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World Checklist And Bibliography Of Euphorbiaceae Columnar

All species of Ficus (Moraceae; refs. 1–3) and Yucca (Agavaceae; refs. 4 and 5) are pollinated exclusively by obligate seed-parasitic wasps and moths, respectively. Among 220,000 angiosperm species, however, such obligate pollination mutualisms between plants and seed-parasitic insects are rare, probably because incipient mutualisms are dominated by abundant pollinators that visit flowers for nectar or pollen (6). Although several plant species have recently been found to be pollinated by seed-parasitic moths (6, 7) and flies (8), such mutualism is restricted to a few plant species in each genus. Accordingly, obligate plant–pollinator mutualism accompanied by reciprocal diversification is surprisingly rare and confirmed only in the fig/fig wasp (9, 10) and yucca/yucca moth (11, 12) systems.

Glochidion, a monoecious tree genus of Euphorbiaceae (recently treated as a separate family, Phyllanthaceae), has minute apetalous female flowers with highly specialized styles (13, 14). The genus consists of 318 species ranging from tropical Asia to Australia, Polynesia, and Madagascar (15). Although its pollination system was unknown, Glochidion trees usually bear many fruits, most of which are infested by small moth larvae. Our field observations revealed that flowers were actively pollinated by seed-parasitic gracillariid moths that oviposited into styles. Here we present evidence of a widespread obligate pollination mutualism between Glochidion trees and their seed-parasitic moths.

Results and Discussion

Glochidion flowers are dimorphic, consisting of a pedunculate male flower with unfolded sepals and connate ellipsoid stamens, with a sessile or shortly pedunculate female flower composed of folded sepals and a united columnar style. The style has a narrow pit at its lobed tip. The inner surface of the pit is the stigma. This cryptic stigma is unlikely to be pollinated by wind or ordinary insect visitors. We therefore made field observations of the pollination of three Glochidion species in Japan.

Trees of G. acuminatum have male and female flowers in separate axillary clusters on each branch (Fig. 1a). Male flowers are aggregated at the base, and the female flowers are at the apical part of each branch. Female flowers are generally six-ovuled. Neither male nor female flowers secrete nectar. In the daytime, the flowers were rarely visited by insects, whereas various insects passed by the inflorescence.

Figure 1

Flowers and pollinators of G. acuminatum (af), G. zeylanicum (gj), and G. obovatum (km). (a) A branch bearing many male and female flowers, one of which is visited by a moth, Epicephala sp. 1. (b) A female moth visiting a male flower. (c) Pollinated stigma. (d) An ovipositing Epicephala moth. (e) Pollen-loaded proboscis of a female moth. (f) A fruit with two seeds infested by a moth larva, which had emerged from the hole. (g) An Epicephala sp. 2 moth actively pollinating a female flower. (h) An ovipositing Epicephala moth. (i) Dorsal view of a female flower, showing the pollinated cryptic stigma. (j) Cross section of a female flower with two eggs (shown by arrows). (k) An ovipositing Epicephala sp. 3 moth. (l) Pollinated stigma. (m) Lateral view of a female moth, whose proboscis is loaded with pollen. [Scale bars, 1 mm (except in e).]

However, beginning in the evening and continuing until midnight, the flowers were visited frequently by a gracillariid moth, Epicephala sp. 1. Female moths visited male flowers to collect pollen by inserting their proboscis into the anthers (Fig. 1b). After flight migration among trees or within a tree, female moths visited female flowers. All female Epicephala moths netted around the Glochidion trees had G. acuminatum pollen grains attached to their ciliated proboscides (mean pollen number 88 ± 66, n = 25; Fig. 1e), whereas proboscides of males had no pollen grains (0 ± 0, n = 19). The behavior of the female moths on female flowers was remarkable. Visiting a cluster of female flowers, a female uncoiled its proboscis, deposited the pollen grains onto the cryptic stigma, and bent its abdomen to insert its long ovipositor into the stylar pit (Fig. 1d). The time spent in pollination and oviposition was 46 ± 10 s (n = 14). The female walked along the branch, visiting each female flower sequentially and repeating the stereotypic behavior.

We examined pollen attachment and moth eggs in female flowers by dissecting the styles under a microscope. Normally, a female laid an egg in a flower just above the ovules at the base of each oviposited style, and an average female flower received 1.71 eggs (Table 1). Oviposited flowers were consistently pollinated (Fig. 1c), whereas unoviposited flowers were very rarely pollinated (Fig. 2). The mean number of pollen grains deposited on an oviposited style was 7.8 ± 4.8, significantly greater than that of an unoviposited style (1.2 ± 2.4, Mann–Whitney U test; U = 154, P < 0.0001). Although few, the pollen grains attached to each style were enough to fertilize all six ovules in an ovary.

Figure 2

Frequency distributions of the number of pollen grains attached to oviposited (solid) and unoviposited (shaded) stigmas of three Glochidion species (acuminatum, zeylanicum, and obovatum). The typical ovipositing postures and oviposited eggs of each Epicephala moth species are shown in each Inset.

Active pollination and the oviposition into styles by gracillariid moths was also observed on G. zeylanicum and G. obovatum, which have different style structures (Figs. 1 and 2). G. zeylanicum has ovoid bud-like female flowers whose styles are almost completely enclosed by sepals (Fig. 1i). The flowers have small openings at the apical tip that lead to the narrow stigma pit of the fused styles. Female flowers were visited at night by Epicephala sp. 2, which actively pollinated the female flower (Fig. 1g), inserted its long ovipositor into the narrow stigma pit (Fig. 1h), and laid an egg (Fig. 1j). Female flowers of G. obovatum are columnar like those of G. acuminatum, but differ in having distinctly swollen ovaries. At night the flowers were actively pollinated by Epicephala sp. 3, which inserted its abdomen between the style and calyx and laid an egg into the locules directly through the ovary wall, rather than through the stigma (Fig. 1km).

Fertilized ovules began to develop, whereas unpollinated female flowers abscised shortly thereafter. The hatched moth larva bored into the ovary and consumed a few developing seeds within a fruit. In G. acuminatum, a larva usually consumed two seeds to complete larval growth and escaped from the fruit to pupate on the litter (Fig. 1f). The life cycles of both the plant and its pollinator moth are inseparably linked (Fig. 3). Seed destruction was caused mostly by Epicephala moths, but nonpollinating seed-parasitic moths of Pyralidae and Tortricidae also infested seeds. On average, one fruit had 6.09 ovules, of which 1.80 were infested by moth larvae, 3.33 were intact, and 0.91 were sterile or aborted. The overall outcomes were similar among the three Glochidion species (Table 1).

Figure 3

Life cycles of G. acuminatum (broken arrows) and its pollinator moth, Epicephala sp. 1 (solid arrows). a, Branch bearing male and female flowers; b, male flower; c, female moth collecting pollen on a male flower; d, female flower; e, female moth depositing pollen on stigma; f, moth laying an egg into a style; g, moth larva infesting seeds; h, fruit with intact and infested seeds; i, moth puparium with an exuvia; j, emerged moth.

Surveying other Glochidion species, we found that all six species harbored an individual, undescribed, seed-parasitic Epicephala species that could be distinguished by its genitalic morphology. Host-specificity of the moths was not surprising because several Glochidion species often co-occur at our study sites without apparent hybridization. We confirmed host-specificity of the moths by investigating nucleotide sequence variation within 1,325 bp of the mitochondrial cytochrome oxidase subunit I gene (COI) among Epicephala. Pairwise sequence divergence between Epicephala moths reared from different Glochidion species was 3.1–9.1% (mean 7.3%), whereas sequence divergence between moths reared from the same Glochidion species in different locations was <0.5%.

These findings suggest that at least three Glochidion species are pollinated by a species-specific seed-parasitic Epicephala species, at the cost of infested seeds. It is notable that the female moth has an exceptionally long ovipositor to insert an egg into a style and a ciliated proboscis to collect pollen. The moth actively pollinated flowers, similar to fig wasps and yucca moths. The seed-parasitic habit of Epicephala is unique in Gracillariidae, most species of which are leaf miners. Other Epicephala larvae infest Phyllanthus seeds and Caesalpinia flower buds (17).

Twelve additional Glochidion species in New Caledonia, Fiji, Australia, Malaysia, and Myanmar, all had traces of limited seed infestation by the moths. Thus, GlochidionEpicephala mutualism may be widespread among the 318 known Glochidion species, whereas Epicephala has not yet been found in Madagascar. This mutualism shares many characteristics with fig/fig wasp and yucca/yucca moth mutualisms, because the reward for the pollinator consists of developing ovules or seeds. Furthermore, the sister groups of these pollinator taxa are endophytic herbivores and the pollinators are females with elongate ovipositors (1–5). Outstanding diversification has occurred only in Ficus (700 spp.) and Glochidion, both of which are tropical monoecious or gynodioecious woody plants that have highly specialized styles into which small pollinating insects oviposit (3).

