Florence Grovida Gardening

Alseuosmiaceae

Alseuosmiaceae Airy Shaw, Kew Bull. 18: 249 (1965). Platyspermatiaceae Doweld (2001).

J. Karehed

Shrubs, sometimes creeping or epiphytic sub-shrubs. Leaves alternate, sub-opposite or in pseudo-whorls, simple, entire or serrate, estipulate. Multicellular, uniseriate hairs present in leaf axils, rarely also on leaves and stems, and erect unicellular hairs sometimes present on both leaves and stems. Flowers regular, hermaphroditic or functionally unisexual, tetra- or pentamerous, rarely up to hexamerous, fascicled in the leaf axils or terminally, sometimes solitary, rarely racemose. Calyx with free lobes. Corolla funnel-shaped or cam-panulate to urn-shaped, sympetalous, sometimes only shortly tubular (Platyspermation), with more or less lobed petal wings (not Platyspermation), sometimes carunculate inside the lobes. Aestivation valvate. Stamens isomerous, attached to the corolla, sometimes inserted at the very base of the corolla tube, alternating with the corolla lobes, sometimes sessile (Platyspermation). Anthers introrse, longitudinally dehiscent. Disc present or absent. Style single with capitate or discoid stigma, often more or less bi- or trilobed. Ovary inferior or sometimes semi-inferior, with two to three locules, each containing two to many anatropous ovules. Placentation axile. Fruits berries or (Platysper-mation) capsules with one to several seeds with minute embryo and copious endosperm.

Ten species classified into five genera in eastern Australia, New Zealand, New Caledonia and New Guinea.

Vegetative Morphology. Alseuosmiaceae are shrubs, ranging from the creeping or epiphytic sub-shrubs of Wittsteinia to the sometimes 6-m-tall Periomphale. The simple leaves are either entire or serrate, lack stipules, and are alternate, sub-opposite, or in pseudo-whorls. Venation is pinnate, very faint in Crispiloba. Leaves size varies between 3 and 20 cm.

Vegetative Anatomy (Gardner 1976; Dickison 1986, 1989; Platyspermation not investigated).

Rusty brown, multicellular uniseriate hairs are present in the leaf axils. In Platyspermation uniseriate hairs with persistent reddish bases are found also on other parts of the plant (Stevens 2001). Erect unicellular hairs may be present on both leaves and stems. The latter type is especially abundant in Wittsteinia vacciniacea, forming an indumentum on stems, petioles and basal portions of the leaves. In contrast, Crispiloba and Periomphale are (almost) completely devoid of this hair type.

The leaf epidermis is thin and consists of one cell layer. In transectional view, the epidermal cells are square or rectangular. Their anticlinal walls are usually undulate and deeply lobed in surface view. The anomocytic stomata are level with the epidermis and have prominent outer cuticular ledges. The one-layered palisade cells are poorly differentiated from the lacunose spongy mesophyll. In Crispi-loba, numerous long sclereids with thick lignified walls form an "interwoven mass... that permeates the mesophyll" (Dickison 1989). More or less rod-shaped sclereids may be found around the midvein in Periomphale. The latter are lobed or armed, have thinner walls, and are not as elongate as the scle-reids of Crispiloba. Sclerenchyma is present either as bundle sheaths or as idioblastic sclereids in the leaves of all taxa except Wittsteinia vacciniacea, which completely lacks foliar sclerenchyma.

The nodes are trilacunar with three traces. In Alseuosmia the traces fuse about halfway up the petiole, in Wittsteinia papuana and Periomphale they fuse at the base of the lamina, and in Crispiloba and W. vacciniacea they remain separated throughout the petiole and into the lamina. Fibrous bundle caps develop distally in the petiole, except in W. vacciniacea.

All genera have an endodermis with prominent Casparian banding present in young stems and surrounding the petiolar vascular bundles in all genera but Periomphale. Calcium oxalate crystals have not been detected.

