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Journal ArticleDOI

A contribution to the systematics of Xylopia (Annonaceae) in Southeast Asia

01 Jan 2015-The Gardens' Bulletin, Singapore (Research Publishing Services)-Vol. 67, Iss: 02, pp 361

TL;DR: 23 Xylopia species in the Sundaic region of Southeast Asia are recognised, and evidence that additional collecting and taxonomic analysis in the region is needed is provided.

AbstractHerbarium and field study of Xylopia L. (Annonaceae) for the Flora of Peninsular Malaysia and the Flora of Thailand projects has clarified regional diversity patterns within this ecologically significant lowland rainforest genus. Two species groups represented within Southeast Asian floras are delineated, one centred on Xylopia ferruginea (Hook.f. & Thomson) Baill. and the other on Xylopia malayana Hook.f. & Thomson. In the Xylopia ferruginea group, a new species, Xylopia erythrodactyla D.M.Johnson & N.A.Murray, is distinguished from X. ferruginea, and a new combination, Xylopia sumatrana (Miq.) D.M.Johnson & N.A.Murray, is proposed, based on an earlier name for the species currently known as Xylopia stenopetala Oliv. In the Xylopia malayana group, review of the species Xylopia elliptica Maingay ex Hook.f. & Thomson resulted in the recognition of three additional species: Xylopia platycarpa D.M.Johnson & N.A.Murray, from southern Thailand and northwestern Peninsular Malaysia, Xylopia ngii D.M.Johnson & N.A.Murray, from Peninsular Malaysia, Sumatra and Borneo, and Xylopia heterotricha D.M.Johnson & N.A.Murray, from Sumatra and Borneo. The taxon Xylopia malayana Hook.f. & Thomson var. obscura Kochummen is placed in synonymy under Xylopia elliptica sensu stricto. Xylopia fusca Maingay ex Hook.f. & Thomson var. sessiliflora Kochummen & Whitmore is distinguished from Xylopia fusca, and raised to species status as Xylopia sessiliflora (Kochummen & Whitmore) D.M.Johnson & N.A.Murray. We recognise 23 Xylopia species in the Sundaic region of Southeast Asia, and provide evidence that additional collecting and taxonomic analysis in the region is needed.

Topics: Xylopia (79%), Xylopia ferruginea (78%), Southeast asian (51%)

Summary (2 min read)

Introduction

  • The Annonaceae, a flowering plant family of 2500 species, including the economically important soursop, custard apple and ylang-ylang, is widespread across the tropics.
  • Corlett & Turner (1997) determined that Annonaceae ranked fourth in species-richness among flowering plant families in Singapore; Appanah et al. (1993) found that Annonaceae ranked first in species diversity among lianas of Malaysian forests.
  • A full treatment, including keys and distribution maps, is forthcoming.
  • Conservation assessments using IUCN (2012) criteria are not included in this account as more data are required for these than the authors currently have available.

Xylopia ferruginea group

  • The Xylopia ferruginea group is characterised by stilt roots, relatively long (5-19 mm) flower pedicels, a flat receptacle lacking a staminal cone (Fig. 2K ), flat narrowly oblong stamens with a tongue-shaped apex to the anther connective (Fig. 2J ), stigmas studded with small papillae (Fig. 2I ), and rugose seeds (Fig. 2C-D ).
  • Seeds in a single row, parallel to long axis of monocarp, 6-12 per monocarp, 7.2-8.8 mm long, 5.5-5.6 mm wide, 4.7-5.1 mm thick, ellipsoid, elliptic in cross section, dark brown, rugose, flattened or a little concave at micropylar end, rounded at chalazal end, raphe and antiraphe distinctly raised; aril and aril plate absent.
  • The bulk of the Sarawak and Brunei collections have come from either lowland peat swamp forest or heath forest (terminology following Saw, 2010) .
  • These features are frequent in Sumatran specimens of Xylopia ferruginea s.s., which in general have larger and broader leaves than those of Peninsular Malaysian and Bornean plants, as well as a tendency toward a subcordate rather than truncate leaf base.
  • The leaves are retuse at the apex and broadly cuneate and decurrent at the base.