In the Malay Archipelago, Glochidion is the largest genus (150 spp.) of Euphorbiaceae (15), and the principal species diagnostic characteristic is the structure of the style (13, 14). Because pollinating moths oviposit into styles by using diverse and specific methods, the length of their ovipositor and their oviposition behavior are crucial for such specialization. Thus, the specialized structure of the Glochidion style and the specialized oviposition behavior of the moths may well serve as barriers against both Glochidion hybridization and host-shift by the moths. In turn, plant speciation based on these traits provides a selective basis for speciation and high diversity. The coevolution of species is one of the major processes contributing to the earth's biodiversity (18, 19), and the GlochidionEpicephala relationship will provide a model system for comparative analyses of coevolutionary processes and mutualistic interactions.

Table 1

Comparison of moth pollination, oviposition, and seed infestation among three Glochidion species

Acknowledgments

We thank D. W. Roubik and S. Sakai for critical reading of the manuscript; T. Kumata for taxonomical suggestion on the moth; T. Terachi, T. Sota, and T. Sugiyama for technical support; and A. Naiki and H. Samejima for plant materials. This work was supported by Grant-in-Aid for Scientific Research 02440217 from the Japan Ministry of Education, Science, and Culture.

Footnotes

  • ↵* To whom correspondence should be addressed. E-mail: kato{at}bio.h.kyoto-u.ac.jp.

  • This paper was submitted directly (Track II) to the PNAS office.

  • Data deposition: The sequences reported in this paper have been deposited in the GenBank database (accession nos. AY221964–AY221981).

  • Received November 25, 2002.
  • Copyright © 2003, The National Academy of Sciences

References

Abstract

Recent molecular phylogenetic studies and reevaluation of morphological characters have led to the inclusion of Glochidion within a broader delimitation of Phyllanthus. It is necessary for preparation of the Vascular Flora of the Marquesas Islands to make new combinations for the Marquesan species. We also provide the relevant combinations and listing of all of the currently accepted species of Phyllanthus on Pacific oceanic islands for a total of 69 native species in oceanic Pacific islands. Glochidion tooviianum J. Florenceis here placed into synonymy of Phyllanthus marchionicus (F. Br.) W. L. Wagner & Lorence based on new assessment of recently collected specimens from Nuku Hiva. Glochidion excorticans Fosberg var. calvum Fosberg is placed into synonomy of Phyllanthus ponapense (Hosokawa) W. L. Wagner & Lorenceand Glochidion puberulum Hosokawa and Glochidion excorticans Fosberg are placed in synonymy of Phyllanthus senyavinianus (Glassman)W. L. Wagner & Lorence based on new study of all Micronesian specimens available to us. No infraspecific taxa are recognized within Phyllanthus pacificus of the Marquesas Islands. Species already with valid names in Phyllanthus are also listed for completeness and convenience. Brief distributional comments are given for each species. We propose new names for species for which a new combination is not possible: Phyllanthus florencei W. L. Wagner & Lorence, nom. nov., Phyllanthus mariannensis W.L. Wagner & Lorence, nom. nov., Phyllanthus otobedii W. L. Wagner & Lorence, Phyllanthus raiateaensis W. L. Wagner & Lorence, Phyllanthus st-johnii W. L. Wagner & Lorence, nom. nov., and Phyllanthus vitilevuensis W.L. Wagner & Lorence, nom. nov. We provide information for four additional naturalized species within the region (Phyllanthus amarus, Phyllanthus debilis, Phyllanthus tenellus, and Phyllanthus urinaria). The name Glochidion ramiflorum widely applied to Pacific island populations is here considered to be a species further west in the Pacific with all of the oceanic species here referred to several regional species.

Keywords: Caroline Islands, Fiji, Glochidion, Marquesas Islands, Micronesia, Pacific, Phyllanthaceae, Phyllanthus

Introduction

Recent molecular phylogenetic studies have greatly advanced the understanding of relationships in the family Phyllanthaceae (a segregate from Euphorbiaceaes. l.) based on chloroplast and nuclear DNA sequence data (Wurdack et al. 2004; Samuel et al. 2005; Kathriarachchi et al. 2005, 2006). Among the tribes in Phyllanthaceae (Hoffmann et al. 2006), tribe Phyllantheae is the largest natural group and accounts for more than half of the 2000 species in the family (Kathriarachchi et al. 2006; Hoffmann et al. 2006). One of the most taxonomically difficult groups in the family is Phyllanthus L.and related genera, species of which have small unisexual flowers and an often confusingly similar habit in unrelated groups. Historically generic circumscriptions in Phyllantheae have undergone substantial fluctuation, and the definition of natural groups is still unclear in many parts of the tribe (Hoffmann et al., 2006). The recent molecular studies of relationships within the family, with special emphasis on the large genus Phyllanthus, confirm paraphyly of Phyllanthus in its traditional circumscription with Breynia J. R. Forst. & G. Forst., Glochidion J. R. Forst. & G. Forst., Reverchonia A. Gray, and Sauropus Blumeembedded within Phyllanthus. These results led the research groups working on Phyllanthaceae (Hoffmann et al. 2006; Kathriarachchi et al. 2006) to conclude that the embedded genera should be submerged into a broadened Phyllanthus rather than further generic segregation, which would create a series of highly technical genera distinguishable only by specialists. Distinguishing morphological characters among these five taxa are not clear-cut or comparable, and thus recognition of equivalent clades at generic level would exacerbate the problem of understanding the evolution of this large and widespread lineage (Hoffmann et al. 2006). The enlarged Phyllanthus comprises nearly 1300 species with the inclusion of Breynia, Glochidion, Reverchonia, Phyllanthodendron Hemsl., and Sauropus. It is not surprising that the vast morphological diversity of Phyllanthus prior to adding the embedded taxa would readily accommodate them (Hoffmann et al. 2006). Unfortunately, the broadening of Phyllanthus necessitates a considerable number of new combinations, but nevertheless this seems to be the best approach over the long term in the interest of developing a classification that facilitates understanding of the evolution and relationships in the overall group.

Our work has focused on the oceanic Pacific and specifically we are currently completing investigation of the flora of the Marquesas Islands as part of a collaboration between National Tropical Botanical Garden (NTBG), Smithsonian Institution (SI), and the Délégation à la Recherche, Papeete, Tahiti (French Polynesia). We do not believe it is wise to make only the necessary combinations for just the Marquesan species previously placed in Glochidion so we here provide a synopsis of all of the Pacific oceanic island species (Fiji and Caroline Islands eastward across the Pacific). We also include all of the species currently recognized within Phyllanthus for completeness. We have not made an exhaustive study of the taxonomy of these species, but accept for the most part previous taxonomic conclusions. The Pacific group of species is in serious need of comprehensive analysis as previous work has focused on localized areas with few exceptions. Smith (1981) and Florence (1997) have made good regional taxonomic studies, but both point out problems that need a broader geographical perspective to solve. The Micronesian species are in particular need of comprehensive study and have received only partial treatments, either of part of the region (Hosokawa 1935) or more superficial descriptive work (Fosberg and Oliver 1991) without consideration of all species previously described. We have made some changes to the taxonomy of these species, but an in depth study is required to gain a more solid understanding of the diversity within Micronesia. Webster (1986) also made a partial study in the region, primarily of southwestern Pacific species, and Croizat (1943) contributed a useful analysis of several species. We have made extensive use of the Euphorbiaceae world checklist (Govaerts et al. 2000) in compiling the list presented here.

Systematics

1.Phyllanthus amentuligerMüll. Arg., Flora 48: 390. 1865. TYPE. FIJI ISLANDS:Vanua Levu: Mbua Bay, 1840, U.S. Expl. Exped. s.n. (holotype: probably G; Isotype: US!).

Diasperus amentuliger (Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 598. 1891. Glochidion amentuligerum Müll. Arg.) Croizat, Sargentia 1: 46. 1942.

Distribution. Endemic to Fiji and known from Vanua Levu and eastern Viti Levu at elevations of 100–400 m in dense or open forest, edges, and thickets (Smith 1981).

2.Phyllanthus amicorumG. Webster, Pacific Sci. 40: 100. 1986 [1988]. TYPE. TONGA: Eua, E. Soakai 341 (Holotype: K).