The wood has very faint or no growth rings. The narrow vessels are mostly solitary, sometimes in radial multiples, or rarely clustered. The mean number of bars of the scalariform perforation plates ranges from 20 (Crispiloba) to 43 (Periomphale). Intervessel pits commonly have circular borders and are opposite and transitional to alternate, or sometimes scalariform. In Alseuosmia and Crispiloba the vessel elements have fine helical thickenings. The imperforate elements are living, store starch at maturity, and have indistinctly bordered or simple pits. Both septate and non-septate fibres are present in all genera. Periomphale has tall (> 1.5 cm) and wide, multiseriate rays, whereas the rays of Crispiloba are heterocellular, shorter and narrower. No rays are present in the other genera. Axial parenchyma is very sparse or absent.

Inflorescences. Alseuosmia and Wittsteinia have few-flowered fascicles or solitary flowers in the leaf axils. In Periomphale the inflorescences are predominantly terminal. Commonly, they consist of fascicled flowers but racemes are sometimes found. Terminal, umbel-like, mostly pedunculate inflorescences of one to five flowers are found in Crispiloba. Platyspermation has few-flowered inflorescences with inclined flowers. Pedicels have few bracts which are very early caducous in Periomphale.

Flower M0Rph0L0gy. The flowers are regular with whorls of normally four or five flower parts; hexamerous flowers are sometimes encountered (septamerous flowers in Periomphale were reported by Baillon 1888). According to Tirel and Jeremie (1996), Periomphale has both hermaphroditic (perhaps functionally male) and functionally female flowers. In the other genera, the flowers are hermaphroditic. The calyx lobes are valvate, free, more or less triangular, and persistent or circumscissile-caducous (Alseuosmia). The corolla tube of Platyspermation is short whereas in the other genera the clearly sympetalous corolla is funnel-shaped or campanulate to urn-shaped. The colour of the corolla ranges from dull red in Alseuosmia macrophylla over various pale shades of pink, yellow and green, with or without red markings, to pure white in Crispiloba. The valvate corolla lobes have appendages, so-called petal wings, which are more or less lobed (Fig. 2A-E). In Crispiloba they are conspicuously fringed whereas in Platyspermation the corolla lobes lack evident petal wings but are papillate. Petal wings remi niscent of those in Alseuosmiaceae are also found in Goodeniaceae and Menyanthaceae (Gustafsson 1995). In Crispiloba, the base of the midrib on the inside of the lobes is fringed in a similar way. Also, there are irregular appendages at the throat of the corolla tube. In Wittsteinia and Periomphale, a caruncle at the base of the lobes forms a 'corona' which may cover the tube (Fig. 2D). The length of the tube varies from c. 0.5 cm (Wittsteinia papuana and small-flowered Periomphale) to 4.5 cm (Alseuosmia macrophylla and Crispiloba). The stamens alternate with the corolla lobes, and are inserted either in the throat of the corolla tube (Alseuosmia and Crispiloba) or at the very base of the tube. The introrse anthers open by longitudinal slits. In Platyspermation the anthers are sessile and extrorse (?), brown hairy below, and with a large flat connective appendage at the apex (van Steenis 1982). A disc is mostly present. It is very reduced or missing in Wittsteinia, and inhabited by parasitic insects in flowers of Periomphale (see below). The single style has a capitate or discoid, often bilobed stigma, sometimes trilobed in Wittsteinia vacciniacea. The ovary is inferior or sometimes initially semi-inferior in Periom-phale and two-locular; Wittsteinia vacciniacea commonly has three locules. Each locule contains two to many anatropous ovules with axile placentation, in Platyspermation the placenta is brown and very thick (van Steenis 1982). Besides the normal type of flowers, Tirel (1996) and Tirel and Jeremie (1996) described flowers inhabited by parasitic insects in Periomphale. These are globose or obconical, do not open, and are often borne on elongated pedicels (up to 6 cm long). The stamens and the style are frequently only weakly developed, as is the corolla, which is shed prematurely or withers. A disc is usually lacking and the ovules, if present, do not develop into seeds. Similar flowers are reportedly present also in Wittsteinia and Platyspermation (van Steenis 1984; Stevens 2001).