Xylopia malayana group

  • Species of the Xylopia malayana group are distinctive and well circumscribed with the exception of X. elliptica and X. malayana.
  • The protologue, reproduced verbatim below, shows that the diagnosis emphasised the glabrous branches, the small elliptic, obtuse, membranous, glabrous leaves and the solitary flowers: 14.
  • X. elliptica, Maingay mss.; branches glabrous, leaves small elliptic obtuse membranous glabrous, tip rounded, nerves faint reticulate, flowers small solitary erect pubescent, sepals subacute united to the middle, ovaries 1-3.
  • Ridley (1922) subsequently maintained King's concept of the species intact.

Xylopia platycarpa

  • Restricted to a small area of southern Thailand and northwestern Peninsular Malaysia.
  • All localities are in lowland evergreen forest.
  • Flowers in July and August, fruit collected in July.

Local name. Kerangi lotong (Kedah: Awang 42444).

  • The species is named for its unusually flattened and beanlike monocarps.
  • Twigs dark brown to brownish grey, eventually lenticellate, glabrous or finely but sparsely pubescent and soon glabrate; double-branching occasional.
  • Material of this species is usually identified in herbarium collections as Xylopia malayana, but X. ngii differs consistently from that species in its smaller (2-2.9 mm long) sepals sparsely covered with pale brown pubescence and the longer (13.5-34.2 mm) and narrower (1-1.6 mm wide at the midpoint) outer petals.

Distribution

  • The leaf blades are relatively small and oblong, often with a blotchy mottled appearance to the adaxial leaf blade surface when dried.
  • Malaysia, Negeri Sembilan, Kuala Pilah, Pasoh F.R., Compt, also known as -TYPE.
  • To confuse things further, this specimen was identified in turn as both Xylopia elliptica and X. malayana in the same publication by Corner (1978) .

Conclusion

  • Six species are widespread across the Malay Peninsula, Sumatra and Borneo: Xylopia caudata Wall. ex Hook.f. & Thomson, X. ferruginea, X. fusca, X. malayana, X. ngii and X. sumatrana.
  • Two species are shared between the Malay Peninsula and Borneo: Xylopia erythrodactyla and X. magna.
  • Several species are large trees more than 40 meters in height, and this, coupled with sparseness of individuals in populations, may cause them to be overlooked.
  • This explanation accords well with the extraordinary diversity of the genus found at sites such as the Pasoh Reserve, where in 2014 the authors verified the occurrence of seven Xylopia species in or near the Pasoh permanent 50-hectare research plot (Manokaran et al., 1992) .
  • The authors results make clear, however, that further collecting and taxonomic analysis of the Malay Peninsula flora is still critically needed.

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361
Gardens Bulletin Singapore 67(2): 361–386. 2015
doi: 10.3850/S2382581215000307
A contribution to the systematics of Xylopia
(Annonaceae) in Southeast Asia
D.M. Johnson & N.A. Murray
Department of Botany & Microbiology, Ohio Wesleyan University
Delaware, OH 43015 USA
dmjohnso@owu.edu
namurray@owu.edu
ABSTRACT. Herbarium and eld study of Xylopia L. (Annonaceae) for the Flora of Peninsular
Malaysia and the Flora of Thailand projects has claried regional diversity patterns within
this ecologically signicant lowland rainforest genus. Two species groups represented within
Southeast Asian oras are delineated, one centred on Xylopia ferruginea (Hook.f. & Thomson)
Baill. and the other on Xylopia malayana Hook.f. & Thomson. In the Xylopia ferruginea group,
a new species, Xylopia erythrodactyla D.M.Johnson & N.A.Murray, is distinguished from X.
ferruginea, and a new combination, Xylopia sumatrana (Miq.) D.M.Johnson & N.A.Murray,
is proposed, based on an earlier name for the species currently known as Xylopia stenopetala
Oliv. In the Xylopia malayana group, review of the species Xylopia elliptica Maingay ex
Hook.f. & Thomson resulted in the recognition of three additional species: Xylopia platycarpa
D.M.Johnson & N.A.Murray, from southern Thailand and northwestern Peninsular Malaysia,
Xylopia ngii D.M.Johnson & N.A.Murray, from Peninsular Malaysia, Sumatra and Borneo,
and Xylopia heterotricha D.M.Johnson & N.A.Murray, from Sumatra and Borneo. The taxon
Xylopia malayana Hook.f. & Thomson var. obscura Kochummen is placed in synonymy under
Xylopia elliptica sensu stricto. Xylopia fusca Maingay ex Hook.f. & Thomson var. sessiliora
Kochummen & Whitmore is distinguished from Xylopia fusca, and raised to species status as
Xylopia sessiliora (Kochummen & Whitmore) D.M.Johnson & N.A.Murray. We recognise 23
Xylopia species in the Sundaic region of Southeast Asia, and provide evidence that additional
collecting and taxonomic analysis in the region is needed.
Keywords. Annonaceae, biogeography, Borneo, Malay Peninsula, Sumatra, Sundaland,
Thailand, Xylopia
Introduction
The Annonaceae, a owering plant family of 2500 species, including the economically
important soursop, custard apple and ylang-ylang, is widespread across the tropics.
The family is most diverse, and ecologically most signicant, in tropical Asia, where
it is represented by c. 40 genera and 800 species. In southeastern Asia it is one of
the dominant families in lowland wet forests. Corlett & Turner (1997) determined
that Annonaceae ranked fourth in species-richness among owering plant families
in Singapore; Appanah et al. (1993) found that Annonaceae ranked rst in species
diversity among lianas of Malaysian forests. In long-term ecological plots in Southeast