Distribution. Endemic to the island of Eua, Tonga (Webster 1986) where it is only known from forest margins and exposed rocks, 300m, on the Liku Plateau.

3.Phyllanthus anfractuosus(Gibbs) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112692-1

Basionym:Glochidion anfractuosum Gibbs, J. Linn. Soc. Bot. 39: 168. 1909. TYPE. FIJI ISLANDS: Viti Levu: Nandarivatu, Sep 1907, J. G. Gibbs 730 (holotype: BM).

Distribution. Endemic to Fiji and known only from Viti Levu and Ovalau at elevations from 100 to 1075 m, in dense or dry forest, thickets or ridge forest (Smith 1981).

4.Phyllanthus aoraiensisNadeaud, Énum. Pl. Tahiti 73. 1873. TYPE. SOCIETY ISLANDS: Tahiti: 1000 m, Nov 1857, J. Nadeaud 459 (holotype: P; isotypes: G [2], P[2]).

Distribution. Endemic to the Society Islands and known only from Tahiti at about 1000 m elevation on a ridge crest. Not collected since 1857 and presumed extinct (Florence 1997).

5.Phyllanthus atalotrichus (A.C. Sm.) W.L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112693-1

Basionym:Glochidion atalotrichum A.C. Sm., Contr. U.S. Natl. Herb. 37: 74. 1967. TYPE.FIJI ISLANDS: Viti Levu: Namosi, northern Korombasambasanga Range, drainage of Wainavindrau Creek, 450 to 600 m, 28 Sep 1953, A.C. Smith 8747 (holotype: US-02191397!; isotypes: GH, BISH!, L, NY!, S).

Distribution. Endemic to Fiji and known only from Namosi Prov., Viti Levu from 250–800 m in dense forest (Smith 1981).

6.Phyllanthus atrovirens(A.C. Sm.) W.L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112694-1

Basionym:Glochidion atrovirens A.C. Sm., Fl. Vitiensis Nova 2: 481,491. 1981. TYPE. FIJI ISLANDS: Viti Levu: Serua, hills between Waininggere and Waisese Creeks, between Ngaloa and Wainiyambia, 50 to 100 m, 10 Dec 1953, A.C. Smith 9550 (holotype: BISH-508142!; isotype: US!).

Distribution. Endemic to Fiji and know only from coastal and slightly inland areas of southern Viti Levu from 50–100 m in dry or dense forest (Smith 1981).

7.Phyllanthus bracteatus(Gillespie) W.L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112695-1

Basionym:Glochidion bracteatum Gillespie, Bernice P. Bishop Mus. Bull. 91: 15. 1932. TYPE. FIJI ISLANDS: Viti Levu: Rewa, SE slopes of Mt. Korombamba, 7 Aug 1927, J. W. Gillespie 2169 (holotype: BISH-508144!; isotypes: BISH [2]!).

Distribution. Endemic to Fiji from southern and eastern Vitu Levu, from 100 to 430 m in dense or open forest (Smith 1981).

8.Phyllanthus brothersonii(J. Florence) W.L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112696-1

Basionym:Glochidion brothersonii J. Florence, Fl. Polynésie Française 1: 68. 1997. TYPE.SOCIETY ISLANDS: Raiatea: Opoa, Mont Oropiro, épaulement nord. 200 m, 151°24’ W, 16°51’ S, 2 Jun 1990, J. Florence 10373 (holotype: P; isotypes: BISH!, CHR, DAV, K, L, P, PAP, PTBG!, US!).

Distribution. Endemic to the Society Islands and know only from Raiatea at apparently only low elevations from 200 to 250 m, collected in riparian forest with Hibiscus tiliaceus L. and in mesic ridge forest with Metrosideros collina (J. R. Forst. & G. Forst.) A. Gray and Commersonia bartramia (L.) Merr. (Florence 1997) The area has been converted to pine plantation (D. Hembry, pers. comm. 2011).

9.Phyllanthus brunnescens(A.C. Sm.) W.L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112697-1

Basionym:Glochidion brunnescens A.C. Sm., Fl. Vitiensis Nova 2: 482, 491. 1981. TYPE. FIJI ISLANDS: Viti Levu: Namosi, Mt., Nambui, third peak of Korombasambasanga Range, 12 Nov 1965, DA 14548 pro parte (coll. D. Koroiveibau & I. Qoro) (holotype: BISH-508147!; isotype: SUVA).

Distribution. Endemic to Fiji and know only from inland areas of Viti Levu and Vanua Levu, from 300 to 960 m in dense forest, ridge forest and scrubby forest (Smith 1981).

10.Phyllanthus calciphilus (Croizat) W.L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112698-1

Basionym:Glochidion calciphilum Croizat, Sargentia 1: 46. 1942. TYPE. FIJI ISLANDS: Fulaga: limestone cliff of lagoon, 0 to 80 m, 26 Feb 1934, A.C. Smith 1217 (holotype: GH; isotypes: BISH!, K, S, US!).

Distribution. Endemic to Fiji and know only from the two islands of southern Lau (Fulaga and Kabara) near sea level on limestone and lagoon cliffs (Smith 1981).

11.Phyllanthus christophersenii(Croizat) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112699-1

Basionym:Glochidion christophersenii Croizat, Occas. Pap. Bernice P. Bishop Mus. 17(16): 213. 1943. TYPE. SAMOAN ISLANDS: Savai`i: above Matavanu, 1300 m, 24 Jul 1931, E. Christophersen & E. P. Hume 2134 (holotype: A; isotype: BISH).

Distribution. Endemic to montane Savai`i and known from cloud forest at 1000–1550 m (Whistler 1978; Christophersen 1935).

12.Phyllanthus cleistanthoides(Fosberg) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112700-1

Basionym:Glochidion cleistanthoides Fosberg, Willdenowia 20: 263. 1991. TYPE. CAROLINE ISLANDS: Pohnpei: 1913, C. L. Ledermann 13599a (holotype: B-bc100249519!)

Distribution. Endemic to Pohnpei where it occurs from ca. 12 to 770 m elevation, most commonly in lowland wet forest and agroforest, but occasionally in summit cloud forest.

Note. This species is distinctive in having oblong-ovate to narrowly oblong-elliptic leaves and comparatively small flowers in umbel-like fascicles often borne on short, sometimes branched stalks or peduncles to 5 mm long, a densely puberulent pistil with a puberulent columnar style exserted for 1–1.5 mm beyond the calyx lobes, and relatively small, densely puberulent fruits. Pohnpei collections of this species have been identified as Glochidion ramiflorum or less commonly Glochidion puberulum.

13.Phyllanthus comitus(J. Florence) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112701-1

Basionym:Glochidion comitum J. Florence, Novon 7:29. 1997. TYPE. PITCAIRN ISLANDS: Pitcairn: Sommet Crete Sud-Est, 25°4’ S, 130°7’ W, 300 m, 19 Apr 1991, J. Florence 10740 (holotype: K; isotypes: BISH, BM, DAV, E, L, MO, P, PAP, TER, US!).

Distribution. Endemic to Pitcairn where it is known from fewer than 10 collections.

14.Phyllanthus concolor(Müll. Arg.) Müll. Arg., Flora 48: 374. 1865.

Basionym:Glochidion concolor Müll. Arg., Linnaea 32: 62. 1863. Phyllanthus concolor var. ellipticus (Müll. Arg.), Prodromus Systematis Naturalis Regni Vegetabilis 15(2): 290. 1866, nom. illeg. Diasperus concolor (Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 599. 1891. TYPE. FIJI ISLANDS:s.l., Nov 1855, W. H. Harvey s.n. (holotype: P; isotypes: BM, K).

Glochidion ramiflorum J.R. Forst. & G. Forst. var. lanceolatum Müll. Arg. Linnaea 32: 63. 1863. Phyllanthus ramifloris (J.R. Forst. & G. Forst.) Müll. Arg. var. lanceolatus (Müll. Arg.) Müll. Arg., Flora 48: 374. 1865, nom. illeg. TYPE. FIJI ISLANDS: Ovalau: Port Kinnaird, June 1860 and Somosomo, Taveuni, May 1860, B.C. Seemann 415 (lectotype: presumably G, designated by Smith (1981, p. 476); isolectotypes: BM, K).\\

Distribution. Widespread in Fiji, Tonga, and possibly Raratonga (Smith 1981), and also from Niue (Sykes 1970) from 0 to 1000 m elevation in dense or open forest, edges and forest-grassland transition, and on open slopes. These collections were previously referred to Glochidion ramiflorum, butwere considered by Smith (1981) to be a separate species. He considered Glochidion ramiflorum to be a species from New Guinea to New Hebrides.

