencies. However, in the case of Sonneratia Cronquist, A. 1968. The evolution and classifi- ape tala and S. caseolaris the frequencies are cation of flowering plants. Houghton Mifflin slightly outside the range given for mangrove Co., Boston. plants by Panshin (1932). Duabanga growing Dahlgren, R. 1975. A system of classification inland has larger vessels in lower frequencies. of the angiosperms to be used to demon- In keeping with its mangrove habit, Sonneratia strate the distribution of characters. Bot. has a very low vulnerability value (2.51-4.52, Notis. 128: 119-147. average 3.52) when compared to Duabanga ~ 1980. A revised system of classification of which showed a value of 43.20 (range 34.80- the angiosperms. Bot. J. Linn. Soc. 80: 91- 51.60). The low values obtained for Sonneratia 124. are indicative of its capacity to resist water ~ & R. F. Thorne. 1984. The order Myrtales: stress. Similarly, the mesomorphy values ob- circumscription, variation and relationships. tained for Duabanga (32,999) is indicative of a Ann. Missouri Bot. Gard. 71: 633-699. mesic ecology of the species unlike Sonneratia Emberger, L. 1960. Les vep~taux vasculaires. which has a very low value (2,315). Vol. 2 (in 2 fascicles). In: Trait6 de Botani- que (Systematique) (eds. M. Chaudefaud & L. Emberger). Masson & Cie, Paris. Acknowledgements Graham, S.A., H. Tobe & P. Baas. 1986. Koehne- Thanks are due to Mr. Magan Singh for the ria, a new genus of Lythraceae from Mada- technical work and Mr. Prasant Sharma for gascar. Ann. Missouri Bot. Gard. 73: 788- photographic work. 809. Gray, R.L. & C.H. de Zeeuw. 1974. Terminol- ogy for multiperforate plates in vessel ele- References ments. IAWA Bull. 1974/2: 22-27. Airy Shaw, H.K. 1966. A dictionary of the Hutchinson, J. 1973. The families of flowering flowering plants and ferns. (Revised 6th plants. 3rd Ed. Clarendon Press, Oxford. edition by J.C. Willis). Cambridge Univ. Koek-Noorman, J., P. Hogeweg, W.M.H. van Press, Cambridge. Maanen & B.J.H. ter Welle. 1979. Wood Baas, P. 1979. The anatomy of Alzatea Ruiz & anatomy of the Blakeeae (Melastomata- Pav. (Myrtales). Acta Bot. Neerl. 28: 156- ceae). Acta Bot. Neerl. 28: 21-44. 158. Melchior, H. 1964. Myrtiflorae. In: Engler's Syl- ~ 1986. Wood anatomy of Lythraceae - addi- labus der Pflanzenfamilien (ed. H. Melchior), tional genera (Capuronia, Galpinia, Haitia, 12th Ed. Vol. 2: 345-366. Gebr. Borntrae- Orias, and Pleurophora). Ann. Missouri Bot. ger, Berlin, Nikolassee. Gard. 73: 810-819. Metcalfe, C.R. & L. Chalk. 1950. Anatomy of ~ & R. C. V. J. Zweypfenning. 1979. Wood the dicotyledons. II. Clarendon Press, Ox- anatomy of the Lythraceae. Acta Bot. Neerl. ford. 28:117-155. Mujica, M.B. & D.F. Cutler. 1974. Taxonomic Bentham, G. & J.D. Hooker. 1862. Genera plan- implications of anatomical studies on the tarum. Reeve & Co., London. Oliniaceae. Kew Bull. 29: 93-123. Bridgwater, S. D. & P. Baas. 1978. Wood anat- Panshin, A. J. 1932. An anatomical study of the omy of the Punicaceae. IAWA Bull. 1978/1: woods of the Philippine mangrove swamps. 3-6. Philipp. J. Sci. 48: 143-207. Carlquist, S. 1975a. Wood anatomy of Onagra- Pearson, R. S. & H.P. Brown. 1932. Commer- ceae with notes on alternative modes of cial timbers of India. II. Central Publication photosynthate movement in dicotyledon Branch, Calcutta. woods. Ann. Missouri Bot. Gard. 62: 386- Purkayastha, S.K. 1982. Indian Woods. IV. Con- 424. troller of Publications, Delhi. ~ 1975b. Ecological strategies of xylem evo- Reyes, L. J. 1938. Philippine woods. Tech. Bull. lution. Univ. California Press, Berkeley. 7, Dept. Agric. Philipp. Inst. Manila.

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Schmid, R. & P. Baas. 1984. The occurrence of Venkateswarlu, J. & P.S. Prakasa Rao. 1964. scalariform perforation plates and helical The wood anatomy and taxonomic posi- vessel wall thickenings in wood of Myrta- tion of Sonneratiaceae. Curr. Sci. 3: 6-9. ceae. IAWA Bull. n.s. 5: 197-216. Vijendra Rao, R., S. S. Bisen, Babulal Sharma & Soo', C. R. 1967. Die modernen Systeme der R. Dayal. 1987. SEM observations of perfo- Angiospermen. Acta Bot. Acad. Sci. Hung. ration plates in Sonneratia Linn. (Sonnera- 13: 209-233. tiaceae). IAWA Bull. n.s. 8: 331-336. - 1975. A review of the new classification Vliet, G.J.C.M. van.1975.Comparativeanatomy systems of flowering plants (Angiosperma- of the Crypteroniaceae sensu lato. Blumea tophyta, Magnoliophyta). Taxon 24: 585- 22: 173-195. 592. - 1976. Wood anatomy of the Rhizophora- Takhtajan, A. L. 1969. Flowering plants: Origin ceae. Leiden Bot. Series 3: 20-75. and dispersal. (Transl. from Russian by C. - 1978. Vestured pits of Combretaceae and Jeffrey.) Smithson. Inst. Press, Washington, allied families. Acta Bot. Neerl. 27: 273- D.C. 285. Thorne, R. F. 1968. Synopsis of a putative - 1979. Wood anatomy of the Combretaceae. phylogenetic classification of the flowering Blumea 25: 141-223. plants. Aliso 6: 57-66. - 1981. Wood anatomy of the paleo tropical - 1976. A phylogenetic classification of the Melastomataceae. Blumea 27: 395-462. Angiospermae. Evol. BioI. 9: 35-106. - & P. Baas. 1984. Wood anatomy and clas- - 1981. Phytochemistry and angiosperm phy- sification of the Myrtales. Ann. Missouri logeny, a summary statement. In: Phyto- Bot. Gard. 71: 783-800. chemistry and angiosperm phylogeny (eds. Welle, B.J.H. ter & J. Koek-Noorman. 1981. D. Young & D.S. Seigler): 233-295. Prae- Wood anatomy of the Neotropical Melas- ger, New York. tomataceae. Blumea 27: 335-394.

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