BLUMEA 32 (1987) 277-302 A SYSTEMATIC REVISION OF THE MANGROVE GENUS SONNERATIA (SONNERATIACEAE) IN AUSTRALASIA* N.C. DUKE** & Betsy R. JACKES *** SUMMARY In Australia, New Guinea and the southwestern Pacific seven taxa are recognised in Sonneratia of which three species are redescribed in view of their Indo-Malesian counterparts (S. alba, S. caseolaris, S. ovata), one species is redescribed from its most likelyIndo-Malesian equivalent (S. lanceolata, being distinct from S.caseolaris), one widespread hybrid is fully described (S. x gulngai being the putative synonym of S. alba x S. caseolaris), and two other hybrids (S. alba x S. gulngai, S. alba x S. lanceolata ) of very limited occurrence are described as sub-units of their closest ‘parental’ forms. A key, descriptions and full synonymy are given as well as 2 distribution maps, 6 other figures and one table. TAXONOMIC STATUS AND DISTRIBUTION Backer & Van Steenis (1951) compiled a thorough review of the Sonneratiaceae, a family of the order Myrtales. Two genera were described and include Duabanga, a small evergreen rainforest genus, and Sonneratia. The latter was made up of five mangrove tree species: S. alba, S. caseolaris, S. ovata, S. apetala , and S. griffithii. Their distributions were described as tropical and ranging through the Indo-West Pacific region. However, the account was not complete and new observations were reported in various appendices up to 1972. Some observations included those of additional characters, e.g., leaf mucronate tips or stamen colour; but most importantly there was the possibility of undescribed taxa. These included putative hybrids discovered in NW. Borneo (Muller & Hou-Liu, 1966). In Australia, only one taxon (S. alba) was initially reported (Backer & Van Steenis, 1951; Jones, 1971), although there was some evidence that suggested the presence of others, including hybrids (Van Steenis, 1968; Muller & Van Steenis, 1968). Additional taxa were later discovered in extensive field surveys across northern Australia (Wells, 1982; Bunt et al., 1982). However, problems were encountered in classifying them using keys of Backer & Van Steenis (1951). These problems (Bunt et al., 1982; Duke et al., 1984) included:1) * Australian Institute of Marine Science Contribution No. 329. ** Australian Institute of Marine Science P. M. B. 3, M. C. Townsville, Queensland 4810, Australia. *** James Cook University of North Queensland, Townsville, Queensland 4811, Australia. 278 BLUMEA - VOL. 32, No. 2, 1987 presence or absence of petals used to disting S. alba and S. ovata; 2) the occurrence of two forms o S.caseolaris; and 3) the possibility of a hybrid between one of the S.caseolaris forms and S. alba (or S . ovata?). Therefore classification of Sonneratia species in the study region (described as Australia, New Guinea and the southwestern Pacific) was incomplete, and no taxa were adequately defined. The most widespread species S . alba (fig. 1), was considered always to have Fig. 1. Distribution of S o n n e r a t i a in Australia, New Guinea and the southwestern Pacific region: (1) S. alba J.Smith (cross), S. o v a t a Backer (circle), and S. alba x S. gulngui (square); (2) S. caseolaris (L.) Engl.(cross), S. x gulngai N.C. Duke (circle), S. lanceolata Blume (triangle) and S. alba x S. lanceolata (square). N.C. Duke & B.R. Jackes: Revision of Sonneratia in Australasia . . 279 petals, although these may have been hard to discern (Backer & Van Steenis, 1951). Exhaustive searches of both fresh and herbarium material from the region have revealed that the petalous condition of S. alba is variable. It is now known to occur in three forms: petalous, apetalous, and semipetalous (the latter term applies to those forms where petal numbers are less than the number of calyx lobes, commonly six). These three forms were also observed to have a disjunct distribution. Apetalous varieties predominate at the southern limits of the species on the east and west coasts of Australia. In New Guinea, there is no discernible geographical pattern. However, ecologically, apetalous forms predominate in small offshore island populations, and petalled forms are chiefly estuarine throughout their range. The use of petal presence or absence by Backer & Van Steenis (1951) to distinguish between apetalous S. alba and the less common, normally apetalous, S. ovata many other charcreated problems. Although both species possess white acters appeared to be quite distinctive. Characters unique to S. ovata include the lack of a mucronate leaf apex, a finely verruculose calyx, and its common occurrence in higher topographic (tidal) positions near the terrestrial fringe. The known distribution of S. ovata (fig. 1) was extended, and it occurs along the southern Papua New Guinea coastline to Milne Bay (10º23'S,150º3l'E). The species was not recorded in Australia, but it was located on Daru Island (9º05'S, 143’ 10'E) in the northern Torres Strait. Van Steenis (1968) indicated that he expected a greater variety of forms of Sonneratia in Australia and New Guinea, including some of the putative hybrids observed in NW. Borneo (Muller & Van Steenis, 1968). Since then three other major taxa have been discovered. These five taxa and their diagnostic characters are presented in table 1. Sonneratia lanceolata and S. caseolaris have never been observed to co-inhabit the same estuary, although their geographical ranges overlap. Sonneratia caseolaris is only found on the northeast coast of Australia (fig. 1), but S. 1anceolata also occurs in the Northern Territory (although it is limited to three separate areas). In New Guinea the distribution of these two taxa is also disjunct, with S. lanceolata occupying most larger estuaries on the mainland south coast, and S. caseolaris in similar habitats on the north coast. Features which characterise populations of the hybrid, S. x gulngai (Duke, 1984) include morphological attributes which are either intermediate or alternately shared between S. alba and S, caseolaris. These characters include: high levels of pollen infertility and poor fruit set; a floral development cycle which is complex; generally luxuriant foliage and greater tree size. This latter factor allows this common hybrid form to be easily recognised in the field. The regional distribution of the hybrid and the least common parent, S. caseolaris, are equivalent in Australia. Both taxa are found on the east coast from the southern limit, Murray River (18º05'S, 146º0l'E), north to the Olive River (12ºl0'S, 143º05'E). It is anticipated that in New Guinea additional populations will be located where the parental ranges also overlap. This revision is based on both herbarium material and field observations. Contributing herbaria (abbreviations from Holmgren & Keuken, 1974): AIMS, Herbarium of the Australian Institute of Marine Science, TownsvilIe; BRI, Queensland Herba- Table 1. Diagnostic characters of major Sonneratia in Australia, New Guinea and the southwestern Pacific region. S. alba S. caseolaris S. x gulngai S. lanceolata S. ovata Leaves elliptic elliptic elliptic elliptic lanceolate broadly ovate Leaf apices rounded, mucro fold apiculate, mucronate apiculate, mucronate apiculate, mucronate rounded, mucro absent Leaf base attenuate oblique attenuate oblique attenuate obIique attenuate oblique reniform Peduncle/branchlets terete terete or tetragonous terete terete or tetragonous terete Petals white, linear-spathulate, red, linear often absent red, linear red, linear, rarely double absent Stamen white red, rarely white ted white white calyx smooth, shiny coriaceous-warty, shiny smooth, dull shiny coriaceous-smooth, dull smooth, dull coriaceous Fruit calyx (hypanthium) cup-shapcd flat-expanded cup-shaped flat-expanded flat-expanded Fruit width = corolla width width 5 mm > corolla width width = corolla width width 5 mm > corolla width width 5 mm > corolla width Seeds falcate angular irregular angular irregular angular irregular rounded irregular Inflorescence 1-5 (2) 1-3 (1 or 2) 1-3 (1 or 2) 1-2 (1) 1-3 (2) triporate 50-80 µmP 40-50 µmP variable 40-60 µmP 30-40 µmP variable 60-100 µmP 40-65 µmP triporate 40-65 µmP 30-55 µ m P Pollen shape size N.C. Duke & B.R. Jackes: Revision of Sonneratia in Australasia 281 rium, Brisbane; DNA, Herbarium of the Conservation Commission of the Northern Territory, Darwin; LAE, Division of