FLORaL Anatomy. The floral anatomy of Alseuosmia and Periomphale was investigated by Gardner (1976). According to him, the anatomical features are uniform within Alseuosmia, and essentially agree with those in Periomphale. Notably, at the top of the ovary the locules interconnect above the placental region for about 50-100 pm, due to the septum being transversely divided (in one examined flower of Periomphale, this resulted in parietal placentation of the uppermost ovule).

1. 2. Alseuosmiaceae. A Alseuosmia macrophylla, habit. B A. banksii, flower with opened corolla. C Crispiloba dis-perma, habit. D Periomphale balansae, female flower; longitudinal section of ovary. E Wittsteinia vacciniacea, habit. (Redrawn after A Hooker 1887, B Hooker 1853-1855, C van Steenis 1984, D Tirel and Jeremie 1996, E Mueller 1885)
2. 2. Alseuosmiaceae. A Alseuosmia macrophylla, habit. B A. banksii, flower with opened corolla. C Crispiloba dis-perma, habit. D Periomphale balansae, female flower; longitudinal section of ovary. E Wittsteinia vacciniacea, habit. (Redrawn after A Hooker 1887, B Hooker 1853-1855, C van Steenis 1984, D Tirel and Jeremie 1996, E Mueller 1885)
3. In Alseuosmia the anther wall consists of an epidermis, an endothecium with fibrous thickenings when mature, two middle layers and a one-layered tapetum. The tapetum cells are binucleate before meiosis in the pollen mother cells, remain in place during pollen development, and subsequently undergo nuclear fusion. The pollen is shed in the trinucleate condition. The ovules of Alseuosmia are unitegmic and tenuin-ucellate, and the embryo sac is eight-nucleate at maturity (Gardner 1976).
4. The diploid chromosome number of Alseuosmia is2n = 18 (Gardner 1976).
5. The 3-colporate pollen of Alseuosmia was studied by Erdtman (1952) who described the pollen grains as often angulaperturate, oblate spheroidal-prolate (longest axis45-50 pm). Sexine tegillate, slightly undulating, thicker than the nexine, sometimes with small fissures; ectosexine thicker than endosexine, the latter only faintly baculate. According to Hufford (1992) and Karehed et al. (1999), Alseuosmia pollen has well-developed columellae. The pollen of Periomphale is similar in appearance to that of Alseuosmia (Bortenschlager et al. 1966), as is that of Platyspermation (3-colpor(oid)ate, oblate, spheroidal, c. 31 x 36 pm and with the sexine thicker than the nexine; Erdtman 1952).

Fruit and Seed. The fruits of Alseuosmiaceae are two-locular berries, two- to three-locular in Wittsteinia, or two-locular capsules (Platyspermation). Their shape varies from globose (W vacciniacea) to narrowly ellipsoid. Each fruit contains one to many ellipsoid or ovoid, more or less compressed seeds with a brown to black testa. In Platyspermation, seeds are sculptured and have fine, inflated hairs on the margin (van Steenis 1982). The seed coat structure and the lignified exotesta cells of Alseuosmia have been described by Nemirovich-Danchenko and Lobova (1998).

Reproductive Biology. According to Gardner (1976), Alseuosmia pusilla is self-compatible, whereas A. macrophylla is an obligate outbreeder with an incompatibility mechanism operating at the stylar level.