15.Phyllanthus cordatus(Seem. ex Müll. Arg.) Müll. Arg., Flora 48: 376. 1865.

Basionym:Glochidion cordatum Seem. ex Müll. Arg., Linnaea 32: 64. 1863. Diasperus cordatus (Seem. ex Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 599. 1891. TYPE. FIJI ISLANDS: Viti Levu: July 1860 and Ovalau: Port Kinnaird, June 1860 [the K isotype bears 2 field labels], B.C. Seemann 416 (holotype: presumably G; isotypes: BM, K).

Distribution. Endemic to Fiji and know from Viti Levu, Ovalau, and Vanua Levu at 50–900 m, in dry forest, ridge forest, and in pastures and along roadsides (Smith 1981).

16.Phyllanthus cuspidatusMüll. Arg., Flora 48: 377. 1865.

Basionym:Glochidion cuspidatum (Müll. Arg.) Pax, Bot. Jahrb. Syst. 25: 645. 1898. Diasperus cuspidatus (Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 599. 1891. TYPE. SAMOAN ISLANDS:s.l., U.S. Expl. Exped. s.n. (holotype: G-DC).

Distribution. Endemic to the Samoan Islands (Savai`i, `Upolu), and Tutuila at up to 400–450 m in ridge forest (Whistler 1980).

17.Phyllanthus distichusHook. & Arn., Bot. Beechey Voy.: 95. 1832.

Basionym:Diasperus distichus (Hook. & Arn.) Kuntze, Revis. Gen. Pl. 2: 599. 1891. TYPE. HAWAIIAN ISLANDS:O`ahu: 1827–1827, G.T. Lay & A. Collie s.n. (probable holotype: K; probable isotype: E).

Distribution.Phyllanthus distichus is endemic to the Hawaiian islands where it is occasional to locally common in mesic forest, often on steep slopes or ridge tops, sometimes in dry shrubland at 60–950 m on the islands of Kaua`i, O`ahu, Moloka`i, Lana`i, West Maui, and rare on East Maui. A number of additional names have been applied to populations of Phyllanthus distichus, but they were placed into synonymy by Wagner et al. (1990, 1999) and are not repeated here.

18.Phyllanthus emarginatus(J. W. Moore) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112702-1

Basionym:Glochidion emarginatum J. W. Moore, Bernice P. Bishop Mus. Bull. 102: 30. 1933. TYPE.SOCIETY ISLANDS: Raiatea: Mount Temehani, 470 m, 1 Jan 1927, J.W. Moore 476A (holotype: BISH-508150!; isotype: P [2]).

Glochidion raiateense J. W. Moore, Bernice P. Bishop Mus. Bull. 102: 30. 1933. TYPE:SOCIETY ISLANDS: Raiatea: Mount Temehani, 470 m, 1 Jan 1927, J.W. Moore 476B (Holotype: BISH-508230!).

Distribution. Endemic to the Society Islands and known only from Raiatea at 580–750 m where it occurs in marshy shrubland with Metrosideros collina and species of Cyperaceae. Also collected at ca. 930 m in mesic ridge shrubland with Ilex and Weinmannia (Florence 1997).

19.Phyllanthus euryoides(A. C. Sm.) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112703-1

Basionym:Glochidion euryoides A.C. Sm., J. Arnold Arbor. 33: 373. 1952. TYPE. FIJI ISLANDS: Viti Levu: Mba, upper slopes of Mt. Koromba (Pickering Peak), 3 Jun 1947, A. C. Smith 4659 (holotype: A; isotypes: BISH!, K, S, US!).

Distribution. Endemic to Fiji and known only from the type collection.

20.Phyllanthus florenceiW. L. Wagner & Lorence, nom. nov.

urn:lsid:ipni.org:names:77112704-1

Replaced name:Glochidion societatis J. Florence, Fl. Polynésie Française 1: 90. 1997. TYPE. SOCIETY ISLANDS: Tahaa: Patio, Mt. Purauti, crête sud-est, 151°30’ W, 16°37’ S, 225 m, 18 Jun 1990, J. Florence 10,627(holotype: P; isotypes: BISH!, CHR, DAV, K, L, NY!, P [2], PAP, PTBG!, US!).

Distribution. Known from the Society Islands (Huahine, Mauapiti, Raiatea, and Tahaa) and Austral Islands (Rimatara) at 0–225 m elevation where it is locally common to abundant in coastal vegetation with Scaevola and Euphorbia on coral sand, in lowland mesic forest with Neolauclea and Hibiscus tiliaceus, in mesic ridge forest with Metrosideros collina and Dicranopteris linearis (Burm. f.) Underw., and in secondary forest (Florence 1997).

21.Phyllanthus gillespiei(Croizat) W.L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112705-1

Basionym:Glochidion gillespiei Croizat, Sargentia 1: 46. 1942. TYPE. FIJI ISLANDS:Viti Levu: Namosi, near summit of Mt. Naitarandamu, 28 Sep 1927, J.W. Gillespie 3161 (holotype: GH; isotypes: BISH).

Distribution. Endemic to Fiji and known only from mountainous areas of Viti Levu from 750–1155 m in dense or ridge forest (Smith 1981).

22. Phyllanthus grantii(J. Florence) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112706-1

Basionym:Glochidion grantii J. Florence, Bull. Mus. Natl. Hist. Nat., B, Adansonia 18: 250. 1996. TYPE. SOCIETY ISLANDS: Tahaa: Tapuamu, crête entre les Monts Tete et Ohiri. 530 m, 151°31’ W, 16°37’ S, 5 Nov 1992, J. Florence & R. Tahuaitu 11816 (holotype: P; isotype: BISH!, DAV, K, L, PAP, US!).

Distribution. Endemic to the Society Islands (Raiatea and Tahaa) where it occurs at 435–730 m elevation in marshy shrubland with Metrosideros and Cyperaceae and in wet summit forest with Metrosideros and Macaranga (Florence 1997).

23.Phyllanthus grayanusMüll. Arg., Flora 48(24): 380. 1863.

Diasperus grayanus (Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 599 .1891. Glochidion grayanum (Müll. Arg.) J. Florence, Bull. Mus. Natl. Hist., B, Adansonia 18: 250. 1996. TYPE. SOCIETY ISLANDS: Tahiti:s. l., Sep 1839 or Jan1841, U. S. Expl. Exped. s.n. (Holotype: G-DC).

Distribution. Endemic to the Society Islands and known only from Tahiti at 60–1040 m. It occurs in lowland riparian forest with Neonauclea and Hibiscus tiliaceus and extends up to mesic ridge and summit forest dominated by Metrosideros collina (Florence 1997).

24.Phyllanthus heterodoxusMüll. Arg., in DC., Prodr. 15(2):321. 1866.

Diasperus heterodoxus (Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 599. 1891. Glochidion heterodoxum (Müll. Arg.) Pax & K.Hoffm. in H.G.A. Engler, Nat. Pflanzenfam. ed. 2, 19c: 58. 1931. TYPE. FIJI ISLANDS:s. l.,1840, U.S. Expl. Exped. s.n. (Holotype: Probably G).

Distribution. Endemic to Fiji and known only from Vanua Levu and Lau Group, Fulaga from 0 to 870 m (Smith 1981).

25.Phyllanthus hivaoaense(J. Florence) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112707-1

Basionym:Glochidion hivaoaense J. Florence, Fl. Polynésie Française 1: 74. 1997. TYPE. MARQUESAS ISLANDS: Hiva Oa: Atuona, piste de Hanamenu, NW du Mt. Temetiu, 139°5’ W, 9°48’ S, 1100 m, 30 Jul 1988, J. Florence & S. Perlman 9673 (holotype: P; isotypes: BISH!, DAV, K, L, P, PAP, PTBG!, US!). Fig. 1.

Figure 1.

Phyllanthus hivaoaense (J. Florence) W. L. Wagner & Lorence. A branch B abaxial leaf surface C leaf axil showing stipules D female flowers E female flower close-up F–G calyx lobes H longitudinal view of female flower I capsule. A–I...

Distribution. Endemic to the Marquesas Islands of Hiva Oa and Tahuata, from about 700 to 1200 m, collected in wet shrubland and forest with Freycinetia, Weinmannia, and the tree ferns Alsophila and Sphaeropteris (Florence 1997).

26.Phyllanthus hosokawae(Fosberg) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112708-1

Basionym:Glochidion hosokawae Fosberg, Willdenowia 20: 261. 1991. TYPE: CAROLINE ISLANDS: Pohnpei: Awak, 25.8.1980, F.R. Fosberg 60467 (holotype: US [not located]: isotypes: BISH, L).