Botany, Department of Forests, Lae; PERTH, Western Australian Herbarium, Department of Agriculture, Perth; QRS, Queensland Research Station, Forest Research Institute, Atherton; and UPNG, Herbarium of the University of Papua New Guinea, Port Moresby. A BRIEF EVALUATION OF DIAGNOSTIC CHARACTERS* . Habitat - Most species of this region may be distinguished by the habitat in which they commonly occur. Further details shall be reported by Duke (a; in preparation). Only two species, S. lanceolata and S. caseolaris, were found in the same, or a similar, habitat. Habit - Tree form and foliage colour may be used to distinguish the species. However, the differences are subtle but they are useful to the experienced field observer. Pneumatophores - Length, diameter and shape are useful characters in the field. Leaves - Subtle differences of surface texture, colour and shape may be used to distinguish most species. However, only S. Ianceolata (with lanceolate leaves) and S. ovata (with broadly ovate leaves and no apical mucro) may be readily separated. Petiofe - Relative length, and degrees of lamina extensions may be used to categorise subgroupings or support other diagnostic characters. Inflorescence - The numbers of buds per inflorescence vary but are not generally considered diagnostic. Hypanthium - Surface texture of the calyx tube is diagnostic although populations of some species vary slightly. The degree of red coloration is variable on the inner calyx lobe surfaces. Corolla - Presence or absence of petals is of little assistance in distinguishing the species because this character is variable in S. alba. Petal colour may be useful in separating subgroups. Stamens - Colour of the filaments is generally consistent and useful in separating subgroups of the genus, unlike length and thickness. Ovary - Structure of the ovary generally is not diagnostic. Subtle differences are observed in the position of placental material on the septa. The number of locules is variable. Pollen - Muller (1969, 1978) and Wright (1977) reported some differences in shape, size and ornamentation. Cytology - Muller & Hou-Liu (1966) reported chromosome counts of n = 11 for the major Indo-Malesian taxa and associated hybrids. In Australasia, one of us (Duke) observed counts of n = 12 in the four major taxa described (1-4). Methods used were similar to those described earlier by Muller & Hou-Liu (1966), except that acid orcein stain was found more suitable. These latter results concur with the findings of Sidhu (1962) for S. apetafa, and Weiss (1973) for S. alba. * Unless otherwise stated, all characters are more fully described in the specific descriptions. 282 BLUMEA - VOL. 32, No. 2, 1987 Fruit - Characteristics of the mature fruit are very important in distinguishing between the species. Some characters enable generic subgroups to be delimited, others may be used to confirm species identification. Diagnostic characters includecalyx tube shape, the relative widths of fruit and hypanthium, and surface texture and colour. Seeds - Shape and size of seeds are generally not useful as diagnostic tools, but one species, S. alba, is readily distinguished from the others. MORPHOMETRIC ANALYSES Classification of taxa in this revision was supported by morphometric analyses. Full details of these studies are to be reported by Duke (b, in preparation). In summary, 283 specimens from 43 Australasian sites were examined. Collections were obtained both fresh and from various herbaria including LAE, DNA, QRS, BRI, and AIMS. Observations on collections covered up to 42 numeric and 18 multistate attributes of a wide range of vegetative and reproductive part morphological characters including those commonly used in earlier specific descriptions. The study showed S. alba as having a central generalised affinity with five of the six other taxa considered (fig. 2). Sonneratia lanceolata was notably seen as the taxon with the last affinities with S. alba. It was also quite separate from S. caseolaris. Putative hybrids are clearly portrayed in their intermediate positions. Thus overall in the scheme there are basically three wings branching from the central axis of S. alba and S. caseolaris. Intraspecific results demonstrate the possible ecotypic expression of the petalous condition of S. alba. This species is found to be apetalous chiefly in marine offshore situations and in higher latitudes. Other characters are less variabIe and this species along with S. caseolaris, S. x gulngai and 2. Schematic diagram of Australasian S. ovata are seen to be generally morpho- Fig. Sonneratia taxa and their inter-specific affilogically homogeneous. In contrast, S. lan- nities deduced from morphometric analyses. ceoIata has well defined infraspecific Choice of circle size and line Iength is arbigroups based on geographic occurrence. trary. ACKNOWLEDGEMENTS This revision is part of a larger study of the genus Sonneratia in Australasia. The study was undertaken part-time towards a thesis with the James Cook University of North Queensland whilst the senior author was employed by the Australian Institute of Marine Science. We wish to note our gratitude to both these institutions and Dr. John Bunt (A.I.M.S.). We also wish to thank the directors and curators of the herbaria listed for making specimens in their care avaiIable to us. N.C. Duke & B.R. Jackes: Revision of Sonneratia in Australasia 283 REFERENCES BACKER, C.A. & C.G.G.J. VAN STEENIS. 1951(-1972). Sonneratiaceae. Flora Malesiana I, 4: 280289. Addenda in Flora Malesiana I, 5 (1958): 557 and I, 6 (1972): 973. BUNT, J.S., N.C. DUKE & W.T. WILLIAMS 1982. Mangrove distributions in north-east Australia. J. Biogeography 9: 111-120. D UKE, N.C. 1984. A mangrove hybrid, Sonneratia x gulngai (Sonneratiaceae), from north east Australia. Austrobaileya 2: 103-105. - (a) Localised and regional distribution patterns of the mangrove genus Sonneratia in Australasia. In preparation. - (b) Morphometric analysis of the mangrove genus Sonneratia in Australasia. In preparation. - , J.S. BUNT & W.T. WILLIAMS. 1984. Observations on the floral and vegetative phenologies of north-eastern Australian mangroves. Aust. J. Bot. 32: 87-99. HOLMGREN, P.K. & W. KEUKEN. 1974. Index Herbariorum, Part I, ed. 6, incl. addn. in Herbaria of the World, ed. 6, III, 1978, Taxon 27: 427-432. JONES, W.T. 1971. Field identification and distribution of mangroves in eastern Australia. Qld. Naturalist 20: 35-51. MULLER, J. 1969. A palynological study of the genus Sonneratia (Sonneratiaceae). Pollen et Spores 11: 223-298. 1978. New observations on pollen morphology and fossil distribution of the genus Sonneratia - (S onneratiaceae). Rev. Palaeobot. Palynol. 26: 277-300. - & S. Y. HOU-LIU. 1966. Hybrids and chromosomes in the genus Sonneratia (Sonneratiaceae). Blumea 14: 337-343. - & C.G.G.J. VAN STEENIS. 1968. The genus Sonneratia in Australia, with notes on hybridization of its two species. N. Qld. Naturalist 35: 6-8. SIDHU, S.S. 1962. ChromosomaI studies on some mangrove species. Indian Forester 88: 585-592. STEENIS, C.G.G.J. VAN. 1968. Do Sonneratia caseolaris and S. ovata occur in Queensland or the Northern Territory? N. Qld. Naturalist 35: 3-6. WEISS, H. 1973. Contribution to cytotaxonomy and karyology of mangroves from Madagascar and world (French). Rev. Gen. Bot. 80: 209-240. WELLS, A.G. 1982. Mangrove vegetation of Northern Australia. In: B.F. Clough (ed.) Mangrove ecosystems in Australia: Structure, function and management: 57-78. ANU Press, Canberra. WRIGHT, D.F. 1977. A North Queensland mangrove pollen flora. Hons. thesis. JCUNQ, Aust. 98 pp., 27 plates. SONNERATIA Sonneratia L. f., Suppl. Pl. (1781) 38, nom. cons.; J. Smith in Rees,Cycl. 33, 2 (1819); Blanco, Fl. Filip. (1837) 423; ed. 2 (1845) 296; Blume, Mus. Bot. Lugd.-Bat. 1 (1851) 336; Miq., Fl. Ned. Ind. 1, 1 (1855) 495; Benth., Fl. Austral. 3 (1866) 301; Brongn. & Gris, Ann. Sci. Nat. Bot. 6 (1866) 266; Bull.Soc. Bot. France 13 (1866) 479; Blanco,F l Filip. ed. 3 (1878) 186; Engl. in E. & P., Nat. Pfl. Fam. Nachtr. 1 (1897) 261; Merr., Fl. Manila (1912) 343; Gagnep., Bull. Soc. Bot. France 73 (1916) 1.53; Gagnep. & Guillaumin, Fl. Gén. Indo- Chine 2 (1921) 978; Ridley, Fl. Malay Penins. 