1. In the leaves of Alseuosmia, Cambie and Parnell (1969) detected lupeol, lupenyl acetate, stigmasterol, stearic acid, and triterpene acetates. Also from Alseuosmia, Gardner (1976) reported the presence of a condensed tannin (leu-cocyanidin) and ellagitannin, together with simple phenols (quercetin, caffeic acid, kaempferol, and p-coumaric acid) and triterpenoid saponins. He could not detect either alkaloids or iridoids.
2. Alseuosmiaceae form a mono-phyletic group in Asterales, together with Argophyllaceae and Phellinaceae (Gustafsson et al. 1996; Backlund and Bremer 1997; Gustafsson and Bremer 1997; Kallersjo et al. 1998; Karehed et al. 1999; Lundberg and Bremer 2003).

Before the recent addition of Platyspermation (see below), several studies supported Alseuos-miaceae as a well-defined family (e.g. van Steenis 1984; Dickison 1986, 1989; Karehed et al. 1999). Already when Cunningham (1839) described Alseuosmia, he suggested it to be a distinct family, related to Cornaceae, Caprifoliaceae and Loran-thaceae. Alseuosmia was, however, mostly treated as a more or less aberrant member of Caprifoliaceae (e.g. Hooker 1873; Fritsch 1891), as were the two New Caledonian genera Memecylanthus and Pachydiscus (Schlechter 1906; Guillaumin 1948) now included in Periomphale (van Steenis 1978). Initially, Baillon (1888) placed Periomphale in Ges-neriaceae and, due to lack of detailed knowledge, the genus was mostly kept in that family until Airy Shaw (1965) realized its correct affinities. Airy Shaw (1965) also gave the first formal description of Alseuosmiaceae, indicating affinities with Escalloniaceae. Gardner (1976, 1978a) suggested Alseuosmiaceae to be most closely related to Argophyllaceae, then included in Escalloniaceae but now considered a separate family of Asterales (e.g. Kallersjo et al. 1998; Karehed et al. 1999). van Steenis (1978, 1984) added to the family the newly discovered Wittsteinia papuana and W. vac-ciniacea, formerly thought to belong in Ericaceae (von Mueller 1861; Bentham 1869; Drude 1889; Stevens 1971) or Epacridaceae (Burtt 1949), and Crispiloba, earlier misplaced in Rubiaceae (Moore 1917). van Steenis (1984) included Periomphale in Wittsteinia but Tirel (1996) and Tirel and Jeremie (1996) argued that it should be regarded as a distinct genus. Alseuosmiaceae obtained their present circumscription when molecular studies showed the little known genus Platyspermation to be the sister to Alseuosmiaceae (Lundberg and Bremer 2003). Here, Platyspermation is included in the family, rather than recognising a monotypic Platyspermataceae (Doweld 2001). Platyspermation was originally described as a member of Myrtaceae (Guillaumin 1950). Airy Shaw (in Willis 1973) placed it in Rutaceae, while a relationship to Escalloniaceae was suggested by Erdtman (1952), Schmid (1980) and van Steenis (1982).

Distribution and Habitats. Alseuosmia is found in lowland to montane forests in New Zealand (Gardner 1976). Crispiloba is endemic to rain forests of Queensland, Australia. On New Caledonia, Periomphale inhabits humid forests (Tirel and Jeremie 1996), as does Platyspermation which is also found in the maquis vegetation. Wittsteinia is a mountainous genus; the Australian W. vacciniacea grows in crevices of rocks and on rocky summits of Victorian mountains (Bentham 1869), and the Papua New Guinean W. papuana has been collected in woodland at 3,000 m (van Steenis 1978).

