Recircumscription of Polyscias (Araliaceae) to include six related genera, with a new infrageneric classification and a synopsis of species

Abstract

Lowry, P.P, II & Plunkett, G.M.: Recircumscription of Polyscias (Araliaceae) to include six related genera, with a new infrageneric classification and a synopsis of species. — Plant Div. Evol. 128: 55-84. 2010.— ISSN 1869-6155. The genus Polyscias, described more than two centuries ago, has gone through several cycles of ex-pansion and contraction in its circumscription, settling in the last forty years on a broad consensus that includes nearly all Paleotropical Araliaceae with pinnately compound leaves, an articulated pedicel and a gynoecium with two to many carpels. Recent phylogenetic studies have shown that Polyscias is closely related to six other long-recognized genera (Arthrophyllwn, Cuphocarpus, Gastonia, Munroi-dendron, Reynoldsia and Tetraplasandra). Because these other genera are all nested within a broad Polyscias sensu lato clade, substantial generic-level re-alignment is needed to avoid recognition of para-and polyphyletic groups. An earlier suggestion that each clade be recognized as a separate genus has proven impractical in light of an expanded phylogenetic analysis, which identified more than a dozen geographically coherent clades, some of which encompass significant morphological diversity but lack obvious defining synapomorphies. A more pragmatic approach is adopted here, in which all members of Polyscias sensu lato are treated as a single genus. Within this broadly defined genus, each clade is recognized as a subgenus, thereby retaining valuable information on evolutionary relation-ships while minimizing nomenclatural changes. A revised system of classification is proposed in which 159 currently recognized species are placed in Polyscias and assigned to 10 subgenera (seven species are left unassigned), necessitating 65 new combinations (nine at the rank of subgenus, 54 for species, and two for subspecies) and five new names.

Full text

Plant Div.
Evol.
Vol. 128/1-2, 55-84
Stuttgart, August 20, 2010
Recircumscription of
Polyscias
(Araliaceae) to
include six related genera, with a new infrageneric
classification and a synopsis of species
By Porter P. Lowry II and Gregory M. Plunkett
With 4 tables
Abstract
Lowry, P.P, II & Plunkett,
G.M.:
Recircumscription of Polyscias (Araliaceae) to include six related
genera, with a new infrageneric classification and a synopsis of species. — Plant Div. Evol. 128: 55-
84.
2010.— ISSN 1869-6155.
The genus Polyscias, described more than two centuries ago, has gone through several cycles of ex-
pansion and contraction in its circumscription, settling in the last forty years on a broad consensus that
includes nearly all
Paleotropical
Araliaceae with
pinnately
compound
leaves,
an articulated pedicel
and a gynoecium with two to many carpels. Recent
phylogenetic
studies have shown that Polyscias is
closely related to six other long-recognized genera
(Arthrophyllwn,
Cuphocarpus,
Gastonia,
Munroi-
dendron,
Reynoldsia and Tetraplasandra). Because these other genera are all nested within a broad
Polyscias sensu
lato
clade,
substantial generic-level re-alignment is needed to avoid recognition of
para- and polyphyletic groups. An earlier suggestion that each clade be recognized as a separate genus
has proven impractical in light of an expanded phylogenetic analysis, which identified more than a
dozen geographically coherent clades, some of which encompass significant morphological diversity
but lack obvious defining
synapomorphies.
A more pragmatic approach is adopted here, in which all
members of Polyscias sensu lato are treated as a single genus. Within this broadly defined genus, each
clade is recognized as a subgenus, thereby retaining valuable information on evolutionary
relation-
ships while minimizing nomenclatural changes. A revised system of classification is proposed in
which
159
currently recognized species are placed in Polyscias and assigned to
10
subgenera (seven
species are left unassigned), necessitating 65 new combinations (nine at the rank of subgenus, 54 for
species, and two for subspecies) and five
new
names.
Keywords: Araliaceae,
Polyscias,
infrageneric classification, subgenus,
Arthrophyllum,
Cuphocar-
pus,
Gastonia,
Munroidendron,
Reynoldsia, Tetraplasandra.
Introduction
The genus Polyscias was described by Forster & Forster
(1775)
based on a single col-
lection they made in the South Pacific while serving on board Captain James Cook's
second voyage. The Forsters distinguished their new genus from other Araliaceae rec-
Received
February 27,
2009.
in revised form June 22, 2009. accepted June 27, 2009
©
2010
E.
Schweizerbart'sche
Verlagsbuchhandlung, Stuttgart, Germany www.schweizerbart.de
DOI:
10.1127/1869-6155/2010/0128-0003 1869-6155/2010/0128-0003
$07.50