362
Gard. Bull. Singapore 67(2) 2015
Asia Annonaceae usually rank among the top ten tree families in both number of
individuals and number of species, though not in basal area (see examples in Losos &
Leigh, 2004).
The genus Xylopia L., the only pantropical genus in the family, comprises
180–200 species of trees and shrubs worldwide. The highest concentration of species
in Southeast Asia occurs in the Sundaic region extending from the Kra Isthmus in
southern Thailand to Wallace’s Line, a region of high diversity for many plant taxa.
Xylopias are distinctive among the Annonaceae in their cone-shaped buds, elongate,
aromatic owers, and dehiscent fruits with seeds bearing arils or eshy seed coats. In
West Africa the peppery fruits of Xylopia aethiopica (Dunal) A.Rich. have long been
used as a spice (Dunal, 1817; Burkill, 1985) and are sold commercially. Despite its
signicance, the genus has never been monographed.
In preparing keys and descriptions of the genus Xylopia (Annonaceae) for the
Flora of Peninsular Malaysia and Flora of Thailand projects we had the opportunity to
study material in herbaria with important holdings for the region as well as to observe
several species in the eld. At the beginning of our study 13 Xylopia taxa were known
from Peninsular Malaysia, Singapore and Thailand combined (Sinclair, 1953, 1955;
Kochummen et al., 1970; Kochummen, 1972a, 1972b; Chalermglin, 2001; Gardner et
al., 2015).
Analysis of morphological and preliminary molecular data (Stull et al., 2011;
Thomas et al., 2015; Stull et al., in prep.) has shown that the Southeast Asian species of
Xylopia fall into two groups, one including Xylopia ferruginea (Hook.f. & Thomson)
Baill. and a second including Xylopia malayana Hook.f. & Thomson, each with
their respective allies. Our study revealed a much greater diversity of species than
previously recognised in both groups. To document this diversity, and reconcile it with
the previously existing taxonomy and nomenclature for the genus, the following paper
is presented. A full treatment, including keys and distribution maps, is forthcoming.
Conservation assessments using IUCN (2012) criteria are not included in
this account as more data are required for these than we currently have available.
Although historical EOOs and AOOs can be calculated, we are conscious that many of
the collections have been made in lowland forest areas that have suffered from rapid
deforestation. In these cases the Population Reduction (A) criterion would be more
appropriate, as has been used for many dipterocarp species (IUCN, 2014), but again
we would require more on-the-ground knowledge than we currently have.
Xylopia ferruginea group
The Xylopia ferruginea group is characterised by stilt roots, relatively long (5–19
mm) ower pedicels, a at receptacle lacking a staminal cone (Fig. 2K), at narrowly
oblong stamens with a tongue-shaped apex to the anther connective (Fig. 2J), stigmas
studded with small papillae (Fig. 2I), and rugose seeds (Fig. 2C–D). In addition, most
species of the Xylopia ferruginea group have relatively numerous (up to 20) linear and
somewhat torulose monocarps.