Key to the Genera

1. Corolla tube short; stamens sessile; fruit a capsule
2. Platyspermation

- Corolla tube well developed; stamens inserted in the corolla tube; fruit a berry 2

2. Corolla funnel-shaped or narrowly cylindrical; stamens inserted at the apex of the corolla tube 3

- Corolla campanulate to urn-shaped or narrowly barrel-shaped; stamens inserted at the base of the corolla tube 4

3. Leaves alternate, (sub)serrate; flowers fascicled or solitary in the leaf axils; calyx circumscissile-caducous; corolla funnel-shaped with ± lobed petal wings 1. Alseuosmia

- Leaves in pseudo-whorls, entire; flowers terminal; calyx persistent; corolla narrowly cylindrical with strongly fringed petal wings 2. Crispiloba

4. Shrubs; leaves entire; inflorescences terminal; corolla with entire petal wings 3. Periomphale

- Subshrubs; leaves serrate; inflorescences axile; corolla with lobed petal wings 4. Wittsteinia

Genera of ALseuosmiaceae

Alseuosmia A. Cunn., Ann. Nat. Hist. 2: 209 (1839); Merrett & Clarkson, N. Z. J. Bot. 38: 153-164 (2000), rev.

1. Leaves alternate, simple, serrate, sometimes subentire. Flowers fascicled in the leaf axils or solitary, regular, hermaphroditic, tetra- or pen-tamerous, rarely up to hexamerous. Corolla with a funnel-shaped tube and valvate lobes with more or less fringed petal wings. Stamens inserted at apex of corolla tube. Disc present. Stigma capitate or bilobed on elongate style. Ovary inferior, two-locular with two to many ovules in each locule. Fruit a two-locular berry. n = 9. Five species in New Zealand.
2. Crispiloba Steen. Fig. 2C

Crispiloba Steen., Blumea 29: 391 (1984).

1. Leaves in pseudo-whorls of 3-6, entire. Inflorescences terminal, predominantly pedunculate, umbel-like with (one) up to five flowers. Flowers (tetra-) pentamerous, with a salver-shaped corolla and a long, narrow, cylindrical tube. Corolla lobes fringed inside and with strongly fringed petal wings. Stamens inserted at the throat of the corolla tube. Ovary inferior with two locules, each containing 2-3 ovules. Fruit a berry with black, more or less planoconvex seeds. One species, C. disperma (S. Moore) Steen., Queensland, Australia.
2. Periomphale Baill. Fig. 2D

Periomphale Baill., Bull. Mens. Soc. Linn. Paris 1:731 (1888); Tirel & Jérémie, Fl. Nouvelle-Calédonie 20: 100-106 (1996), rev.

Memecylanthus Gilg & Schltr. (1906). Pachydiscus Gilg & Schltr. (1906).

Shrubs with flexuose branches. Leaves alternate, often sub-opposite or in pseudo-whorls at the tip of the branches, entire. Inflorescences predominantly terminal, rarely racemose or more commonly of (1)2-8(15)-fascicledflowers. Flowers tetra- or pen-tamerous (rarely hexamerous), hermaphroditic or functionally unisexual. Corolla urn-shaped to cam-panulate; corolla lobes carunculate inside. Stamens inserted at the very base of the corolla tube. Disc present, rarely only weakly developed. Ovary inferior, sometimes initially semi-inferior, with two locules, each containing 4-6 ovules. Fruit a sub-cylindric to ovoid-fusiform berry with more or less compressed, ellipsoid seeds. One species, P balansae Baill., endemic to New Caledonia.

4. Wittsteinia F. Muell. Fig. 2E

Subshrubs, creeping with ascending branches, W. papuana (Steen.) Steen. epiphytic. Leaves alternate or in pseudo-whorls, serrate. Inflorescences of one or two axillary flowers. Corolla more or less cam-panulate or narrowly barrel-shaped ( W papuana (Steen.) Steen.); corolla lobes carunculate inside (inconspicuously so in W. vacciniacea F. Muell.) and with lobed petal wings. Stamens inserted on the corolla tube at the very base. Ovary inferior, two-to three-locular, few ovules in each locule. Fruit a globose berry with ovoid seeds. Two species; one endemic to Victoria, Australia, and one to Papua New Guinea.

5. Platyspermation Guillaumin

Platyspermation Guillaumin, Acta Horti Gotob. 18: 253 (1950).