Distribution. To date know only from the type collection from Pohnpei; the holotype could not be located at US.

Note. This entity should be carefully evaluated in the context of an overall review of Phyllanthus in Micronesia. It may be conspecific with Glochidion cleistanthoides, in which case we would adopt the latter name because the description applies most closely to this species and the holotype is available at B.

27.Phyllanthus huahineense(J. Florence) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112709-1

Basionym:Glochidion huahineense J. Florence, Fl. Polynésie Française 1: 75. 1997. TYPE. SOCIETY ISLANDS: Huahine: Maeva, motu Oavarei, secteur Haaparu, 151°W, 16°41’ S, 2 m, 1 Nov 1992, J. Florence & R. Tahuaitu 11745 (holotype: P ; isotypes: BISH, DAV, K, L, NY, P, PAP, PTBG !, US !).

Distribution. Endemic to the Society Islands and known only from Huahine on the islet Motu de Maeva at 1–4 m elevation, restricted to coral sand substrate in coastal vegetation with Casuarina, Guettarda, and Tournefortia (Florence 1997).

28.Phyllanthus inusitatus(A.C. Sm.) W.L. Wagner & Lorence, comb. nov.

Basionym:Glochidion inusitatum A.C. Sm., Fl. Vitiensis Nova 2: 486, 493. 1981. TYPE. FIJI ISLANDS: Vanua Levu: divide between Wainunu and Ndreketi Rivers, between Nanduna (old village site) and Mt. Ndelanathau, 17 May 1934, A.C. Smith 1851 (holotype: BISH-142932; isotypes: many indicated, but not specific as to where deposited).

Distribution. Endemic to Fiji and known only from the type.

29.Phyllanthus jardinii Müll. Arg., Linnaea 32: 21. 1863. TYPE. s. l., D.E.S.A. Jardin s.n. (holotype: G-DC-FP3347; isotypes: P [2]). Type presumed to be from Nuku Hiva in the Marquesas Islands.

urn:lsid:ipni.org:names:77112710-1

Note. Precise locality uncertain, known only from a single collection possibly made in the Marquesas Islands (Nuku Hiva) by Jardin in the 19th century (Florence 1997).

30.Phyllanthus kanehirae(Hosokawa) W. L. Wagner &Lorence, comb. nov.

urn:lsid:ipni.org:names:77112711-1

Basionym:Glochidion kanehirae Hosokawa, Trans. Nat. Hist. Soc. Taiwan 25: 22. 1935. TYPE. CAROLINE ISLANDS: Palau: Jul–Aug 1929, R. Kanehira 241 (holotype: TAI).

Distribution. Caroline Islands: Palau (Peleliu, Koror including the Rock Islands of Mecherchar, Ngeruktabel, Ulebsechel and Urukthapel, and also known from Babeldaob [Fosberg et al. 1979; Wagner et al. unpubl.]), Yap (Yap Island), and Chuuk (Moen Island, also Tol, Udot, Uman, Dublon, Fano, Fanurmot [Fosberg et al. 1979; Wagner et al. unpubl.]). On Palau it is restricted to limestone substrate from near sea level to about 200 m elevation in coastal forest with speices of Bruguiera, Heretiera, Semecarpus, Osmoxylon, and Phyllanthus, and in lowland evergreen forest with Horsfieldia and Phyllanthus and agroforest. On Chuuk it occurs near sea level (3 m) in coastal forest interspersed with mangroves and on slopes of unknown elevation in agroforest and secondary vegetation. On Yap it occurs from near sea level at the edge of mangrove vegetation and on slopes up to 40 m with secondary vegetation.

Note. This glabrous species is characterized by female flowers with a small, depressed-globose pistil 1–1.5 mm long with a very short 10–11-fid stylar column 0.5 mm long. Certain collections from Yap have been identified as Glochidion cf. ramiflorum, and certain collections from Palau have been identified as Glochidion palauense.

31.Phyllanthus longfieldiaeL. Riley, Bull. Misc. Inform. Kew [2]:55. 1926.

Glochidion longfieldiae (L. Riley) F. Br., Bernice P. Bishop Mus. Bull. 130: 141. 1935. TYPE. AUSTRAL ISLANDS: Rapa: L.A.M. Riley 776 (holotype: K; isotype: BM).

Distribution. Endemic to the Austral Islands (Rapa) where it occurs from 140 to 575 m, in mesic Metrosideros forest on slopes and in valleys and ravines with species of Corokia and Fitchia (Florence 1997).

32.Phyllanthus macrosepalus(Hosokawa) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112712-1

Basionym:Glochidion macrosepalum Hosokawa, Trans. Nat. Hist. Soc. Taiwan 25: 21. 1935. TYPE. CAROLINE ISLANDS: Palau: Babeldaob, valley near Mt. Agade, “Arukoron-sogan”, 20 Sep 1933, T. Hosokawa 7053 (holotype: TAI-118940; isotypes: BISH, GH, TAI). Holotype presumed to be the TAI sheet with the typewritten label as is typical of Hosokawa collections.

Distribution. Endemic to Palau (Babeldaob, Anguar, Peliliu, and Malakal Islands), where it occurs at 3–150 m, in lowland evergreen moist and wet forest on limestone and limestone-derived soils.

Note. This glabrous species is characterized by female flowers with relatively large calyx lobes enclosing the short, depressed-ovoid pistil 1.5 mm long with a very short style column and flat 5-6-sulcate or -lobed stigmatic region. In male flowers the staminal column is composed of 5 connate stamens (versus 3-4 in other Micronesian species).

33.Phyllanthus manono(Baill. ex Müll. Arg.) Müll. Arg., Flora 48: 377. 1865.

Basionym:Glochidion manono Baill. ex Müll. Arg.,Linnaea 32: 65. 1863. Diasperus manono (Baill. ex Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 600. 1891. TYPE. SOCIETY ISLANDS: Tahiti:s. l.,J. Lepine 210 (lectotype: G-DC, designated by Florence, Fl. Polynésie Française 1: 79. 1997; isolectotypes: P [2]).

Distribution. Endemic to the Society Islands of Moorea and Tahiti where it occurs at 30–1000 m, in valleys in wet forest with Neonauclea, Hibiscus, and Inocarpus and also on slopes and summits, sometimes in disturbed or secondary forest with Metrosideros, Dicranopteris, and Psidium (Florence 1997).

34.Phyllanthus marchionicus(F. Br.) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112713-1

Basionym:Glochidion marchionicum F. Br., Bernice P. Bishop Mus. 130: 142. 1935. TYPE. MARQUESAS ISLANDS: Ua Huka: 9 Nov 1922, E. H. Quayle 1689 (Lectotype: BISH-508228! & 508229!, designated by St. John, Phytologia 33: 420. 1976). Fig. 2.

Figure 2.

Phyllanthus marchionicus (F. Br.) W. L. Wagner & Lorence. Branch with dehisced fruit showing red arillate seeds, with trunk in background. Field image by K. R. Wood: Marquesas Islands. Ua Huka, Haahue, northwestern coastal valley,Wood 10765 (PTBG,...

Glochidion tooviianum J. Florence, Bull. Mus. Natl. Hist. Nat., B, Adansonia 18: 260. 1996, syn. nov. TYPE. MARQUESAS ISLANDS: Nuku Hiva: Toovii, 800 m, 140°9’ W, 8°51’ S, 3 Mar 1986, J. Florence 7445 (holotype: P; isotypes: BISH!, P, PAP, US).

Distribution. Endemic to the Marquesas Islands (Nuku Hiva, Ua Huka, Ua Pou, Hiva Oa, Tahuata, and Fatu Hiva) from 50 to 1130 m, where it is widespread in moist valleys with Hibiscus tiliaceus, disturbed mesic ridge forest with Casuarina, Sapinus,and Xylosma, and secondary vegetation with Dicranopteris, Leucaena, Miscanthus,and Psidium or in primary wet forest with Cheirodendron, Crossostylis, Ilex, and tree ferns (Florence 1997).

35.Phyllanthus mariannensisW.L. Wagner & Lorence, nom. nov.

urn:lsid:ipni.org:names:77112714-1

Replaced name:Glochidion marianum Müll. Arg., Linnaea 32: 65. 1863. Phyllanthus gaudichaudii Muell.-Arg. var marianus (Müll. Arg.) Müll. Arg., in DC. Prodr. 15(2): 300. 1866. TYPE. MARIANA ISLANDS: Guam: C. Gaudichaud-Beaupré 139(holotype: G-DC).