363
Xylopia in Southeast Asia
Study of the group resulted in demarcation of a new species, and recognition of
an earlier name for an existing species.
Xylopia erythrodactyla D.M.Johnson & N.A.Murray, sp. nov.
Species resembling Xylopia ferruginea in the rusty pubescence of the leaves and
owers and the long narrow monocarps, but differing consistently in the more densely
pubescent and thicker leaves, thicker pedicels (1.3–2.5 mm), longer sepals (4.8–7.9
mm), broader outer petals (3–3.7 mm wide at the midpoint) and narrowly oblong and
weakly torulose monocarps 7.5–10.7 cm long and 0.6–1.1 cm wide. In contrast, in
Xylopia ferruginea the pedicels are 1–1.3 mm thick, the sepals are 3–5.5 mm long, the
outer petals are 2–2.2 mm wide at the midpoint, and the monocarps are linear, strongly
torulose, 6.2–11.6 cm long and 0.4–0.6 cm wide. – TYPE: Malaysia, Sarawak, Teluk
Bandung, Santubong, 1st Division, 18 September 1984 (fr), Awa & Ismawi S.47080
(holotype KEP; isotypes ASU, K, L, SAR). (Fig. 1, 2)
Tree up to 30 m tall, dbh up to 75 cm, bole smooth with stilt roots at the base;
secondary branches drooping. Bark smooth, light brown, brown tinged with red,
brick-red, or orange, very nely ssured. Twigs light grey to brown, eventually
dark grey, densely ferruginous-pubescent/velutinous, eventually glabrate. Leaf with
larger blades 12.4–26 cm long, 3.9–7 cm wide, subcoriaceous to coriaceous, strongly
discolorous, oblanceolate, oblong-oblanceolate, or narrowly elliptic, base rounded to
cuneate and short-decurrent, apex short-acuminate, the acumen 2–7 mm long, glabrous
adaxially, densely ferruginous-pubescent, the pubescence especially pronounced
along the midrib, secondary veins, and larger higher-order veins abaxially; midrib
impressed adaxially, raised abaxially; secondary veins 11–14 per side, diverging at
50–60° from the midrib, brochidodromous, these and higher-order veins indistinct
adaxially, strongly raised abaxially; petiole 5–12 mm long, deeply canaliculate
(margins nearly meeting), pubescent. Inorescences axillary or from axils of fallen
leaves, 1–3-owered, densely ferruginous-pubescent; peduncles 1–2 per axil, 2 mm
long; pedicels 2 per peduncle, 7–14 mm long, 1.6–2.5 mm thick; bract 1, attached
⅓–½ distance from base of pedicel, 2.9–3 mm long, ovate, apex acute to obtuse; buds
linear-lanceolate, sometimes somewhat falciform and slightly twisted, apex obtuse.
Sepals ⅛–¼-connate, 4–7.9 mm long, 4–5.3 mm wide, coriaceous, broadly ovate to
triangular, apex acute to acuminate, occasionally obtuse, pubescent along margins
and at apex adaxially, ferruginous-pubescent abaxially. Petals pale yellow to white
in vivo; outer petals curving outward at anthesis, 38–45 mm long, 4–5.3 mm wide at
base, 3–3.7 mm wide at midpoint, linear-lanceolate, obtuse, densely grey-puberulent
adaxially, densely ferruginous-pubescent abaxially; inner petals erect at anthesis, 32–
40 mm long, c. 3.5 mm wide at base, c. 1.3 mm wide at midpoint, linear, densely
grey-puberulent on both surfaces except for glabrous concave base. Stamens up to
77, 2.7–3.5 mm long, narrowly oblong, often setose along edges of anther locules,
apex of connective 0.4–0.6 mm long, oblong, densely long-papillate, lament 0.3–0.5
mm long, glabrous; staminal cone absent; outer staminodes c. 18, c. 2.7 mm long,