56 P.P. Lowry II &
G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae)
ognized at the time (primarily the Linnaean
genera
Aralia L., Hedera L. and Panax L.)
by several characters including the presence of
6
to 8
petals
and as many stamens, 3 or
4 styles and a globose fleshy fruit containing 4 seeds. Over the next 90 years no addi-
tional species had been assigned to
Polyscias,
although during the same period over 60
taxa were described, mostly in the genus
Panax,
that are currently included in Polys-
cias by Frodin & Govaerts (2003), either as accepted species or synonyms. Between
1865
and
1894,
the name Polyscias was applied to
species
just eight times, primarily
by Baker
(1877)
for taxa occurring in the Mascarene Islands. Harms
(1894-97)
was the
first to adopt a broader circumscription
of
Polyscias, recognizing a total of
41
species,
including 33 for taxa he transferred from other genera (mostly from Panax). Only a
few authors, however, adopted his approach in the following decades.
A subsequent shift toward a narrower definition of Polyscias began at the start of
the
20lh
Century with
Viguier
(1905),
who resurrected
Cuphocarpus
Decne. &
Planch,
and Sciadopanax Seem., and described two new genera, Bonnierella R. Vig. and
Tieghemopanax
R. Vig., while transferring some other taxa to Polyscias that had been
published in the preceding decade or that Harms had overlooked. Hutchinson (1967)
went even further, adopting all the same genera as Viguier along with three others that
Harms had reduced into synonymy
{Botryopanax
Miq.,
Eupteron
Miq.,
and Sciadopa-
nax) plus a fourth that he described as new (Gelibia Hutch.). At about the same time,
the trend reversed again toward defining Polyscias more broadly, starting with Smith
& Stone (1965) and then Bernardi
(1971),
who placed in synonymy nearly all of the
segregate genera accepted by Viguier and Hutchinson, along with Palmervanden-
broekia Gibbs and
Kissodendmn
Seem. Philipson (1977a, 1978, 1979) used a simi-
larly broad circumscription of Polyscias in his treatment for Flora Malesiana, for-
mally recognizing four sections within the region and transferring two Australian
species previously misplaced in Pentapanax. This broad definition has been widely
followed since (e.g., Marais 1984; Smith
1985;
Lowry
1989;
Lejoly & Lisowski 1999;
Lowry et
al.
1999;
Frodin & Govaerts
2003;
Callmander
et
al.
2009).
In a series of recent molecular phylogenetic studies (Plunkett et al. 2001, 2004a,
2004b), we identified several major
clades
within Araliaceae, one of which comprises
Polyscias and a number of related genera, collectively referred to as Polyscias sensu
lato.
These studies also identified several major, well-supported
clades
within Polys-
cias sensu lato along with a series of smaller clades comprising one or a few species.
They further revealed that Polyscias, as currently circumscribed, is paraphyletic with
respect to six genera that have been traditionally regarded as distinct by most authors
(viz.
Artkrophyllum
Blume,
Cuphocarpus, Gastonia
Comm.
ex Lam.,
Munroidendron
Sherff Reynoldsia
A. Gray and
Tetraplasandra
A. Gray), most of which have rarely if
ever
been associated directly with Polyscias, and three of which are themselves
poly-
phyletic {Cuphocarpus, Gastonia and Reynoldsia).
These phylogenetic results clearly showed that traditional generic definitions would
have to be completely overhauled in order to circumscribe monophyletic groups and
provide a classification system that accurately reflects evolutionary relationships. Ini-
tially (Plunkett et al. 2001, Lowry
ct
al. 2004) we suggested that each of the major
clades
of
Polyscias sensu lato could be treated as a single genus, pointing out that such
an approach would highlight phylogenetic relationships rather than obscure them.