1 (1922) 825; Hutch., Fam. Flow. Pl. (1926) 145; Watson, Malayan For. Rec. 6 (1928) 118; Burkill, Dict. 2 (1935) 2051; Corner, Wayside Trees (1940) 431; Steenis, F l Scholen Indonesië (1949) 291; Backer & Steenis, Fl. Males. I, 4 (1951) 280; Wyatt-Smith, Malayan For. 16 (1953) 213; H. Perrier, Fl. Madag. (1954) fam. 148, 1; WyattSmith, Malayan For. 23 (1960) 126; Backer & Bakh. f., Fl. Java 1 (1963) 257; Vu VanCuong, Fl. Cambodia Laos Vietnam 4 (196.5) 194, pl. 1; G. Williams Sangai, Fl. Trop. E. Africa (1968) Sonn., 1, f. 1; Whitm., Malayan For. Rec. 26 (1972) 444; Chai, Malaysian For. 38 (1975) 202; Percival & Womersley, Bot. Bull. Dept. For., Papua New Guinea 8 (1975) 82,85; 284 BLUMEA - VOL. 32, No. 2, 1987 Morley, Flow. Pl. (1978) 156; Ko, Acta Phytotax. Sin. 23 (1985) 311.- Blatti Adans., Fam. Pl. 2 (1763) 88, 526; Kuntze, Rev. Gen. Pl. 1 (1891) 238; Niedenzu in E. & P., Nat. Pfl. Fam. 3, 7 (1892) 20. - Aubletia Gaertn., De Fruct. Sem. Pl. (1788) 379. - Kambala Raf., Sylva Tellur. (1838) 19. - Mycostylis Raf., l.c., nom. altem. -Type species: ,Sonneratia acida L. f. [= ‘Pagapate’ Sonn.] Rhizophora auct. non L., Gen.Pl. ed.5 (1754) 202;Herb.Amb. (l754) 13. - T y p e s p e c i e s Rhizophora caseolaris L. [= ‘Mangium caseolare (rubrum)’ Rumph.] Chiratia Montr., Mém. Acad. Sci. Lyon, Sect. Sci. 10 (1860) 202; Brongn.& Gris, Ann. Sci. Nat. Bot. 1 (1864) 362; Bull. Soc. Bot. France 11 (1864) 69. - Type species: Chiratia leucantha Montr. [= ‘Tombea’ Brongn.] Tree medium, columnar to spreading, c. 20 m, trunk not buttressed bark flaky, grey pale brown; roots radiating, horizontal, subsurface. Pneumatophores numerous, vertical, stout, elongate, cone-shaped, often branching, soft flaky surface, soft spongy light wood, length 0.2-2.5 m above substrate. Leaves simple, opposite, entire, smooth, glabrous, leathery; apex mucronate, either extended (minutely) and thickened, or obscure, or rarely non-existent. Stipules absent. Flowers bisexual, termina1, either singly or in groups of 2 or 3, 4-8-merous; calyx tube of mature bud obconical or cup-shaped, lobes ovate-oblong-triangular, surface green, leathery, sometimes coloured red on the inside. Disc saucer-shaped. Petals when present red, white, or white with slight green or red coloration near base, filamentous, spathulate, thinly lanceolate or narrowly linear, early caducous. Stamens numerous (approximately 300), inflexed in closed bud, straight in open flower, early caducous; filaments filiform, red or white; anthers reniform, pale yellow. Ovary superior, sessile, depressed, 10-20 locules. Style folded twice, contracted in mature bud, straight and extended in open flower; stigma entire, capitate. Fruit, when mature, rests on the persistent calyx as an indehiscent, green, smooth, depressed-globose berry with leathery pericarp, crowned by style base; style sometimes persistent but withered. Seeds c. 5 mm Iong, numerous, embedded in firm pulp, released after fruit has fallen from the tree. D i s t r i b u t i o n . Seven species, extending from East Africa to Indo-Malesia, Australia, New Guinea and the western Pacific. Ec olo g y . Found throughout northern Australia, New Guinea, and the southwestern Pacific region in mangrove swamps, on the banks of tidal rivers and creeks, and within sheltered bays of offshore islands and reef cays along the Great Barrier Reef. Sonneratia species along with Avicennia are usually regarded as pioneers of the mangrove swamp. The seeds are intolerant of shade, germinating on bare or near bare mud banks. Consequently their occurrence is usually at the waters edge of the mangrove forest (frontal). There is one exception, S. ovata, which is an uncommon species not yet observed in great detail, but noted to occur closer to the terrestrial margin. Notes. Several vernacular names appeared in prior revisions. ‘Pagapate’ was the’vernacular term used by Sonnerat, Voy. Nouv. Guinée (1776) 16, t. 10 & 11, to describe what is now called Sonneratia caseolaris. Sonnerat did not present an acceptable name. Linnaeus fil. (1781: l.c.) recognised the value of the description and figure, and named the taxon Sonneratia acida. However, the problem was compounded because the species had already been named Rhizophora caseolaris by Lin- N.C. Duke & B.R. Jackes: Revision of Sonneratia in Australasia 285 naeus (1754: 1. c.). Additional problems (discussed later for respective species) arose because this single epithet was based on the dual description ‘Mangium caseolare album/rubrum’ by Rumphius, Herb. Amboin. 3 (1743) 111, t. 73-75. Furthermore, the generic name of Rhizophora was found to be inappropriate, and several subsequent treatments listed other epithets: e.g., Blatti was proposed by Adanson (1763: l.c.) based presumably on a description by Rheede, Hort.Malab. 3 (1682) 43, t. 40. Other treatments followed Linnaeus filius. Engler (1897: 1.c.) proposed an agreeable solution by conserving Sonneratia L. f. as the name of the genus, and, S. caseolaris (L.) Engl. as the type species. However, there was continued confusion with specific epitheton, notably S. caseolaris. These will be discussed later for respective taxa. The name ‘Tombea’, referred to in earlier treatments (notably Backer & Van Steenis, l.c.), was not listed in synonymy. It was also believed to be vernacular (New Caledonia). Brongniart & Gris (1864: 1. c.) believed the entity was Chiratia Montrouzier, although they later synonymised it under Sonneratia (Brongn. & Gris, 1866: l.c.). KEY TO THE SPECIES* 1 a. Calyx smooth, usually shiny; mature fruiting receptacle cup-shaped, the fruit diameter usually < 4 cm, width _< hypanthium width; petioles usually > 0.7 cm long ........................................................................................................................................ 2 b. Calyx coriaceous, often warty and shiny or finely textured; mature fruiting receptacle flat-expanded, fruit diameter usually > 4 cm, width mostly 0.5 cm (or more) greater than hypanthium width; petioles usually c 0.7 cm long . . . . . . . . . 3 2 a. Petals white (maybe tinged green or pink at base) and linear, spathulate or absent; stamina1 filaments white; style length (shortly after anthesis) < 4.5 cm; calyx tube often distinctly ribbed beneath lobe fusion points; fruit surface dull; seeds falcate and smooth. Northern Australia, New Guinea, western Pacific, through Indo-Malesia to China, India and southeastern Africa . . . . . . . 1. S. alba b. Petals red, linear; stamina1 filaments red style length (shortly after anthesis) > 4.5 cm; calyx tube not ribbed beneath lobe fusion points; fruit surface satiny; seeds angular and rugose. Northeastern Australia, northern New Guinea, southwestern Pacific to at least northwestern Borneo . . . . . . . . . . . . . . . 3. S. x gulngai 3 a. Petals red, linear; stamina1 filaments red or white; style length shortly after anthesis > 5 cm; peduncle often tetragonous; calyx lobes flat, expanded; calyx surface shiny, slightly coriaceous; leaf base attenuate to shortly so; leaf mucronate apex distinct ..................................................................................................................................4 b. Petals absent; stamina1 filaments white; style length shortly after anthesis < 4.5 cm; peduncle terete; calyx lobes distinctly reflexed, adpressed; calyx surface verruculose; leaf base rounded to truncate to subcordate;.leaf mucronate apex absent or minute. Southern New Guinea, Indo-Malesia to Thailand and China 5. S. ovata * Observations on colour, texture and form are generally field based. Detailed measurements were taken from dried specimens, unless otherwise stated. Mean values are often given in brackets immediately following attribute ranges of specimen means listed in the specific descriptions. BLUMEA - VOL. 32, No. 2, 1987 286 4 a. Stamina1 filaments red, rarely white; calyx tube grooved; leaves mostly obovate, length to breadth ratio usually < 2; flower buds medially constricted fruit surface shiny, coriaceous. Northeastern Australia, northern New Guinea, southwestern Pacific, through Indo-Malesia to China and India . . . . . . . . . . . . . . 2. S. caseolaris b. Stamina1 filaments white; calyx tube not grooved or ridged; leaves mostly lanceolate, length to breadth ratio usually > 2; flower buds with no distinct medial constriction; fruit surface shiny, smooth. Northern Australia, southern New Guinea, to at least Borneo and Celebes . . . . . . . . . . . . . . . . . . . . . . . . . 