Shrub up to 5 m high. Leaves alternate, pseudo-whorled at the end of branches, with small dentations on the recurved margins, sclerophyl-lous. Inflorescences few-flowered with inclined flowers. Corolla with short tube. Stamens sessile. Anthers with brown hairs below. Ovary inferior, two-locular, with few ovules on thick, brown placenta. Fruit a two-locular capsule with few, flat seeds. One species, P. crassifolium Guillaumin, endemic to New Caledonia.

Selected Bibliography

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Backlund, A., Bremer, B. 1997. Phylogeny of the Asteridae s. str. based on rbcL sequences, with particular reference to the Dipsacales. Pl. Syst. Evol. 207: 225-254.

Baillon, H.E. 1888. Observations sur les Gesnériacées. Bull. Mens. Soc. Linn. Paris 1: 731-732.

Bentham, G. 1869. Flora Australiensis, vol. IV. London: Lovell Reeve.

Bortenschlager, S., Erdtman, G., Praglowski, J. 1966. Pollenmorphologische Notizen über einige Blütenpflanzen incertae sedis. Bot. Notiser 119: 160-168.

Burtt, B.L. 1949. Studies in the Ericales, IX. The taxonomic position of Wittsteinia. Kew Bull. 3: 493-495.

Cambie, R.C., Parnell, J.C. 1969. A New Zealand phytochem-ical survey, part 7. Constituents of some dicotyledons. N. Z. J. Sci. 12: 453-466.

Cunningham, A. 1839. Florae Insularum Novae Zelandiae Precursor; or a specimen of the botany of the islands of New Zealand. Genus Corneis affine. Ann. Nat. Hist. 2: 209-210.

Dickison, W.C. 1986. Wood anatomy and affinities of the Alseuosmiaceae. Syst. Bot. 11: 214-221.

Dickison, W.C. 1989. Stem and leaf anatomy of the Alseu-osmiaceae. Aliso 12: 567-578.

Doweld, A.B. 2001. Prosyllabus Tracheophytorum. Tenta-men Systematis Plantarum Vascularium (Tracheophy-torum). Moscow: GEOS.

Drude, O. 1889. Ericaceae. In: Engler, A., Prantl, K., Die natürlichen Pflanzenfamilien, IV, 1. Leipzig: W. Engelmann, pp. 15-65.

Erdtman, G. 1952. Pollen morphology and plant taxonomy. Angiosperms. Stockholm: Almqvist & Wiksell.

Fritsch, K. 1891. Caprifoliaceae. In: Engler, A., Prantl, K., Die natürlichen Pflanzenfamilien, IV, 4. Leipzig: W. Engelmann, pp. 156-169.

Gardner, R.O. 1976. Studies in the Alseuosmiaceae. Ph.D. Thesis, University of Auckland, Auckland, New Zealand.

Gardner, R.O. 1978a. Systematic notes on the Alseuosmi-aceae. Blumea 24: 138-142.

Gardner, R.O. 1978b. The species of Alseuosmia (Alseuosmiaceae). N. Z. J. Bot. 16: 271-277.

Guillaumin, A. 1948. Flore (analytique et synoptique) de la Nouvelle Calédonie, Phanerogams. Paris: Office de la Recherche Scientifique Coloniale.

Guillaumin, A. 1950. Plantae Neocaledonicae a C. Skotts-berg a. 1949 lectae (96ème contribution à la flore de la Nouvelle-Calédonie). Acta Horti Gotob. 18: 247265

Gustafsson, M.H.G. 1995. Petal venation in Asterales and related orders. Bot. J. Linn. Soc. 118: 1-18.

Gustafsson, M.H.G., Bremer, K. 1997. The circumscription and systematic position of Carpodetaceae. Austral. Syst. Bot. 10: 855-862.