Distribution. Endemic to the Mariana Islands (Guam) where it occurs on limestone and basaltic soils in old fields and grasslands and disturbed native and secondary vegetation at ca. 60–150 m elevation.

Note. The ovate leaves with short acuminate to rounded apices and glabrous columnar style 1.5–2 mm long exserted beyond the calyx lobes are characteristic of this species. Collections from Caroline Islands (Pohnpei) previously identified as this species are here considered to represent Phyllanthus senyavinianus.

36.Phyllanthus marianusMüll. Arg., Linnaea 32:17. 1863.

Diasperus marianus (Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 600. 1891. TYPE. MARIANA ISLANDS: Guam:s.l., 17 March to 3 April 1819, C. Gaudichaud-Beaupré s. n. (holotype: G-DC).

Distribution. Endemic to the Mariana Islands (Guam, Aguijan, Agrihan, Alamagan, Anathan, Asuncion, Guguan, Maug, Pagan, Rota, Saipan, Sarigan, Tinian) and the Caroline Islands (only on Ulithi Atoll). On Guam it is common on limestone cliffs and terraces.

37.Phyllanthus melvilliorum(Airy Shaw) W.L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112715-1

Basionym:Glochidion melvilliorum Airy Shaw., Kew Bull. 25: 487. 1971. TYPE. FUJI ISLANDS: Viti Levu: Nausori Highlands, Nandronga and Navosa Province, 2 May 1962, R. & E.F. Melville & J.W. Parham 7048 (holotype: K; isotypes: BISH, SUVA).

Distribution. Endemic to Fiji and known only from the vicinity of the type locality at 600–670 m in dense or mixed forest (Smith 1981).

38.Phyllanthus multilobus(A.C. Sm.) W.L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112716-1

Basionym:Glochidion multilobum A.C. Sm., Fl. Vitiensis Nova 2: 484, 493. 1981. TYPE. FIJI ISLANDS: Vanua Levu: Thakaundrove Prov., SW slope of Mt. Batini, 28 Nov 1933, A.C. Smith 606 (holotype: BISH-142931; isotypes: many indicated, but not specific as to where deposited).

Distribution. Endemic to Fiji and know only from Mt. Batini and Mt. Seatura on Vanua Levu from 300 to 800 m in dense or crest forest (Smith 1981).

39.Phyllanthus nadeaudii(J. Florence) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112717-1

Basionym:Glochidion nadeaudii J. Florence, Bull. Mus. Natl. Hist. Nat., B, Adansonia 18: 253. 1996. TYPE. SOCIETY ISLANDS: Moorea: Pao Pao, crête N du Mt. Fairurani. 670 m, 149°47’ W, 17°30’ S, 14 May 1987, J. Florence 8287 (holotype: P; isotypes: BISH!, PAP, US!).

Distribution. Endemic to the Society Island of Moorea at 420–800 m, where it typically occurs on high ridge slopes and crests with Weinmannia, Metrosideros, Dicranopteris, and Nephrolepis (Florence 1997).

40.Phyllanthus orohenense(J. W. Moore) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112718-1

Basionym:Glochidion orohenense J. W. Moore, Bernice P. Bishop Mus. 16: 6. 1940. TYPE: SOCIETY ISLANDS: Tahiti: south side of Mt. Orohena, 1300 m, 14 May 1927, L. H. McDaniels 1312(holotype: BH; isotype: BISH!).

Distribution. Endemic to the Society Island of Tahiti where it is rare and localized at 900–1750 m, in cloud forest with Metrosideros, Weinmannia, and Alsophila (Florence 1997).

41.Phyllanthus otobediiW. L. Wagner & Lorence, nom. nov.

urn:lsid:ipni.org:names:77112719-1

Replaced name:Glochidion palauense Hosokawa, Trans. Nat. Hist. Soc. Taiwan 25: 22. 1935. TYPE. CAROLINE ISLANDS: Palau: Aurapushekaru I. (Oropusyakaru-to), S. of Koror I., 8 Oct 1933, T. Hosokawa 7453 (holotype: TAI; isotype: BISH!, MICH, US!, Z).

Distribution. Endemic to Palau on Babeldaob, Anguar, Ngerechong, Ulebsechel, Ulong, and the Rock Islands (Koror, Ngerukeuid, Ngeruktabel). On Babeldaob this species occurs on basaltic soils in evergreen wet forest and savannas at about 100 m elevation. On the Rock Islands it occurs on limestone substrate near sea level (2–5 m elevation) in evergreen coastal forest and cliff vegetation. This glabrous species is distinguished by its female flowers with a conical-columnar pistil 3 mm long with a cylindrical style and shortly 6–7-lobed stigma.

Etymology. We are pleased to name this species for Mr. Demei O. Otobed, president of the board of directors of the Belau National Museum, who has done so much to advance the study and conservation of Palau’s biodiversity.

Note. Some Palau collections of this species were identified as Glochidion macrosepalum.

42.Phyllanthus pacificusMuell. Arg, Linnaea 32 : 31. 1863.

Diasperus pacificus (Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 600. 1891. TYPE. MARQUESAS ISLANDS: Nuku Hiva: D.E.S.A. Jardin 122 (lectotype: P, designated by Florence, Fl. Polynésie Française 1: 123. 1997; isolectotype: G-DC [2], P).

Phyllanthus pacificus Müll. Arg. var. uapensis F. Br., Bernice P. Bishop Mus. Bull. 130 : 138. 1935, syn. nov. TYPE. MARQUESAS ISLANDS: Ua Pou: 9 Sep 1922, E. H. Quayle s.n. (holotype: BISH-143921! & 508277!; isotype: BISH).

Phyllanthus pacificus Müll. Arg. var. quaylei F. Br., Bernice P. Bishop Mus. Bull. 130: 139. 1935, syn. nov. TYPE. MARQUESAS ISLANDS: Nuku Hiva: s.l., 15? Oct 1922, E. H. Quayle 1341 (holotype: BISH-508703!).

Phyllanthus pacificus Müll. Arg. var. uahukensis F. Br., Bernice P. Bishop Mus. Bull. 130: 139. 1935, syn. nov. TYPE: MARQUESAS ISLANDS: Hiva Oa: Kopaafaa, 2770 ft, 2 Aug 1929, E.P. Mumford & A.M. Adamson 488 (lectotype: BISH-508705!; designated by St. John, Phytologia 33: 420. 1976). Additional syntypes: MARQUESAS ISLANDS: Ua Huka:s.l., 9 Nov 1922, E. H. Quayle 1781 (BISH-508704! & -143920!).

Distribution. Endemic to the Marquesas Islands (Nuku Hiva, Hiva Oa, Tahuata, Ua Pou, Ua Huka, and Fatu Hiva) where it is widespread and occurs from about 25 to 1085 m, growing in open areas or on rocky slopes and cliffs in secondary vegetation with Dicranopteris and grasses. At higher elevations it grows among fern cover in Metrosideros-Weinmannia wet shrubland and forest (Florence 1997).

43.Phyllanthus palauensisHosokawa, Trans. Nat. Hist. Soc. Taiwan 25: 19. 1935. TYPE. CAROLINE ISLANDS: Palau: Babeldaob Island, near Almatin at low altitudes, 17 Sep 1933, T. Hosokawa 6921 (holotype: TAI; isotype: GH).

Distribution. Endemic to Palau (Babeldaob, Ngemelachel, Ulong) and Rock Islands (Koror, Mecherchar, Ngerukeuid, Ngerekebesang, Ngeruktabel) where it occurs to at least 30 m elevation on volcanic soils in wet forests along streams and in savannah vegetation.

44.Phyllanthus papenooense(J. Florence) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112720-1

Basionym.Glochidion papenooense J. Florence, Bull. Mus. Natl. Hist. Nat., B, Adansonia 18: 254. 1996. TYPE: SOCIETY ISLANDS: Tahiti: Papenoo, Ofetanu, 160 m, 149°26’ W, 17°38’ S, 9 Sep 1989, J. Florence 9901 (holotype: P; isotypes: BISH!, PAP).

Distribution. Endemic to the Society Island of Tahiti where it is apparently rare and known only from Papenoo Valley up to about 650 m in riparian forest with Hibiscus and Neonauclea invaded by Miconia calvescens DC. (Florence 1997).

45.Phyllanthus pergracilisGillespie, Bernice P. Bishop Mus. Bull. 91: 18. 1932. TYPE. FIJI ISLANDS: Viti Levu: Naitasiri Province, Tamavua woods, 11 km from Suva, 150 m, 9 Aug 1927, J.W. Gillespie 2122 (holotype: BISH-508710!; isotypes: GH, UC).

Distribution. Endemic to Fiji and known only from Viti Levu at 30–1200 m (Smith 1981; Webster 1986).