4. S. lanceolata Additional, but rare taxa: Solitary tree distinguished from S. x gulngai by absence of petals. Northeastern 3a. S. alba x S. gulngai Australia................................................................... Two specimens distinguished from S. 1anceoIata by cup-shaped calyx tube on mature fruit. Northern Australia, southern New Guinea 4a. S. alba x S. lanceolata 1. Sonneratia alba J.Smith - Fig. 3. Sonnerati alba J.Smith in Rees, Cycl. 33, 2 (1819); DC., Prod. 3 (1828) 231; Blume, Mus. Bot. Lugd.-Bat. 1 (1851) 338; Miq., Fl. Ned. Ind. 1, 1 (1855) 497; Brongn. & Gris, Ann. Sci. Nat. Bot. 6 (1866) 266; Bull. Soc. Bot. France 13 (1866) 479; Kurz, For. Fl. Burma 1 (1877) 526; C.B.Clarke, Fl. Brit. India 2 (1879) 580; Fern.-Vill., Nov. App. (1880) 92; Bisschop Grevelink, Pl. Ned. Ind. (1883) 163; Ceron, Cat. PI. Herb. Manila (1892) 86; Koord. & Valeton, Bijdr. Booms. Java 1 (1894) 200; Koord.,Versl. Minahasa (1898) 470; F.M. Bailey, Queensl. Fl. (1900) 679; Ann. Rep.Brit. New Guinea (1901) 143; Becc., Nelle For. di Born. (1902) 579; Koord., Exk. Fl. Java 2 (1912) 663; F.M.Bailey, Compr. Cat. Queensl. Pl. (1913) 214; Merr., Interpr. Herb. Amboin. (1917) 383; Backer, Bull. Jard. Bot. Buitenz. III, 2 (1920) 330; E.G. Baker, J. Linn. Soc. Bot. 15 (1921) 307; Engl. & Drude, Veg. Erde 9 (1921) 655; Gagnep. & Guillaumin, Fl. Gén. Indo-Chine 2 (1921) 980; Merr., Enum. Born. Pl. (1921) 418; CT. White, Proc. Roy. Soc. Queensl. 34 (1922) 45; Meindersma, Tectona 15 (1922) 573, f. 17; Trop. Natuur 12 (1923) 77, f. 17; R.Parker, Indian For. 51 (1925) 507; C.T. White, J. Bot. 64 (1926) 217; K.Heyne, Nutt. Pl. Ned.-Ind. (1927) 1156; Ridley, Disp. Pl. (1930) 293; Troll, Trop. Natuur 22 (1933) 33, cum ic.; Kint, Trop. Natuur 23 (1934) 173, f. 9; Kaneh., J. Jap. Bot. 14 (1938) 421, f. 1-3; Brenan & Greenway, Checkl. For. Trees Brit. Emp., Tangany. Terr. V, 2 (1949) 592; Steenis, Fl. Scholen Indonesië (1949) 292; Backer & Steenis, Fl. Males. I, 4 (1951) 285, f. 3b; Wyatt-Smith, Malayan For. 16 (1953) 214, t. 1, f. 3; H. Perrier, Fl. Madag. (1954) fam. 148, 2, f. 1; Mahabale & Deshpande, Palaeobotanist 6 (1957)52, t. 1; Cuf., Bull. Jard. Bot. État Brux. 29, Suppl. (1959) 612; Dale & Greenway, Kenya Trees Shrubs (1961) 539, f. 98; Backer & Bakh. f., Fl. Java 1 (1963) 257; Vu VanCuong, Fl. Cambodia Laos Vietnam 4 (1965) 196, pl. 1; Muller & Hou-Liu, Blumea 14 (1966) 337, t. 1, f. lc & 3c; Steenis, N. Queensl. Naturalist 35 (1968) 5, cum ic.; Muller & Steenis, ibid. 35 (1968) 7; G. Williams Sangai, Fl. Trop. E. Africa (1968) 1, f. 1; J.E.D.Fox, Ann. Rep. For. Res. Malay. (1970) 99; W.T.Jones, N. Queensl. Naturalist 20 (1971) 50; Backer& Steenis, Fl. Males. I, 6 (1972) 973, f. 17e-g & 18c; Whitm., Malayan For. Rec. 26 (1972) 445; Lind & Morrison, East Afr. Veg. (1974) f. 4 (2d); Chai, Malaysian For. 38 (1975) 202; Frodin et al., Univ. Papua New Guinea Occas. Pap. 3 (1975) 42; Percival& Womersley, Bot. Bull., Dept. For., Papua New Guinea 8 (1975) 85, 88, f. 65; Chapman, Mangrove Veg. (1976) 388, f. 291c; Floyd, Ecol. Rep., Dept. For. Papua New Guinea 4 (1977) 45, 47; Semeniuk et al., Mangroves W. Austral. (1978) 73, pl. 30; A.G.Wells in Clough, Mangrove Ecosyst. Austral. (1982) 74; in Teas, Tasks Veg. Sci. 8 (1983) 71; Ko, Acta Phytotax. Sin. 23 (1985) 313. - Rhizophora caseolaris L, Herb. Amb. N.C. Duke & B.R. Jackes: Revision of Sonneratia in Australasia 287 (1754) 13, p.p. - Sonneratia acida auct. non L. f.: Benth,, Fl. Austral. 3 (1866) 301, p.p.; Hiern, Fl. Trop. Afr. 2 (1871) 483; Baker, Fl. Mauritius (1877) 102; Volkens, Bot. Jahrb, . Syst. 31 (1901) 47: Dupont, Rep. Invest. St. Pierre Astove, Kew Libr. (1907) 24; Hemsley, Kew Bull. (1919) 122. - Blatti alba Kuntze, Rev. Gen. Pl. 1 (1891) 238; Niedenzu in E.& P., Nat. Pfl. Fam. 3, 7 (1892) 21, f. 7h & j. - Sonneratia caseolaris auct. non (L.) Engl.: Engl., Pflanzenwelt Ost-Afrikas C (1895) 286 in E. & P., Nat. Pfl. Fam. Nachtr. 1 (1897) 261, ex parte; sensu Merr., Enum. Philipp. Flow. Pl. 3 (1923) 139; Panshin, Philipp. J. Sci. 48 (1932) 161, pl. 7; Kaneh., Fl. Micron. (1933) 251, f. 117; Enum. Micron. Pl. (1935) 375; R. Williams, Useful Om. Pl. Zanzibar Pemba (1949) 449, cum ic., ex parte; Blake, Austral. J. Bot. 2 (1954) 136; Fernando & Pancho, Sylvatrop. Philipp. For. Res. J. 5 (1980) 52.- Sonneratia pagatpat auct. non Blanco: Merr., Bull. Bur. For. Philipp. 1 (1903) 42; Philipp. J. Sci. 1 (1906) Suppl. 101; Fl. Manila (1912) 344.- Sonneratia grifithii auct. non Kurz: Watson, Malayan For. Rec. 6 (1928) 50,55,120, f.24; Burkill, Dict.2 (1935) 2053. - Type: Rumphius, Herb. Amboin. 3 (1743) 111, t. 73!, Indonesia, Amboina. Chiratia leucantha Montr., Mém. Acad. Sci. Lyon, Sect. Sci. 10 (1860) 203; Brongn &. Gris, Ann. Sci. Nat. Bot. 1 (1864) 364; Bull. Soc. Bot. France 11 (1864) 71. -Types: Vieillard 432, Pancher s.n., Deplanche 50.5 (syn.; n.v.), New Caledonia. Sonneratia mossambicensis Klotzsch in W. Peters, Naturw. Reise Mossambique 1 (1862) 66, pl. 12. - Sonneratia acida var. mossambicensis (Klotzsch) Mattei, Boll. Reale Orto Bot. Giard. Colon. Palermo 7 (1908) 108. - T ype: Peters s.n. (B, holo; K, iso; n.v.), Africa, Mozambique. Sonneratia iriomotensis Masam., Syokubutu-tirigaku (1936) f. 71; Trans. Nat. Hist. .Soc. Taiwan 29 (1939) 272. - Sonneratia alba J. Smith var. iriomotensis (Masam.) Yamamoto, J. Soc. Trop. Agric. 12 (1940) 162. - Type : unknown, Japan? Sprawling to erect tree c. 20 m high. Trunk base not buttressed. Bark smooth or lightly fissured and flaky, dark grey to pale fleshy coloured. Pneumatophores stocky, bluntly pointed, c. 25 cm high. Leaves: lamina elliptic or ovate (often widely so), 48-107 (74) mm long, 25-76 (47) mm wide, pale green, dull on upper surface, satin lustre on lower surface, midvein rarely reddened, lamina not decurrent on petiole; apex broad, mucro recurved, adpressed to leaf undersurface; petiole 6-15 (9) mm long, terete. Inflorescence with 1-5 (2) buds; stems beneath mostIy terete. Mature bud: apex acute to obtuse, attenuate to obtuse at base, medially constricted, upper portion 13-15 (14) mm wide, lower portion 12-22 (16) mm wide, in all 2033 (28) mm long. Flowers: hypanthium glossy, smooth, often ribbed beneath fusion point of calyx lobes; calyx lobes 5-8 (6 or 7), 13-19 (16) mm long, acute apex, inner surface often reddish; petal numbers and presence variable, linear, 14-30 (20) mm long, 1-2 (1) mm wide when fresh, to spathulate or stamen-like, mostly white, often tinged red or green at base; staminal filaments 15-45 (29) mm long when fresh, entirely white; anthers dorsifixed, pale yellow; petals and stamens fall within hours of anthesis; ovary 12-20-(16-)locular; style c. 43 mm long, stigma fungiform, c. 3 mm wide. Berry erect, globose, 4-27 (14) mm long, 22-46 (32) mm wide, pericarp coriaceous, dull without ribs; calyx persistent, hypanthium cupshaped, 25-41 (32) mm wide, 27-46 (37) mm from calyx base to lobe apex, lobes erect, spreading 17-26 (20) mm long. Seeds numerous, c. 12 mm long, falcate. P h e n o l o g y . Leaf fall peaks in January. Flowering is common in October and November. Fruiting (in the Murray River, Queensland, c. 40% of units attains maturity from immature bud stages; c. 30% are aborted at the immature fruit stage) occurs mainly around February (ranging from January to March). 288 BLUMEA - VOL. 32, No. 2, 1987 D i s t r i b u t i o n . The species is found from southeastern Africa to India and southern China to the western islands of the Pacific Ocean. In Australia the species is found from Port Clinton (22º35' S, 150º45' E) on the east coast, across northern Fig. 3. Sonneratia alba J. Smith. 