364
Gard. Bull. Singapore 67(2) 2015
Fig. 1. Xylopia erythrodactyla D.M.Johnson & N.A.Murray. A. Two leaves, abaxial view. B.
Flower bud and abaxial view of monocarp, the latter showing beginning dehiscence with single
seed visible. C. Single monocarp in lateral view, resting on adaxial surface of a leaf.
(Photos: N.A. Murray)
oblanceolate, at, apex obtuse. Carpels 16–20; ovaries 1.5–2.5 mm long, narrowly
oblong, densely ferruginous-pubescent with hairs obscuring lower portion of stigmas;
stigmas loosely connivent, c. 2.2 mm long, dark, with a few scattered hairs and
studded with amber-coloured papillae. Torus at, c. 3.4 mm in diameter. Pedicel of
fruit 11–20 mm long, 4–7 mm thick, pubescent; torus of fruit c. 8 mm high, 9–14
mm in diam., depressed-globose, sparsely pubescent to glabrate. Monocarps red with
brown tomentum in vivo, up to 20 per fruit, 5.5–10.7 cm long, 0.6–1.1 cm wide and
thick, linear to narrowly oblong, weakly torulose, terete in cross-section, apex rostrate,
the beak 2.5–9 mm long, base contracted into a stipe 8–10 mm long, 3.5–4 mm thick,
longitudinally wrinkled, ferruginous-pubescent to glabrate; pericarp 1.7 mm thick.
Seeds in a single row, parallel to long axis of monocarp, 6–12 per monocarp, 7.2–8.8
mm long, 5.5–5.6 mm wide, 4.7–5.1 mm thick, ellipsoid, elliptic in cross section, dark
brown, rugose, attened or a little concave at micropylar end, rounded at chalazal end,
raphe and antiraphe distinctly raised; aril and aril plate absent.
Distribution. Occurs in Terengganu in northeastern Peninsular Malaysia, and on the
northern coast of the island of Borneo in Sarawak (East Malaysia) and in Brunei.
Considering its restricted habitat and the pace of development in Sarawak, this species

365
Xylopia in Southeast Asia
Fig. 2. Xylopia erythrodactyla D.M.Johnson & N.A.Murray. A. Habit. B. Inorescence with
ower buds, side view. C. Seed, view of micropylar end. D. Seed, side view. E. Fruit. F.
Outer petal, adaxial view. G. Inner petal, adaxial view. H. Stigma apex. I. Carpel. J. Stamen.
K. Close-up of ower with petals and stamens removed, to show sepals, carpels and torus.
L. Schematic side view of ower at anthesis, one outer petal removed. Drawn by Kate Ball
from (A) Awa & Ismawi S.47080, ASU; (B) Sibat ak Luang S.24502, L; (C–E) Chew & Kiah
SFN.40982, A; (F, G, K) Zehnder S.16803, A; (H–J) Rogstad 704, A; and (L) eld sketch.

Citations
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Journal ArticleDOI
TL;DR: The phylogenetic results suggest an African origin for the genus Xylopia and reveal complex biogeographic patterns, likely facilitated by long-distance dispersal.
Abstract: The floristic treatment of Engler and Diels, published in 1901, provides the only infrageneric classification of the pantropical genus Xylopia (Annonaceae) Here we test and extend that classification using molecular and seed morphology characters Phylogenetic relationships were reconstructed using data from four plastid regions obtained from 44 of the approximately 165 species in the genus, recovering four well-supported major clades Seed characters were examined for these taxa, and six aril morphologies, three previously undocumented, were distinguished; we also document the presence of a sarcotesta on the seeds of many species Molecular and seed data support recognition of five sections within the genus; one, Xylopia sect Rugosperma, is proposed here as new Our phylogenetic results suggest an African origin for the genus and reveal complex biogeographic patterns, likely facilitated by long-distance dispersal

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01 Jan 1924

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Journal ArticleDOI
Abstract: We present revisions of the Neotropical genera Ruizodendron and Pseudephedranthus (Annonaceae). Ruizodendron includes a single species R. ovale. Pseudephedranthus now comprises two species, with the description of the new species P. enigmaticus sp. nov. extending the range of the genus beyond the Upper Rio Negro region of Brazil (Amazonas) and adjacent Venezuela (P. fragrans), to include Guyana, Suriname, and the Brazilian state of Para. An overview is provided of current revisions of Neotropical Annonaceae genera that will aid in accessing proper species information for this frequently encountered tropical rain forest family.