46.Phyllanthus pinaiensisS.L.Welsh, Flora Societensis 112. 1998.

Basionym:Phyllanthus urceolatus Baill., Adansonia 2: 239. 1862. Diasperus urceolatus (Baill.) Kuntze, Revis. Gen. Pl. 2: 601. 1891, non Phyllanthus urceolatus Noronha (1790). TYPE. SOC IETY ISLANDS: Tahiti: “Nouvelle-Caledonie, Port de France”, E. Vieillard 336 (holotype: P; isotype: P).

Distribution. Endemic to the Society Islands of Moorea, Raiatea, and Tahiti from 150–830 m, usually in understory of valley forest with Hernandia, Hibiscus, and Neonauclea or sometimes on ridge crests. Not collected on Tahiti since the end of the 19th century but apparently still frequent on Moorea (Florence 1997).

Note. According to Florence (1997) the type of Phyllanthus urceolatus Baill. is from Tahiti, not New Caledonia. This is likely due to a labeling error.

47.Phyllanthus pitcairnense(H. St. John) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112721-1

Basionym.Glochidion pitcairnense (F.Br.) H.St.John, Trans. Roy. Soc. New Zealand, Bot. 1: 187. 1962. Glochidion tahitense var. pitcairnense F. Br., Bernice P. Bishop Mus. Bull. 130: 142. 1935. TYPE: PITCAIRN ISLANDS:s.l., 1922, E. H. Quayle s.n. (holotype: BISH-508246! & -508247!).

Distribution. Endemic to the Pitcairn Islands (Henderson and Pitcairn), from about 30 to 270 m elevation. On Henderson it occurs at 30 m on eroded calcarenite in beach forest and scrub with Nesoluma, Pisonia, and Xylosma. On Pitcairn it has been collected at 270 m in secondary upland vegetation with grasses and ferns.

48.Phyllanthus podocarpusMüll. Arg., Flora 48: 388. 1865.Diasperus podocarpus (Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 600. 1891. Glochidion podocarpum (Müll. Arg.) C.B. Robinson, Philippine J. Sci. Bot. 6: 300. 1911. TYPE. FIJI ISLANDS:s. l., 1840, U.S. Expl. Exped. s.n. (holotype: probably G; isotype: US!).

Distribution. Endemic to Fiji and know only from the type collection.

49.Phyllanthus ponapense(Hosokawa) W. L. Wagner &Lorence, comb. nov.

urn:lsid:ipni.org:names:77112722-1

Basionym.Glochidion ponapense Hosokawa, Trans. Nat. Hist. Soc. Taiwan 25: 24. 1935. TYPE: CAROLINE ISLANDS: Pohnpei: summit of Mt. Troton, 13 Aug 1933, T. Hosokawa 5770 (holotype: TAI).

Glochidion excorticans Fosberg var. calvum Fosberg, Willdenowia 20: 261. 1991, syn. nov. TYPE: CAROLINE ISLANDS: Pohnpei: 1913-1914, C. L. Ledermann 13333 (holotype: B-bc100249513!).

Distribution. Endemic to Pohnpei, this species occurs from lowland wet forest up into montane cloud forest on summits from 20 to 732 m elevation.

Note. This species is characterized by its female flowers with a short, glabrous, depressed-globose ovary 0.5 mm long and glabrous columnar style 1-1.5 mm long with a 6-7-dentate stigma. Some collections of Phyllanthus ponapense were previously identified as Glochidion marianum or Glochidion ramiflorum.

50.Phyllanthus raiateaensisW. L. Wagner & Lorence, nom. nov.

urn:lsid:ipni.org:names:77112723-1

Replaced name:Glochidion moorei P. T. Li, Acta Phytotax. Sin. 20: 117. 1982, non Phyllanthus moorei M. Schmid (1991). Glochidion salicifolium J. W. Moore, Bernice P. Bishop Mus. Bull. 226: 13. 1963, non Glochidion salicifolium (Baill.) Müll. Arg. (1863) nec Phyllanthus salicifolius Baill (1862). TYPE. SOCIETY ISLANDS:Raiatea: Temihani Plateau, 5 Oct 1934, H. St. John 17250 (holotype: BISH-508232!; isotype: BISH).

Note. The sheet (BISH-142814) was considered by Florence (1997) to not be part of the type because it was pubescent vs. glabrous as in the holotype. The full variation of Phyllanthus raiateaensis is not well understood, but Florence accepted both specimens as this species. Without further supporting information that this really represents a mixed collection it seems best to accept the second sheet as an isotype. However, even if the second sheet is not accepted as an isotype, the first sheet is clearly the holotype as it was explicitly stated as such and shown in a figure in the original publication.

Distribution. Endemic to the Society Island of Raiatea where it is known from the Temehani plateau region at 435–750 m, occurring in wet forest with Metrosideros, Weinmannia, and Myrsine and in open marshland with Metrosideros and species of Cyperaceae (Florence 1997).

51.Phyllantus raivavense(F. Br.) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112724-1

Basionym.Glochidion raivavense F. Br., Bernice P. Bishop Mus. Bull. 130: 142. 1935. TYPE. AUSTRAL ISLANDS: Raivavae: 23 Mar 1922, A. M. Stokes 43 (holotype: BISH-508248!).

Distribution. Endemic to the Austral Islands (Raivavae, Rurutu, and Tubuai). It occurs at 10–340 m in primary and secondary vegetation including riparian forest with Aleurites, Hernandia, Hibiscus tiliaceus, and Metrosideros, and on dry slopes or crests with Celtis, Dicranopteris, Xylosma, and grasses (Florence 1997).

52.Phyllanthus rapaense(J. Florence) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112725-1

Basionym.Glochidion rapaense J. Florence, Bull. Mus. Natl. Hist. Nat., B, Adansonia 18: 258. 1996. TYPE. AUSTRAL ISLANDS: Rapa: flanc SE du Mt. Pukumia, 150 m, 144°19’ W, 27°36’ S, 5 Feb 1984, J. Florence 6465 (holotype: P; isotypes: BISH!, K, PAP, US!).

Distribution. Endemic to the Austral Island of Rapa, at 50–330 m in mesic forest with Metrosideros, Meryta, and Freycinetia, and sometimes on wet rocks or cliffs (Florence 1997).

53.Phyllanthus rupiinsularisHosokawa, Trans. Nat. Hist. Soc. Taiwan 25: 19. 1935. TYPE. CAROLINEISLANDS: Palau: upon a coral islet near the island of Urktable [Ngeruktabel], 15 Oct 1933, T. Hosokawa 7534 (holotype: TAI; isotype: GH, US [2]!).

Distribution. Endemic to Palau where it occurs on the Rock Islands (Ngerukeuid, Ngeruktabel, Ulong), on limestone substrate in coastal cliff vegetation near sea level to about 5 m elevation.

54.Phyllanthus saffordiiMerr., Philipp. J. Sci., C 9: 104. 1914. TYPE. MARIANA ISLANDS: Guam: hills back of Piti, 100 m, Oct 1911, R. C. McGregor 476 (lectotype: US- 01860446!, here designated).The type in PNH was destroyed during World War II.

Distribution. Endemic to the Maraiana Islands (Guam, Alamagan, Anatahan, Pagan, Saipan, and Tinian). This species occurs in savannah vegetation.

55.Phyllanthus st-johniiW. L. Wagner & Lorence, nom. nov.

urn:lsid:ipni.org:names:77112726-1

Replaced name:Glochidion myrtifolium J. W. Moore, Bernice P. Bishop Mus. Bull. 226: 10. 1963. TYPE. SOCIETY ISLANDS: Raiatea: S ridge of Ereeo Valley, 9 Oct 1934, H. St. John 17328 (holotype: BISH-508233!; isotype: BISH).

Glochidion longipedicellatum J.W.Moore, Bernice P. Bishop Mus. Bull. 226: 9. 1963, nom. illeg., non Yamamoto (1933). Glochidion longipes P.T.Li, Acta Phytotax. Sin. 20:117. 1982. TYPE. SOCIETY ISLANDS: Raiatea: south side of Toahiva Valley, 200 m, 7 Oct 1934, H. St. John 17305 (holotype: BISH-508155!; isotype: BISH).

Distribution. Endemic to the Society Islands (Bora Bora, Moorea, Raiatea, and Tahaa) where it occurs at 30–680 m in primary or secondary mesic or wet forest with Hibiscus tiliaceus and Nauclea in valleys, or with Dicranopteris, Metrosideros, and Psidium on slopes and ridges (Florence 1997).