1. Mean range of leaf outlines: 2. ‘lip’-Iike leaf mucronate apex; 3. petiole base; 4. mature flower bud (longitudinal section) showing general internal anatomy just prior to anthesis; 5. flower at anthesis (petalled form); 6. floral diagram (semi-petalled form); 7. stamen; 8 & 9. two forms of staminal-form petals; 10.. spathulate petal; 11. linear petal; 12. mature fruit. Scale = 1 cm. N.C. Duke & B.R. Jackes: Revision of Sonneratia in Australasia 289 Australia to Cape Bossut (18º43'S, 121º38'E) on the west coast. The species was similarly widespread about New Guinea, although it is less common on the northern coast. Other locations include: New Caledonia, New Hebrides, Solomon Islands, and generally throughout SW. Oceania. Fig. 1. E c o 1 o g y . This species occurs at lower tidal contours in frontal stands, as isolated trees in downstream lower estuarine situations or about offshore islands in areas of high to moderate rainfall and tidal range greater than one metre. Commonly associated with Rhizophora stylosa, Aegiceras corniculatum and Avicennia marina, and usually growing in either sand, gravel or soft mud. Notes. Since J. Smith described S. alba (1819: l.c.) from the figure and description of ‘Mangium caseolare album’ Rumphius (1743: l.c.), our view of this taxon has been relatively stable. However, one aspect has altered significantly. In earlier keys (e.g., Backer & Steenis, 1951: l.c.), the presence or absence of petals was used as a definitive key attribute. Unfortunately, there were a variety of petalous conditions in S. alba. Furthermore, the occurrence of less-petalled forms was related to marginal habitats resulting from both climatic and localised factors. Thus in colder climates, towards the southern latitudinal limits, apetalous and semipetalous forms were more common. In more equatorial areas, petalled forms were common in riverine estuaries and less-petalled forms were found more frequently offshore on smaller rocky islands and coral cays. This apparent ecotypic variability precluded even the use of intraspecific epitheton based on petallous condition in this taxon. 2. Sonneratia caseolaris (L.) Engl. - Fig. 4. Sonneratia caseolaris (L.) Engl. in E. & P., Nat.Pfl.Fam.Nachtr. 1 (1897) 261; Koord. & Valeton, Bijdr. Booms. Java 8 (1902) 156; Merr., Fl. Manila (1912) 344; Druce, Bot. Exch. Club Brit. Isles 3 (1914) 424; Merr., Interpr. Herb. Amboin. (1917) 383;Enum. Born. Pl. (1921) 418; R. Parker, Indian For. 51 (1925) 507; Panshin, Philipp. J. Sci. 48 (1932) 143; Burkill, Dict. 2 (1935) 2052; Kaneh., J.Jap.Bot. 14 (1938) 423, f. 3 l-n; Steenis, Fl. Scholen Indonesië (1949) 291; Merr., J.Amold Arbor. 31 (1950) 285; Backer& Steenis, Fl. Males. I, 4 (1951) 283, f. 3c; Wyatt-Smith, Malayan For. 16 (1953) 214, t. 1, f. 1; Backer & Steenis, Fl. Males. I, 5 (1958) 537; Steenis, Nova Guinea n.s. 12 (1963) 189; Vu VanCuong, Fl. Cambodia Laos Vietnam 4 (1965) 199, pl. 1; Muller & Hou-Liu, Blumea 14 (1966) 337, t. 1, f. le & 3e; Steenis, N. Queensland Naturalist 35 (1968) 3, cum ic.; Muller & Steenis, ibid. 35 (1968) 8; J.E.D. Fox, Ann. Rep. For. Res. Malay. (1970) 99; Backer & Steenis, Fl. Males. I, 6 (1972) 973, f. 17a-d & 18e; Whitm., Malayan For. Rec. 26 (1972) 445; Chai, Malaysian For. 38 (1975) 203;Perci val & Womersley, Bot. Bull., Dept. For., Papua New Guinea 8 (1975) 88, f. 66; Chapman, Mangrove Veg. (1976) 388, f. 291e; Floyd, Ecol. Rep., Dept. For., Papua New Guinea 4 (1977) 44 ic. only; A.G. Wells in Teas, Tasks Veg. Sci. 8 (1983) 72, ex parte; Ko, Acta Phytotax. Sin. 23 (1985) 313. -Rhizophora caseolaris L., Herb. Amb. (1754) 13, ex parte; Amoen. Acad. 4 (1759) 123; Syst. Nat. ed. 10 (1759) 1043; Sp. Pl. ed. 2 (1762) 635; Burm. f., Fl. Indica (1768) 108; Lour., Fl. Cochinch. ed. 2 (1793) 296, 363. - Sonneratia acida auct. non L. f.: Lam., Encycl. 1 (1785) 429; Tabl. Encycl. 2 (1794) t. 420; J. Smith in Rees, Cycl. 33, 2 (1819); Roth, Nov. Pl. Sp. (1821) 233; Sprengel, Syst. Veg. 2 (1825) 493; DC., Prod. 3 (1828) 231; Roxb., Fl. Ind. ed. 2 (1832) 506; Wight & Arn., Prod. Fl. Penins. Ind. Or. 1 (1834) 327; Decne., Nouv. Ann. Mus. Hist. Nat. 3 (1835) 454; Spanoghe, Linnaea 15 (1841) 203; Wight, Icon. Pl. 2 (1843) 340; Korth., Ned. Kruidk. Arch. 1 (1846) 198; Blume, Mus. Bot. Lugd.-Bat. 1 (1851) 336; A. Gray, U.S. Expl. Exp., Phan. 15 (1854) 550; Griff., Not. Pl. 290 BLUMEA - VOL. 32, No. 2, 1987 Fig. 4. Sonneratia caseolaris (L.) Engl. 1. Mean range of leaf outlines; 2. variable shaped leaf mucronate apex; 3. petiole base; 4. mature flower bud (longitudinal section) showing general internal anatomy just prior to anthesis; 5. flower at anthesis; 6. floral diagram; 7. petal; 8. mature fruit. Scale = 1 cm. N.C. Duke & B.R. Jackes: Revision of Sonneratia in Australasia 291 Asiat. 4 (1854) 652; Miq., Fl. Ned. Ind. 1, 1 (1855) 496, p.p.; Dalzell, Bombay Fl. (1861) 98; Miq., Sumatra (1862) 120,316; Brandis, For. Fl. N.W. India (1874) 242; Kurz, For. F’l. Burma 1 (1877) 526; C.B. Clarke, Fl. Brit. India 2 (1879) 579;Fern.-Vill., Nov. App. (1880) 92; Vidal, Sin. Gen. Pl. Lenos. Filip., Atlas 27 (1883) t. 52, f. G; Boerl., Handl. Fl. Ned. Ind. 1 (1890) 538; Koord. & Valeton, Bijdr. Booms. Java 1 (1894) 198; Merr., Philipp. J. Sci. 3 (1908) Bot. 83; Pulle, Nova Guinea, Bot. 8 (1911) 677, 679; Whitford, Bull. Bur. For. Philipp. 10 (1911) Sonn.; Koord., Exk. Fl. Java 2 (1912) 663; Atlas t. Java (1915) t. 592-4; Moll & Janssonius, Mikrogr. Holz. 3 (1918) 603; Gamble,Fl. Madras (1918/19) 363, 515; N. Brown, For. Products (1919)Sonn.; Backer, Bull. Jard. Bot.Buitenz. III, 2 (1920) 330; Foxw., Malayan For. Rec. 1 (1921) 117; Gagnep.& Guillaumin, Fl. G&n. Indo-Chine 2 (1921) 979; Troup, Silvic. Ind. Trees (1921) 2, f. 228; Ridley, Fl. Malay. Penins. 1 (1922) 825; Meindersma, Tectona 15 (1922) 573, f. 18; Trop. Natuur 12 (1923) 77, f. 18;Merr., Enum. Philipp. Flow. Pl. 3 (1923) 138; C.T. White J. Bot. 64 (1926) 217; K. Heyne, Nutt. Pl. Ned.-Ind. (1927) 1156; Watson, Malayan For. Rec. 6 (1928) 50, 55, 120, f. 22; Ridley, Disp. Pl. (1930) 293; Panshin, Philipp. J. Sci. 48 (1932) 163, pl. 8; Kint, Trop. Natuur 23 (1934) 182 Mahabale & Deshpande, Palaeobotanist 6 (1957) 52, t. 1; Backer& Bakh. f., Fl. Java 1 (1963) 257.Aubletia caseolaris Gaertn., De Fruct. Sem. Pl. 1 (1788) 379, t. 78, f. 2 p.p. , - Sonneratia rubra Oken, Allg. Naturgesch. 3 (1841) 1952.- Blatti caseolaris Kuntze, Rev. Gen. Pl. 1 (1891) 238; Niedenzu in E. & P., Nat. Pfl. Fam. 3, 7 (1892) 21, f. 7a-g, p.p. - Sonneratia alba non J. Smith: Fernando & Pancho, Silvatrop. Philipp. For. Res. J. 5 (1980) 51.- T y p e: Rumphius , Herb. Amboin. 3 (1743) 112, t. -74 51, Indonesia, Amboina. Sonneratia acida L. f., Suppl. Pl. (1781) 252. - Sonneratia pagatpat Blanco, Fl. Filip. (1837) 424; ed. 2 (1845) 296; Blume, Mus.Bot. Lugd.-Bat. 1 (1851) 337; Miq., Fl. Ned. Ind. 1, 1 (1855) 496; Blanco, Fl. Filip. ed. 3 (1878) 186. - Blatti pagatpat Niedenzu in E. & P., Nat. Pfl. Fam. 3, 7 (1892) 21. - T y p e : Sonnerat s.n. (LINN, iso), New Guinea. Sonneratia ovalis Korth., Ned. Kruidk. Arch. 1 (1846) 198; Miq., Anal. Bot. Ind. 1 (1850) 28. Type: Korthals s.n. in herb. L (sh. nr. 908.128-980), Borneo, Bandjermasin. Sonneratia neglecta Blume, Mus. Bot. Lugd.-Bat. 1 (1851) 338; Miq., Fl.Ned.lnd. 1, 1 (1855) 498. - Type: herb. P. Royen in herb. L (sh. nr. 908.128-961). Ceylon? Sonneratia evenia Blume, Mus. Bot. Lugd.-Bat. 1 (1851) 337; Miq., Fl. Ned. Ind. 1, 1 (1855) 497. - Type: Blume s.n. in herb. L(sh. nr. 908.128-971). Indonesia, Timor. Sonneratia obovata Blume, Mus. Bot. Lugd.-Bat. 1 (1851) 337; Miq., Fl. Ned. Ind. 1, 1 (1855) 497; Koord. & Valeton, Bijdr. Booms. Java 1 (1894) 198.- Type: Blume s.n. in herb. L (sh. nr. 908.128-990), Indonesia, Java, Tjilakahan. Sonneratia acida auct non L. f. var.mucronata Miq., Fl. Ned. Ind.1, 1 (1855) 496, p.p.; R. Parker, Indian For. 51 (1925) 507.- T y p e: unknown, Indonesia, Java. Columnar tree c. 20 m high, canopy generally sparse. Trunk base not buttressed. Bark smooth or lightly fissured and flaky, grey or flesh coloured. Pneumatophores thin, pointed and often branched, c. 1 m high. Leaves: lamina elliptic, 45-113 (79) mm long, 20-71 (52) mm wide, deep green satiny upper surface, lower surface shiny, midvein prominent with reddish coloration often on upper surface towards petiole base; apex acute, thickened, mucro recurved; petiole 2-9 (5) mm long, lamina decurrent. Inflorescences with 1-3 (mostIy 1 or 2) buds; peduncles terete or tetragonous. Mature bud: apex mostly acute, attenuate to obtuse base, medially constricted, upper portion c. 15 mm wide, lower portion 13-17 (15) mm wide, in all 24-27 (25) mm long. Flowers: hypanthium satiny, coriaceous, tending to warty with pronounced ribs along calyx lobe margins, deep grooves beneath junctures; calyx lobes 5-7 (6), 14-19 (17) mm long, apex acute, inner surface often streaked with red coloration; petals always present, linear, 19-29 (25) mm long, 1-3 (2) mm wide when 292 BLUMEA - VOL. 32, No. 2, 1987 fresh, red; stamina1 filaments c. 37 mm long when fresh, red, rarely white; anthers dorsifixed, pale yellow; petals and stamens fall within hours after