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Journal ArticleDOI
TL;DR: Taxonomic review supports the hypothesis that the genus dispersed to Madagascar from continental Africa five times, and ten new species are described: Xylopia anomala D.M.Johnson & N.A.Murray, sp.
Abstract: We recognize 30 species of the pantropical genus Xylopia L. from Madagascar and an additional three species from the Mascarene Islands. Of the six sections of the genus, three are represented in the region: sect. Xylopia, sect. Verdcourtia, and sect. Stenoxylopia. All species are endemic, and many are microendemics of conservation concern. The greatest species richness occurs in humid forests below 1000 m. Taxonomic review supports the hypothesis that the genus dispersed to Madagascar from continental Africa five times. Ten new species are described: Xylopia anomala D.M.Johnson & N.A.Murray, sp. nov., X. australis D.M.Johnson & N.A.Murray, sp. nov., X. carinata D.M.Johnson & N.A.Murray, sp. nov., X. galokothamna D.M.Johnson & N.A.Murray, sp. nov., X. lokobensis D.M.Johnson & N.A.Murray, sp. nov., X. longirostra D.M.Johnson & N.A.Murray, sp. nov., X. marojejyana D.M.Johnson & N.A.Murray, sp. nov., X. ravelonarivoi D.M.Johnson & N.A.Murray, sp. nov., X. retusa D.M.Johnson & N.A.Murray, sp. nov., and X. sclerophylla D.M.Johnson & N.A.Murray, sp. nov. An identification key, technical descriptions, illustrations, and distribution maps are provided.

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Cites background from "A contribution to the systematics o..."

  • ...This section is represented by 34 species in tropical Africa (Johnson & Murray 2018), 23 species on Madagascar, and c. 40 species in Southeast Asia. noteS Many of the sect....

    [...]

  • ...Variation in climatic conditions in South and Southeast Asia from the early Miocene onward (Morley 2018), would, however, have permitted intermittent migration of Xylopia eastward, with the genus ultimately reaching New Caledonia and Fiji by long-distance dispersal (Johnson et al. 2013)....

    [...]

  • ...Rugosperma D.M. Johnson & N.A. Murray (Asia), sect....

    [...]

  • ...In conclusion, Madagascar may have played a role in the dispersal of the Stenoxylopia clade into Asia, but there is no current evidence, morphological, fossil, or molecular, to suggest that the sect....

    [...]

  • ...Densely lenticellate oblong woody monocarps are unique among Madagascar Xylopia species, and rare in the genus, although this fruit morphology is known in species such as X. hypolampra in Africa (Johnson & Murray 2018) and X. ngii in Asia (Johnson & Murray 2015)....

    [...]



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"A contribution to the systematics o..." refers background in this paper

  • ...The detached fruit on the sheet of KEP 20309 does not look like a fruit of Annonaceae, Sinclair’s determination of the specimen notwithstanding, but the leaves of the collection possibly represent this species. Turner (2011) designated the specimens K000574709 and K000574712 at K as lectotypes of the name Xylopia elliptica....

    [...]

  • ...The Annonaceae, a flowering plant family of 2500 species, including the economically important soursop, custard apple and ylang-ylang, is widespread across the tropics. The family is most diverse, and ecologically most significant, in tropical Asia, where it is represented by c. 40 genera and 800 species. In southeastern Asia it is one of the dominant families in lowland wet forests. Corlett & Turner (1997) determined that Annonaceae ranked fourth in species-richness among flowering plant families in Singapore; Appanah et al....

    [...]

  • ...Hand-written descriptive notes that are incorporated into the protologue are present on K000574709, and it is therefore designated as a second-step lectotype as permitted by Article 9.17 of the ICN (McNeill et al., 2012)....

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Abstract: VOL. 1 of this series was noticed in NATURE of January 6, 1923, p. 6. Of Vols. 2 and 3 which have now appeared, the former deals with the Gamopetalae from Caprifoliaceae to Labiatae, and the latter with the Apetalae from Nyctagineae to Salicinese. The arrangement of the natural families is that of the “Genera Plantarum “except that in the second volume the Plantagineae are inserted after the Plumbagineae, and the Cardiopteridae after the Convolvulacese. In the third volume the Aristolochiaceae and Nepenthaceae are transposed, the Hernandiaceas are separated from the Laurineae and the Opiliaceae are included. The Urticaceae are, however, still retained as one general family.The Flora of the Malay Peninsula.By Henry N. Ridley. (Published under the Authority of the Government of the Straits Settlements and Federated Malay States.) Vol. 2: Gamopetalæ. Pp. vi + 672. Vol. 3: Apetalæ. Pp. vi + 406. (London: L. Reeve and Co., Ltd., 1924.) 42s. net each vol.

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