P.P. Lowry II & G.M. Plunkett,
Recircumscription
of Polyscias (Araliaceae) 57
However, in light of the conclusions reached in an expanded phylogenetic analysis of
molecular data, presented in a companion paper (Plunkett & Lowry
2010),
it is now
evident that this approach presents serious difficulties. First, to avoid recognition of
paraphyletic groups, many of the clades that would serve as potential genera are diffi-
cult to define on the basis of morphology and present few if any obvious synapomor-
phies,
a fact that would significantly compromise the utility of
the
resulting classifica-
tion. Moreover, such an approach would require publication of more than 100 new
combinations in order to place each species in its corresponding genus. Based on these
considerations, we have therefore elected instead to treat all members of the Polyscias
sensu
lato clade
as a single genus (see Plunkett & Lowry 2010). Adopting a broad ge-
neric circumscription is considerably less disruptive to nomenclatural stability, requir-
ing a total of 54 new combinations and five new names to bring all currently recog-
nized species into Polyscias. Furthermore, as mentioned in our companion paper,
defining Polyscias broadly avoids potential problems associated with the treatment of
species that can not confidently be assigned to a clade (and thus to a distinct genus)
based on either molecular or morphological data, including several currently recog-
nized species (e.g., Polyscias mollis,
P.
murrayi,
P.
purpurea) plus many new species,
mostly from Madagascar and New Caledonia, awaiting formal description.
While several of
the
clades identified in our phylogenetic study (Plunkett & Lowry
2010) are not well suited for recognition at the generic level, many of them contain
important information on evolutionary relationships and all exhibit strong geographic
structuring. In an attempt to translate the
phylogeny
of Polyscias into a workable clas-
sification system that retains this potentially useful information, we have, for several
reasons, chosen to recognize each of these clades at the subgeneric level. First, by do-
ing so we retain the option of formally recognizing sections within the subgenera,
several of which appear to contain morphologically distinctive species groups that
may prove to be
monophyletic.
Also, we avoid possible confusion with the incomplete
system of four sections proposed by Philipson (1978, 1979) for the species of Polys-
cias he recognized in Malesia, several of which are of limited use in other parts of the
world and do not comprise monophyletic groups (see below). Moreover, because we
initially considered recognizing these groups as distinct genera (Plunkett et
al,
2001,
Lowry et al. 2004), treating them as subgenera would seem appropriate. Finally, rec-
ognizing subgenera within Polyscias is consistent with the approach we are adopting
for other Araliaceae, such as some of the clades currently placed in the polyphyletic
genus
Scheffiera
J.R. Forst. & G. Forst.
Table
1
summarizes the correspondence between the major clades identified in our
companion paper and the ten subgenera presented below. Some of these groups are
well delimited and their members can be easily recognized on the basis of obvious
morphological features (as indicated below in the discussion under each subgenus).
Others, however, are more difficult to characterize, such as subgenus Maralia, which
includes nearly all of the species from Madagascar and encompasses an impressive
level of morphological diversity (much of which remains to be described), presumably
the result of extensive local radiation and diversification. Nonetheless, each of the
subgenera recognized in our classification system is geographically coherent and com-
prises a well supported monophyletic assemblage.

58 P.P. Lowry II & G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae)
Table 1. Correspondence between the major
clades
within Polyscias sensu
lato
identified by Plunkett
& Lowry
(2010)
and the subgenera
of
Polyscias recognized in the present classification system.
Clade
name used by Plunkett & Lowry
(2010)
Corresponding subgenus in the present
classification system
Section Polyscias clade A. Polyscias subg. Polyscias
Indian Ocean Basin
(IOB) clade/Mascarenes
subclade B. Polyscias subg. Grotenfendia
IOB clade/Malagasy
subclade C. Polyscias subg.
Maralia
Austro-Malesian clade D. Polyscias subg. Arthrophyllum
IOB
clade/Cuphocarpus
subclade E. Polyscias subg.
Cuphocarpus
New Guinea-Polynesia clade F. Polyscias subg. Tetraplasandra
Polyscias nodosa clade G. Polyscias subg.
Euptcron
IOB
clade/.P/»/v«
subclade H. Polyscias subg. Sciadopanax
Tieghemopanax
clade I. Polyscias subg.
Tieghcmopanax
Seychelles clade J. Polyscias subg.
Indokingia
The classification system presented below is synoptic in form. Full synonymy is
provided for each subgeneric name along with place of publication and an indication
of
the
nomenclatural type (when necessary the corresponding species currently recog-
nized in Polyscias is also given). At the species level, however, Frodin & Govaerts
(2003) have already provided full synonymy for all species of Araliaceae recognized
in their World Checklist and Bibliography. Thus, we have largely refrained from re-
peating this information except for synonyms that are currently in wide use. We for-
mally place ten species in synonymy that were accepted as distinct by Frodin and
Table 2. Species recognized by Frodin & Govaerts (2003) but placed in synonymy in the present clas-
sification.
Name
accepted
by
Frodin & Govaerts (2003) Placement in synonymy in the present classification
Arthrophyllum angustatum
(Baill.)
Philipson
Arthrophyllum daenikeri (Baum.-Bod.)
Philipson
Arthrophyllum glaberrimum (Baum.-Bod.)
Philipson
Arthrophyllum hederoides (Baum.-Bod.)
Philipson
Arthrophyllum schlechteri (Harms) Philipson
Polyscias grandifolia Volkens
Polyscias
tennantii Bernardi
Reynoldsia grayana Christoph.
Reynoldsia tahitiensis Nadeaud
Reynoldsia tauensis A. C. Smith & B. C.
Stone
Polyscias otopyrena (Baill.) Lowry & G. M. Plunkett
Polyscias
vieillardii
(Baill.) Lowry & G.
M.
Plunkett
subsp. halansae (Baill.) Lowry & G. M. Plunkett
Polyscias vieillardii (Baill.) Lowry & G.
M.
Plunkett
subsp. halansae (Baill.) Lowry & G. M. Plunkett
Polyscias vieillardii (Baill.) Lowry & G.
M.
Plunkett
subsp. halansae (Baill.) Lowry & G.
M.
Plunkett
Polyscias otopyrena (Baill.) Lowry & G. M. Plunkett
Polyscias macgillivrayi (Seem.) Harms
Polyscias chapelieri (Drake) Harms ex R. Vig.
Polyscias lanutoensis (Hochr.) Lowry & G. M.
Plunkett
Polyscias verrucosa (Seem.) Lowry & G.
M.
Plunkett
Polyscias lanutoensis (Hochr.) Lowry & G. M.
Plunkett
C
tl
r;
('
B
S
i
t