anthesis; ovary 13-20- (16-)locular; style c. 62 mm long, stigma fungiform, c. 3 mm wide. Berry globose, 7-32 (20) mm long, 17- 54 (41) mm wide; pericarp leathery, glossy with slight ribs; calyx persistent, hypanthium flat-expanded, 20-46 (34) mm wide, 2746 (37) mm from calyx base to lobe apex, lobes erect, spreading, 18-30 (25) mm in length. Seeds numerous, irregularly angular, c. 7 mm long. P h e n o l o g y . Leaf fall occurs chiefly from October to January. Flowering occurs mainly around January. Along the Murray River, only 6% of the buds develop to mature fruit, about 60% of fruit set reach maturity. Fruit maturation occurs predominantly in June and July. D i s t r i b u t i o n . The species is found from the west coast of India to southem China and through the western islands of the Pacific Ocean. Distribution in Australia is limited, occurring only on the northeast coast where it is associated with areas of higher rainfall and large river estuaries from the Murray River (18º05' S, 146º0l'E) in the south to the Olive River (12ºl0'S, 143º05' E) in the north.In New Guinea, the taxon is only found on the north coast and on larger islands. Fig. 1. E c o 1 o g y . The species occurs in frontal stands often in upstream estuarine positions of rivers subjected to high levels of freshwater runoff. An alliance is observed with Aegiceras corniculatum, Barringtonia racemosa and Rhizophora mucronata. Substrate was usually soft river silt and mud of the accreting inside bank of estuarine meanders. Notes. There was some confusion in attaching an epithet to this species. This confusion was based on at least two major errors or omissions in the original type description. Firstly, Linnaeus (1754: l.c.) named only one taxon from the Rumphius descriptions (1743: l.c.) of ‘Mangium caseolare’ - being made up of two distinct forms, viz., ‘album’ and ‘rubrum’. The latter form was generally believed to be that which Linnaeus had described, but it is impossible to be sure. J. Smith (1819: 1. c.) partially clarified the situation by separately identifying the two taxa, and in naming ‘f. album’ as S. alba. The second confusing aspect was in regard to the other form, ‘ f . rubrum’, and, in the Linnaeus fil. (1781: l.c.) description of S. acida. This latter description (as already noted in the generic section) was based on different type material, and subsequent authors apparently used a mixture, viz., Linnaeus type material and Linnaeus fil. epithet. This confusion generally continued until the major revision by Backer & Van Steenis (1951: l.c.). These authors rightly reduced the Linnaeus fil. name to synonymy, and used ‘Mangium caseolare rubrum’ ’ as the type description. We follow Backer & Van Steenis in this, and for other type material not seen by us. Sonneratia x gulngai N.C. Duke - Fig. 5. Sonneratia x gulngai N.C. Duke, Austrobaileya 2 (1984) 103. - Sonneratia alba x S. caseolaris Muller & Hou-Liu, Blumea 14 (1966) 337, t. 1, f. ld, 2b & 3d; Muller & Steenis, N. Queensl. Naturalist 35 (1968) 8; Backer & Steenis, Fl. Males. I, 6 (1972) 974, f. 18d; Chapman, Mangrove Veg. (1976) f. 291d. - Type: N.C.Duke AIMS 547 (BRI, holo!; AIMS, CANB, JCT, K, L, iso!), Australia, Queensland,McIvor River. N.C. Duke & B.R. Jackes: Revision of Sonneratia in Australasia 293 Fig. 5, Sonneratia x gulngui N.C. Duke. 1. Mean range of leaf outlines; 2. variable shaped leaf mucronate apex; 3. petiole base; 4. mature flower bud (longitudinal section) showing general internal anatomy just prior to anthesis; 5. flower at anthesis; 6. floral diagram; 7. petal; 8. mature fruit. Scale = 1 cm. 294 BLUMEA - VOL. 32, No. 2, 1987 Spreading tree c. 25 m high; canopy rather dense. Trunk base not buttressed. Bark smooth or fissured and flaky, grey. Pneumatophores thin, pointed and often branched, c. 80 cm high. Leaves: lamina obovate, 52-85 (80) mm long, 38-64 (54) mm wide, dull green upper surface, satin lustre on lower surface, midvein often tinged red towards petiole; apex acute, thickened, mucro recurved; petiole 6 -14 (11) mm long, lamina decurrent. Inflorescences with one or two buds; stems beneath mostly terete. Mature bud: apex acute to obtuse, attenuate to obtuse at base, medially constricted, the upper and lower portions often similar, in all 25 -35 (3 1) mm long, 10-20 mm wide. Flower: hypanthium dull, smooth, without ribs; calyx lobes 5-7 (6), 15-23 (18) mm long, acute apex, often reddish on inner surface; petals always present, linear, 20-41 (33) mm long, 1-3 (2) mm wide when fresh, red; stamina1 filaments c. 37 mm long when fresh, red; anthers pale yellow; petals and stamens fall within hours after anthesis; ovary 13-17- (15-)locular; style c. 30 mm long, stigma fungiform, c. 1.7 mm wide. Berry erect, globose, 10-20 (12) mm long, 29-48 (39) mm wide, sometimes indented around style base; pericarp leathery, smooth, satiny lustre, without ribs; calyx persistent, hypanthium cup-shaped, 31-45 (38) mm wide, 41-47 (44) mm from calyx base to lobe apex, lobes erect, 23-28 (27) mm long. Seeds numerous, angular. P h e n o l o g y . Leaf fall peaks twice, in August to September and March to May. Floral development of the hybrid is complex and is coupled with observations of high levels of aborted pollen (c. 30% collapsed grains) and low fruit set and maturation (along the Murray River, c. 1% of the potential crop attain maturity). The dual character of the hybrid is briefly summarised by the apparent bimodal peak of flowering and fruiting periods which coincide in this region with those of S. alba and S. caseolaris, respectively. Flowering occurs as two peaks based on December and also in March. Mature fruits fall mainly in March and August. D i s t r i b u t i o n . This putative hybrid has been reported (Muller & Hou-Liu, 1966) to occur in northwestern Borneo. In Australia, S. x gulngai is found on the northeastern coast from the Murray River (18º05'S, 146’01 ‘E) in the south, to the Olive River (12ºl0'S, 143º05'E) in the north. It is limited to estuaries which have both S. caseolaris and S. alba. The latter species is widespread, therefore the possibIe limitations on the distribution in Australia of the hybrid are linked to the occurrence of the less common S. caseolaris. Occurrence is more frequent in estuaries where the distributions of S. alba and S. caseofaris overlap. Distribution throughout Indo-Malesia is expected to be widespread while subject to the constraints already outlined. Fig. 1. E c o 1 o g y . Sonneratia x gulngai is easily distinguished in the field by its size and Iuxuriant foliage (i.e., darker green canopy appearance wlthIarger leaves and flower buds). It is found commonly at the downstream limits of S. caseofaris which often places it centrally within Australian estuaries. In this position the species occurs in lower to middIe tidal contours; i.e., above and behind S. caseofaris. Here it is commonIy associated with Bruguiera parviflora and Xylocarpus granatum. To a lesser extent the species is also associated with Rhizophora mucronata and Nypa fruticans. Sonneratia x gulngai is usually found on firm mud or silt. N.C. Duke & B.R. Jackes: Revision of Sonneratia in Australasia 295 Notes. Sonneratia x gulngai is comparable with S. alba x S. caseolaris Muller & Hou-Liu (1966: 1. c.), but there are some differences. However, the Australian axon has consistent morphological characteristics throughout the region; particularly in northeastern Australia. The taxon is therefore recognised as a distinct hybrid species. Its hybrid status is supported by the following evidence: - intermediate and shared morphological characters; - reduced fertility and distinctive shape and character of the pollen; - poor fruit maturation; - complex floral phenology, apparently taking characteristics of each of the putative parental cycles; - luxuriant growth of tree form and foliage; - distribution in Australia limited to those estuaries where S. caseolaris also occurs; - number of individuals usually quite low; more plants are found in those estuaries where the distributions of S. alba and S. caseolaris overlap. 