Gustafsson, M.H.G., Backlund, A., Bremer, B. 1996. Phy-logeny of the Asterales sensu lato based on rbcL sequences with particular reference to the Goodeniaceae. Pl. Syst. Evol. 199: 217-242.

Hooker, J.D. 1853-1855. The botany of the Antarctic voyage of H.M. Discovery ships Erebus and Terror in the years 1839-43. II. Flora Novae-Zelandiae. London: Lovell Reeve.

Hooker, J.D. 1873. Caprifoliaceae. In: Bentham, G., Hooker, J.D., Genera Plantarum, vol. II, part 1. London: Lovell Reeve, pp. 1-7.

Hooker, J.D. 1887. Curtis's Botanical Magazine, vol. CXIII. London: Lovell Reeve.

Hufford, L. 1992. Rosidae and their relationships to other nonmagnoliid dicotyledons: a phylogenetic analysis using morphological and chemical data. Ann. Missouri Bot. Gard. 79:218-248.

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Karehed, J., Lundberg, J., Bremer, B., Bremer, K. 1999. Evolution of the Australasian families Alseuosmiaceae, Argophyllaceae, and Phellinaceae. Syst. Bot. 24: 660682.

Lundberg, J., Bremer, K. 2003. A phylogenetic study of the order Asterales using one morphological and three molecular data sets. Intl J. Pl. Sci. 164: 553-578.

Moore, S. 1917. A contribution to the phyto-geography of Bellenden-Ker. II. Systematic account. Phanerogams. J. Bot. 55: 302-309.

Mueller, F. von 1861. Fragmenta Phytographiae Australiae, vol. II. Melbourne: Auctoritate Gubern. Coloniae Vic-toriae.

Mueller, F. von 1885. Key to the system of Victorian plants. Melbourne: Government Printer.

Nemirovich-Danchenko, E.N., Lobova, T.A. 1998. The seed coat structure in some representatives of the order Hydrangeales. Bot. Zhurn. (Moscow & Leningrad) 83: 1-9.

Schlechter, R. 1906. Beiträge zur Kenntnis der Flora von

Neu-Kaledonien. Bot. Jahrb. Syst. 39: 1-274. Schmid, R. 1980. Comparative anatomy and morphology of Psiloxylon and Heteropyxis, and the subfamilial and tribal classification of Myrtaceae. Taxon 29: 559-595. Stevens, P.F. 1971. A classification of the Ericales: subfamilies and tribes. Bot. J. Linn. Soc. 64: 1-53. Stevens, P.F. 2001 (onwards). Angiosperm Phylogeny website, version 4, May 2003. http://www.mobot.org/ MOBOT/research/APweb/ Tirel, C. 1996. Rétablissement de Periomphale Baill. (Alseu-osmiaceae), genre endémique de Nouvelle-Calédonie. Bull. Mus. Natl Hist. Nat. B, Adansonia 18: 155-160. Tirel, C., Jérémie, J. 1996. Alseuosmiaceae. In: Morat, P. (ed.) Flore de la Nouvelle-Calédonie, vol. 20. Paris: Muséum National d'Histoire Naturelle, pp. 100-106. van Steenis, C.G.G.J. 1978. The genus Periomphale in New

Guinea (Caprifoliaceae). Blumea 24: 480-481. van Steenis, C.G.G.J. 1982. 157. Preliminary note on the taxonomic disposition of Platyspermation Guillaumin (Myrtaceae) from New Caledonia. In: van Stee-nis, C.G.G.J., Veldkamp, J.F., Miscellaneous botanical notes XXVI. Reinwardtia 10: 21-26. van Steenis, C.G.G.J. 1984. A synopsis of Alseuosmiaceae in New Zealand, New Caledonia, Australia, and New Guinea. Blumea 29: 387-394. Willis, J.C. 1973. A dictionary of the flowering plants and ferns, ed. 8, revised by H.K. Airy Shaw. Cambridge: University Press.

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