56.Phyllanthus samoanus(Müll. Arg.) W. L. Wagner & Lorence, comb. et stat. nov.

urn:lsid:ipni.org:names:77112727-1

Basionym:Phyllanthus ramiflorus (J.R. Forst. & G. Forst.) Müll. Arg. var. samoanus Müll. Arg. in A.P.de Candolle, Prodr. 15(2): 289. 1866. Glochidion ramiflorum J.R. Forst. & G. Forst. var. samoanum (Müll. Arg.) Pax, Bot. Jahrb. Syst. 25: 645. 1898. TYPE. SAMOAN ISLANDS:s.l., U.S. Expl. Exped. s.n. (holotype: probably G-DC).

Phyllanthus gaudichaudii var. samoanus Müll. Arg. in A.P.de Candolle, Prodr. 15(2):300. 1866. Glochidion cuspidatum var. samoanum .(Müll. Arg.) Pax, Bot. Jahrb. Syst. 25:645. 1898. TYPE. SAMOAN ISLANDS:s.l., U.S. Expl. Exped. s.n. (holotype: probably G-DC).

Distribution. Endemic to the Samoan Islands (Savai`i, Upolu, Tutuila, Aunu`u, Ofu, Olosega, and Ta`u) at 60–1000 m disturbed forest, secondary forest, and pastures (Whistler 1980).

Note. Even though the varietal name was published under an illegitimate species it is legitimate under Art. 55.2 of the ICBN and is available for use at the specific level. These Samoan collections were previously referred to Glochidion ramiflorum, butwere considered by Smith (1981) to be a separate species. He considered Glochidion ramiflorum to be a species from New Guinea to New Hebrides [see Excluded Names].

57.Phyllanthus seemannii(Müll. Arg.) Müll. Arg., Flora 48: 374. 1865 (as seammanianus).

Basionym.Glochidion seemannii Müll. Arg., Linnaea 32: 63. 1863 (as seemanni). Diasperus seemannii (Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 600. 1891. TYPE. FIJI ISLANDS: Kadavu: s.l., 1860, B.C. Seemann 413 (holotype: probably G; isotypes: BM, K).

Phyllanthus venulosus Müll. Arg., J., Flora 48:374. 1865. Diasperus venulosus (Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 601. 1891. Glochidion venulosum (Müll. Arg.) P.T.Li, Guihaia 14: 131. 1994. TYPE. FIJI ISLANDS:s.l., 1840, U.S. Expl. Exped. s.n. (Holotype: probably G; Isotypes: GH, US).

Distribution. Endemic to Fiji where it is known from Viti Levu, Ovalau, Vanua Levu, Taveuni, and Moala, but Smith (1981) thought it is most likely more widespread, at 0–1150 m, in dense dry or secondary forest or on more open hillsides.

58.Phyllanthus senyavinianus(Glassman)W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112728-1

Basionym.Glochidion senyavinianum Glassman, Bernice P. Bishop Mus. Bull. 209: 71. 1952. TYPE. CAROLINE ISLANDS: Pohnpei: Mt. Ninani, 731 m, 17 Aug 1949, S. Glassman 2884 (holotype: US-02158415!; isotypes: BISH, OKL).

Glochidion puberulum Hosokawa, Trans. Nat. Hist. Soc. Taiwan 25:23. 1935, non Phyllanthus puberulus Miq. ex Baill., syn. nov. (1858). TYPE. CAROLINE ISLANDS: Pohnpei: 8 Aug 1933, T. Hosokawa 5523 (holotype: TAI; isotype: US!).

Glochidion excorticans Fosberg, Willdenowia 20: 260. 1991, syn. nov. TYPE. CAROLINE ISLANDS: Pohnpei: 1913-1914, C. L. Ledermann 13643x (holotype: B!).

Distribution. Caroline Islands, known from Pohnpei and Chuuk (Fano, Dublon, Nomwin, Melot Moen, Romonum, Tol, Udot, and Uman). On Pohnpei it occurs from near sea level (2 m) to 770 m in primary and secondary lowland and montane wet forest and summit cloud forest. On Moen it occurs in lowland areas among mangrove swamps near sea level and on slopes and ridges where said to be common in agroforest and secondary forest. Habitat is unknown on the other islands of Chuuk.

Note. This species is characterized by its variably pubescent stems and densely hirtellous pistil and capsules. Collections from Chuuk resemble Phyllanthus senyavinianus in having a densely hirtellous ovary and style, but the pistil is comparatively shorter and only as long as the calyx lobes, and the leaves are narrowly ovate-oblong. These collections from Chuuk were previously identified as Glochidion puberulum and are here tentatively included under Phyllanthus senyavinianus, but may represent an undescribed species. Chuuk collections of Phyllanthus kanehirae differ in having female flowers with a glabrous pistil nearly twice as long as the calyx lobes. Some of these collections were previously identified as Glochidion puberulum. Certain Pohnpei collections of Phyllanthus senyavinianus were previously identified as Glochidion ramiflorum or Glochidion marianum.

59.Phyllanthus smithianusG. L. Webster, Pacific Sci. 40: 99. 1986 (1987). TYPE. FIJI ISLANDS: Viti Levu: Rewa, woods at summit of Mt. Korombamba, 381 to 427 m, 09 Jul 1968, G.L. Webster, R. Hildreth & I. Kuruvoli 14078 (holotype: DAV; isotypes: BI, SH, GH, NY, US!).

Distribution. Endemic to Fiji on the southern part of Viti Levu at 50–430 m (Smith 1981; Webster 1986).

Note. This distinctive species was treated as Phyllanthus sp. by Smith (1981, p.464).

60.Phyllanthus societatisMüll. Arg., in DC. Prodr. 15(2): 364. 1866. Diasperus societatis (Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 601. 1891. TYPE. SOCIETY ISLANDS:s. l.,1838–1842, U.S. Expl. Exped. s.n. (holotype: G-DC; isotype: US!).

Distribution. Known from the central southern Pacific region, ranging from Nauru to the Tuamotu Islands (Makatea) and Cook Islands (Aitutaki, Atiu, Mauke, and Mitiaro). Restricted to lowland calcareous substrates, usually in clearings or sunny sites in forest with Guettarda, Hibiscus, Homalium, and Pandanus (Florence 1997).

61.Phyllanthus taitensis(Baill. ex Müll. Arg.) Müll. Arg., Flora 48: 380. 1865.

Basionym.Glochidion taitense Baill. ex Müll. Arg., Linnaea 32: 66. 1863. Diasperus taitensis (Baill. ex Müll. Arg.) Kuntze, Revis. Gen. Pl. 2: 601 (1891). TYPE. SOCIETY ISLANDS: Tahiti: 1847, J. Lépine 209(holotype: G-DC; isotypes: P [3]).

Phyllanthus taitensis (Baill. ex Müll. Arg.) Müll. Arg. var. glabrescens Müll. Arg. in A.P.de Candolle, Prodr. 15(2): 301. 1866. TYPE. SOCIETY ISLANDS: Tahiti: 1838–1842, U.S. Expl. Exped. s.n. (holotype: G-DC; isotype: US!).

Glochidion ramiflorum J.R. Forst. & G. Forst. var. macrophyllum Müll. Arg., Linnaea 32: 63. 1863. Phyllanthus ramiflorus (J.R. Forst. & G. Forst.) Müll. Arg.var. macrophyllus (Müll. Arg.) Müll. Arg., Flora 48: 374. 1865. TYPE. SOCIETY ISLANDS: Tahiti: J.A. Moerenhout s.n. (lectotype: G, designated by Florence, Fl. Polynésie Française 1: 123. 1997; isolectotype: P).

Distribution. Endemic to the Society Islands of Moorea and Tahiti where widespread and common from 50 to 1500 m, occurring from lowland wet forest with Hibiscus tiliaceus and Neonauclea in valleys to mid and high elevation wet forest with Alstonia, Metrosideros, Streblus, and Weinmannia (Florence 1997).

62.Phyllanthus temehaniensis(J. W. Moore) W. L. Wagner & Lorence, comb. nov.

urn:lsid:ipni.org:names:77112729-1

Basionym.Glochidion temehaniense J. W. Moore, Bernice P. Bishop Mus. Bull. 226: 15. 1935. TYPE. SOCIETY ISLANDS: Raiatea: Temehani Plateau, 600 m, 5 Oct 1934, H. St. John 17279 (holotype: BISH-508231!; isotype: P).

Distribution. Endemic to the Society Islands (Huahine, Raiatea, and Tahaa) where it occurs from 0 to 600 m in lowland vegetation such as coconut plantations and wet valleys with Hibiscus and Neonauclea

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