3a. Sonneratia alba x Sonneratia gulngai Spreading tree c. 15 m high. Trunk base not buttressed. Bark smooth or lightly fissured and flaky, pale grey coloured. Leaves: lamina elliptic or ovate, 63 mm long, 37 mm wide, pale green dull on upper surface, satin lustre on lower surface, midvein green, lamina decurrent on petiole; apex broadly acute, mucro recurved, adpressed to leaf undersurface; petiole 7 mm long, terete. Inflorescences with only one bud; stems beneath terete. Mature bud: apex acute to obtuse, attenuate to obtuse at base, medially constricted, upper portion 12 mm wide, lower portion 11 mm wide, in all 22 mm long. Flowers: hypanthium dull, smooth, without ribs or grooves; calyx lobes 5-7 (6), 13 mm long, apex acute, inner surface green tinged red at base; petals absent; stamina1 filaments entirely red; anthers dorsifixed, pale yellow; stamens fall within hours of anthesis; ovary 12-17- (15-)locular; style c. 39 mm long, stigma fungiform, c. 3 mm wide. Berry erect, globose, 15 mm long, 31 mm wide, pericarp coriaceous, dull, without ribs; calyx persistent, hypanthium cup-shaped, 31 mm wide, 33 mm from calyx base to lobe apex, lobes erect, spreading, 20 mm long. Seeds numerous, angular. P h e n o l o g y. Mature buds and fruit were collected in August and September. D i s t r i b u t i o n . Only one tree was observed throughout the region. This tree is located in the McIvor River (15º08'S, 145º14' E), Australia, around 8 km upstream from the estuary mouth and on the waters’ edge. Reference collections held at AIMS include N. C. Duke AIMS 702. Notes. This taxon is distinguished by its red stamina1 filaments and lack of petals on specimens which otherwise resembled S. alba. This latter species (the downstream form) and S. caseolaris (the upstream form) are common in the McIvor River estuary. Sonneratia x gulngai, the predominant hybrid form, is well represented and there are numerous specimens in the overlap zone. As all these forms occur in close proximity it is considered that opportunities for hybridization are high. This significant overlap in Sonneratia taxa is not observed elsewhere in Australia. 296 BLUMEA - VOL. 32, No. 2.1987 Fig. 6. Sonneratia lanceolata Blume. 1. Mean range of lanceolate leaf outline; 2. ibid. of ovate leaf outline; 3. petiole base; 4. mature flower bud (longitudinal section) showing general internal anatomy just prior to anthesis; 5. flower at anthesis (common single petal form); 6. floral diagram; 7. thickened second petal of the uncommon double-petalled form; 8. normal linear petal; 9. mature fruit. Scale = 1 cm. N.C. Duke & B.R. Jackes: Revision of Sonneratia in Australasia 297 4. Sonneratia lanceolata Blume - Fig. 6. Sonneratia lanceolata Blume, Mus. Bot. Lugd.-Bat. 1 (1851) 337; Miq., Fl. Ned. Ind. 1, 1 (1855) 497; Koord., Versl. Minahasa (1898) 471; Becc., Nelle Foreste di Born. (1902) 579; CT. White, Proc. Roy. Soc. Queensl. 34 (1922) 45; R. Parker, Indian For. 51 (1925) 505.- Sonneratia acida auct. non L. f.: Benth., Fl. Austral. 3 (1866) 301, p. p.; Ewart & Davies, Fl. North. Terr. Austral. (1917) 199; Blake, Austral. J. Bot. 2 (1954) 137. - Sonneratia alba auct. non J. Smith: Merr., Enum. Born. Pl. (1921) 418. - Sonneratia caseolaris auct. non (L.) Engl., ex parte: Ridley, Fl. MaIay. Penins. 1 (1922) 825; Backer & Steenis, Fl. Males. I, 4 (1951) 283, ex parte; Percival & Womersley, Bot. Bull., Dept. For., Papua New Guinea 8 (1975) 88, ex parte; Frodin et al., Univ. Papua New Guinea Occas. Pap. 3 (1975) 44; Floyd, Ecol. Rep., Dept. For., Papua New Guinea 4 (1977) 47, non ic.; A.G. Wells in Clough, Mangrove Ecosyst. Austral. (1982) 75; in Teas, Tasks Veg. Sci. 8 (1983) 72, ex parte. - Type : Blume s.n. in herb. L (sh. nr. 908.128-988), Indonesia, Borneo. Columnar tree c. 20 m high, canopy generally sparse. Trunk base not buttressed. Bark smooth or lightly fissured and flaky, grey to paIe fleshy colour. Pneumatophores thin, pointed, c. 20 cm high. Leaves: lamina elliptic to lanceolate, 61-124 (86) mm long, 12-38 (25) mm wide, dull pale green, upper and lower surfaces similar, midvein rarely reddened, apex acute, slightly thickened, mucro recurved; petiole 2-7 (4) mm long, lamina decurrent. Inflorescences with mostly one but sometimes two buds; peduncles terete or tetragonous. Mature bud: apex acute to obtuse, base mostly obtuse, no medial constriction, 20-26 (23) mm long, 12-18 (15) mm wide. Flowers:hypanthium dull, slight leathery with no ribs or grooves; calyx lobes 5 -7 (6), 13-14 (14) mm long, apex acute, inner surface rarely with reddish streaks; petals always present, rarely doubled, linear, 12-38 (29) mm long, 2-4 (3) mm wide when fresh, red; stamina1 filaments c. 45 mm long when fresh, always white; anthers dorsifixed, yellow; petaIs and stamens fall within hours after anthesis; ovary 12-17- (15-)locular; style c. 56 mm long, stigma fungiform, c. 3 mm wide. Berry globose, 9-18 (14) mm long, 25-38 (31) mm wide; pericarp smooth, glossy, with slight ribs; calyx persistent, hypanthium flat-expanded, 24-30 (26) mm wide, 2430 (27) mm from caIyx base to lobe apex, lobes erect, spreading, 14-19 (16) mm in length. Seeds numerous, angular, c. 7 mm long. P h e n o l o g y. No information exists on leaf fall periodicity. Flowering was observed in September to October and in March. Fruits were numerous in specimens collected during December to March. D i s t r i b u t i o n . Except for the type record, there are few records of this species in countries other than Australia and New Guinea. In Australia this species occurs in the Pascoe (12º3O'S, 143º16'E) and Claudie (l2º50'S, 143º2l'E) Rivers of northeastern QueensIand, and several rivers in three separate locations in the Northern Territory; viz. Arnhem Bay (12º00'S, 136º00'E), Bonaparte Gulf (12º00'S 132º00'E),and MelviIIe Island (llº0l' S, 130º0l'E).It is known onIy on the southern coast of New Guinea. Fig. I. E c o l o g y . The species occurs at lower tida1 contours in frontal stands or as isolated trees in upstream estuarine positions in rivers subjected to relativeIy high Ievels of freshwater runoff. Associated species include Avicennia marina, Nypa fru- 298 BLUMEA - VOL. 32, No. 2.1987 ticans and Bruguiera sexangula. Substrate is usually fine soft silt on the accreting inside banks of river meanders. Notes. In this study there were two taxa referable to S. caseolaris described by Backer & Van Steenis (1951: l.c.). We had to decide which taxon was consistent Fig. 7. Sonneratia ovata Backer. 1. Mean range of leaf outlines; 2. petiole base; 3. mature flower bud (longitudinal section) showing general internal anatomy just prior to anthesis; 4. flower at anthesis; 5. floral diagram; 6. mature fruit. Scale= = 1 cm. N.C. Duke & B.R. Jackes: Revision of Sonneratia in Australasia 299 with that epithet. Our problem was resolved by comparing these taxa with the respective Rumphius description and figures (1743: l.c.). The shape of the leaves was a major diagnostic character. We now recognise the ovate-leaf entity from northeast Australia and northern New Guinea as S. caseolaris. The remaining form had lanceolate leaves and was comparable with S. lanceolata Blume (185 1: 1. c.). Populations of S. lanceolata from neighbouring rivers often exhibit distinct morphological differences. One population isolated from the main estuary of the Wildman River (12º18'S, 132º04' E) has leaves which are typically linear rather than lanceolate, and the petal numbers are often double. 4a. Sonneratia alba x Sonneratia lanceolata Tree to 8 m high. Leaves: lamina elliptic, 77-101 (89) mm long, 35-54 (41) mm wide, midvein prominent under; apex acute, slightly thickened, mucro recurved; petiole 7-12 (10) mm long, lamina not always decurrent. Inflorescence: stems beneath mostly terete. Mature bud: apex acute to mostly obtuse, attenuate to obtuse at base, slight medial constriction, in all 31 mm long, 19 mm wide. Flowers: hypanthium dull, slightIy leathery, with no ribs or grooves; calyx lobes 6, 19 mm long, apex acute; petals lanceolate, red; stamina1 filaments possibly white; style c. 56 mm long, stigma fungiform, c. 1-2 mm wide. Berry erect, globose, 7 mm long, 26 mm wide, not indented about style base; pericarp smooth, glossy, with slight ribs; calyx persistent, hypanthium cup-shaped, 28 mm wide, 39 mm from calyx base to lobe apex, lobes erect, spreading, 23 mm long. P h e n o l o g y. Flowers were collected in June and mature fruit in December. D i s t ri b u t i o n . Only two herbaria specimens were seen. In addition there are tentative sightings from the Kikori region of Papua New Guinea. It is expected, however, that the distribution of this taxon would reflect that of the putative parents. AUSTRALIA. Northern Territory: Goromura River (12º28'S, 136º13'E),Wells s.n. (DNA 15228). INDONESIA . West Irian: Merauke (8º30'S, 140º22'E), McKee 1701 (LAE). Notes. This taxon is distinguished chiefly by its S. alba-like fruit on specimens which otherwise resembIe S. lanceolata. However, the leaves are not so long in relation to their width. Foliage characteristics are expected to be somewhat reminiscent of S. x gulngai with larger leaves and buds and a more luxuriant crown. 5. Sonneratia ovata Backer - Fig. 7. Sonneratia ovata Backer, Bull. Jard. Bot. Buitenz. III, 2 (1920) 329; Meindersma, Tectona 15 (1922) 573, f. 19; Trop. Natuur 12 (1923) 77, f. 19; Steenis, Bull. Jard. Bot. Buitenz. III, 12 (1932) 162; Merr. & Perry, J. Arnold Arbor. 22 (1941) 269; Backer & Steenis, FL Males. I, 4 (1951) 285, f. 3a; Wyatt-Smith, Malayan For. 16 (1953) 214, t. 1, f. 2; Mahabale & Deshpande, Palaeobotanist 6 (1957) 52, t. 1; Backer & Bakh. f., Fl. Java 1 (1963) 257; Vu Van Cuong, Adansonia 4 (1964) 346; Fl. Cambodia Laos Vietnam 4 (1965) 202, pl. 1; Muller & Hou-Liu, Blumea 14 (1966) 337, t. 1, f. la & 3a; Backer & Steenis, Fl. Males. I, 6 (1972) 974; Whitm., Malayan For. Rec. 26 (1972) 445; Chai, Malaysian For. 38 (1975) 203; Percival & Womersley, 300 BLUMEA - VOL. 32, No. 2, 1987 Bot. Bull., Dept. For., Papua New Guinea 8 (1975) 85, f. 64; Frodin et al., Univ. Papua New Guinea Occas. Pap. 3 (1975) 44; Chapman, Mangrove Veg. (1976) 388, f. 291a; Floyd, Ecol. Rep., Dept. For., Papua New Guinea 4 (1977) 47 & 49, cum ic.; Ko, Acta Phytotax. Sin. 23 (1985) 313. - Sonneratia alba auct. non J.Smith: Watson, Malayan For. Rec. 6 (1928) 50, 55, 122, f. 23; Burkill, Dict. 2 (1935) 2051. - Type: Bucker 21422 (L, lecto ex Vu Van Cuong, 1965), Indonesia, Java. Columnar tree c. 20 m high. Trunk base not buttressed. Bark slightly flaky, pale brown to grey. Pneumatophores thin, pointed, c. 20 cm, high. Leaves: lamina very widely ovate, 43-56 (45) mm long, 36-47 (38) mm wide, green, base reniform, upper surface glossy, lower surface satiny, midvein not reddened, leaf apex obtuse, mucro absent; petiole 5-6 (5) mm long, terete. Inflorescence with 1-3 (2) buds; peduncle terete. Mature bud: apex obtuse, base mostly attenuate, medially constricted, with upper portion c. 17 mm wide, lower portion 13-17 (15) mm wide, in all 23-31 (27) mm long. Flowers: hypanthium dull, verruculose, with grooves beneath each calyx lobe juncture; calyx lobes always 6, 14-15 (15) mm long, obtuse apex, inner surface sometimes tinged with red at base; petals absent; stamina1 filaments c. 19 mm long when fresh, entirely white; anthers dorsifixed, yellow; petals and stamens fall within hours of anthesis; ovary ± 10-locular; style c. 26 mm long, stigma fungiform, c. 2 mm wide when fresh. Berry globose, 20-21 (21) mm long, 42-53 (47) mm wide, pericarp leathery, satiny, without ribs; calyx persistent, hypanthium flat-expanded, 26-27 (27) mm wide, 27- 30 (29) mm from calyx base to lobe apex adpressed to pericarp, 18-20 (19) mm long. Seeds numerous, rounded irregular, c. 5 mm long. P h e n o l o g y. Flowers were observed and collected from March to October, and fruits from April to October. D i s t r i b u t i o n . This species is uncommon, but occurs from China through Malaysia to New Guinea. In Papua New Guinea it is located in several sites from Daru Island (9º05'S, 143º00'E) to the Gulf of Papua (7º05'S, 144º00'E) and to Milne Bay (10º03'S, 150º31' E). It is unknown in Australia. Fig. 1. E c o 1 o g y . The species occurs on the landward edge of mangrove swamps in brackish water and muddy soil. N o t e s. The acceptance of petalous variability for S. alba removed the only real problem in S. ovara determinations. Therefore, without the pseudo-reliance on this character, more diagnostic characters were used. Sonneratia ovara was distinguished by several attributes including: the presence of a fine verruculose texture on the calyx surface; leaf shape and texture; absence of a leaf mucronate apex; and, habitat. Therefore, while the species was found infrequently in this region, it had a very different morphology from other taxa. IDENTIFICATION LIST OF COLLECTIONS (Anonymous collections were omitted) Sonneratia 1. alba J. Smith 2. caseolaris (L.) Engl. 3. x gulngai N.C. Duke (= alba x caseolaris) 3a. alba x gulngai 4 . lanceolata Blume 4a. alba x lanceolata 5. ovata Backer N.C. Duke & B.R. Jackes: Revision of Sonneratia in Australasia 301 Adair 203: 1 - AIMS series SO, 156, 157, 180, 203, 213: 1; 235: 4; 242, 244, 255-257: 1; 262: 2; 263: 1; 264: 3; 270, 272, 276, 277: 1; 278: 2; 279, 294: 3; 296-299, 306: 2; 307: 3; 308, 313-317: 1; 318, 319: 2; 329, 330: 3; 331, 332: 1; 339: 2; 345, 348: 1; 362: 3a; 363: 3; 365: 1; 366: 3a; 368: 1; 369: 3; 371, 372: 1; 377, 380, 381: 4; 387-389, 393, 396-401, 406-420, 422, 432: 1; 434: 4; 435,436: 1; 437: 4; 445, 482, 497: 1; 510, 511: 2; 524: 4; 530, 535, 545, 546: 1; 547: 3; 548: 2; 549, 563, 577, 580: 1; 593: 3; 604: 1; 608-613: 2; 614, 615: 3; 616: 1; 617: 4; 618,620: 2; 629,631, 632: 3; 633, 634: 1; 636: 4; 642: 1; 644: 4; 645, 646: 1; 651,652: 4; 653: 1; 654, 662-664: 4; 665, 666, 672, 673, 679: 1; 686: 2; 688: 1; 689: 2; 690: 1; 692: 3; 693, 696, 697: 2; 698: 3; 699: 1; 700: 3; 701: 1; 702: 3a; 707, 721: 4; 723: 1; 731: 4; 734: 1; 735: 4; 736: 2; 737, 738: 3; 739: 1; 740,741: 4; 742: 1; 743-745: 3a; 746, 749, 750: 4; 752-755: 1; 756, 761, 770-772: 4; 775: 1; 776, 779: 4; 783, 784: 2; 785: 3; 799, 800, 805, 812-820: 1; 825: 4; 826: 1; 827: 2; 828: 3; 829: 4; 830: 3; 831: 4; 832: 2; 833-837: 1; 838: 2; 845-847: 1; 849: 1. Blake 16953, 18838: 1 - Boyland 353: 1 - Brass 790, 891: 1; 980: 4; 1350: 1; 1617, 5775: 4; 6264: 5; 6268: 1; 7968, 8204, 8633: 4; 21923: 5; 28160: 1; 33818: 2 - BRI series 33205, 33207, 63632, 63636, 63638: 1; 63645, 63648, 63649: 2; 63651: 1; 87411: 2; 140263: 4; 230424: 1 - BSIP series 2600, 2628: 1; 5676: 2; 6544, 7095, 10435: 1; 10635: 2; 10894, 13322: 1; 16575, 17183: 3; 18002, 19269: 1 - BW series 28: 2; 2445, 3198: 1; 3410: 2; 3575: 1; 4241.4259: 2; 4586: 1; 5249, 5346, 7569.7591: 2; 8576.9814: 1; 14861: 2 Bymes NB501, NB895: 1. Cameron 2409: 1 - Clarkson 2211, 2313: 1 -Craven 812: 1. Darbyshire 798: 1 - DNA series 9869, 9875, 9876: 4; 13666: 1; 13674, 13679, 13689, 15227: 4; 15228: 4a. Flecker 7787: 2; 8352: 1- Franc 2295: 1. Gill M25: 1. Hanson 1539: 1 - T.G. Hartley 10254, 10254B, 10354A: 2 - Henshall 813: 1 - Hoogland 4193, 4419: 2 - Hyland 8896: 1; 9348: 2. Jones 4026: 1 - Juncosa 23x8lh: 5. Kajewski 442: 2 - Kirina 4, 12: 1. Liitjeharms 5399: 1. McDonald 1632: 1 - KcKean B49: 1 - McKee 1701: 4a - Moi 115: 1 - Must 882: 1. NGF series 1382: 1; 1597: 2; 1833, 4105: 1; 4118: 4; 5026: 1; 5045: 4; 6543: 2; 7199: 1; 8012: 4; 8022: 5; 8040: 4; 8683: 2; 9104: 4; 11107: 1; 15307, 16319, 17198: 2; 17688: 1; 18056: 2; 18422, 22171: 1; 22303: 2; 22434: 1; 24053, 25236: 2; 27259: 1; 29425: 2; 29856: 1; 30487: 2; 30781: 1; 37554, 39407: 2; 41115: 1; 41161: 3; 41215: 1; 41496: 2; 43323: 1; 43740: 2; 43797, 43814: 1; 43821: 5; 48638: 2; 49443, 49446: 1; 52139: 2; 59104: 4; 59295: 2; 61201: 4; 61219: 1; 62170: 4; 65553, 66068, 70530: 1. Paijmans 1796: 1; 1878: 4 - Panoff 264: 1 - Parker 690: 1 - PERTH series 2817/B, 2849/C: 1 Pullen 643 1: 4. QRS series 41350, 41351: 1. Rankin 1169, 1817, 2218: 1 - Rau S4: 5 - Rijkers 14800: 1 - Robinson 9285: 1 - van Royen 3100, 5429: 1. Saunders 161,968: 2 - Scarth-Johnson 949A: 2 - Schodde 4444: 4 - Sharpe 20: 1 - Smith 4009, 10670, 12506: 1 - Stocker 1339: 1 - Stoddart 4640, 4773, 4839, 4852, 4905, 5043, 5104: 1 Stone 10485: 1. Thorn 4161, 4167: 1 - Thome 37898: 1. UPNG series 57: 1; 804: 1. White 131: 1; 689: 4. 302 BLUMEA - VOL. 32, No. 2,1987 INDEX Numbers refer to the specific taxon codes. Synonyms are in italics. Aubletia caseolaris 2 Blatti acide 2 alba 1 caseolaris 2 pagatpat 2 Chiratia leucantha 1 ‘Mangium caseolare’ ‘album’ 1 ‘rubrum’ 2 ‘Pagapate’ 2 Rhizophora caseolaris 1, 2 Sonneratia acida 1, 2, 4 var. mossambicensis 1 var. mucronata 2 alba 1 View publication stats (Sonneratia) alba 2, 4, 5 var. iriomotensis 1 alba x S. caseolaris 3 alba x S. gulngai 3a alba x S. lanceolata 4a caseolaris 2 caseolaris 1, 4 evenia 2 griffithii 1 x gulngai 3 iriomotensis 1 lanceolata 4 mossambicensis 1 neglecta 2 obovata 2 ovalis 2 ovata 5 pagatpat 1.2 rubra 2