P.P.
Lowry
II &
G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae) 59
Table 3. Species and
infraspecific
taxa recognized by Frodin & Govaerts (2003) for which a new name
or a change in rank is proposed in the present
classification.
Name
used by
Frodin & Govaerts (2003) Name
adopted
in the current classification
Arthrophyllum
balansae
(Baum.-Bod.)
Philipson
Arthrophyllum crassum Philipson, non
Polyscias crassa
(Hemsl.)
Lowry &
G. M. Plunkett
Arthrophyllum ferrugineum Craib, non
Polyscias
jerruginea
(Hiern) Harms
Arthrophyllum grandifolium
(Guillaumin)
Philipson, non Polyscias
grandifolia
Volkens
Cuphocarpus commersonii Bernardi, non
Polyscias commersonii (Drake) R. Vig.
Gastonia elegans (W. Bull) Frodin, non
Polyscias elegans (C. Moore & F.
Muell.)
Harms
Polyscias australiana (F. Muell.) Philipson
var. disperma (F. Muell.) Philipson
Polyscias subincisa (R. Vig.) Lowry
Polyscias vieillardii
(Baill.)
Lowry & G. M. Plunkett
subsp. balansae (Baill.) Lowry & G. M. Plunkett
Polyscias revoluta (Philipson) Lowry & G. M.
Plunkett
Polyscias thailandica Lowry & G. M. Plunkett
Polyscias mackeei Lowry & G. M. Plunkett
Polyscias compacta Lowry & G. M. Plunkett
Polyscias maraisiana Lowry & G. M. Plunkett
Polyscias disperma (F. Muell.) Lowry & G. M.
Plunkett
Polyscias bracteata (R. Vig.) Lowry subsp.
subincisa (R. Vig.) Lowry & G. M. Plunkett
Govaerts (2003) (see Table 2) and do likewise for seven infraspecific taxa that we do
not recognize. In addition to proposing five new names, we also raise one variety to the
rank of species and reduce two previously recognized species to the level of subspecies
(Table 3). Full information is provided on the
basionym
for each of
the
65 new combi-
nations made here, which include nine for infrageneric taxa, 54 for species, and two for
subspecies. We have designated lectotypes for infrageneric taxa as needed in order to
improve nomenclatural stability. Full citation of types for species and infraspecific
taxa (and designation of lectotypes where appropriate) will, however, be provided
along with listings of exsiccatae, synonymy, and other information in a forthcoming
series of taxonomic revisions being prepared for each subgenus. A total of 59 names
accepted by Frodin & Govaerts (2003) in the genera Arthrophyllum, Cuphocarpus,
Gastonia, Munroidendron, Reynoldsia and Tetraplasandra are treated here in Polys-
cias.
As an aid to those wishing to adopt our names, Table 4 indicates the correspon-
dence between those used by Frodin & Govaerts (2003) and the ones we have adopted
here.

60 P.P. Lowry II & G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae)
Table 4. Index
lo
names accepted by
Frodin
& Govaerts (2003) in genera other than Polyscias and their place-
ment in the present classification.
Name
used by Frodin & Govaerts (2003) Current name in Polyscias in the present classification
Arthrophyllum alternifolium Maingay
ex
Ridl.
Arthroph)
Arthroph}
Arthroph)
Arthroph}
Arthroph}
Arthroph}
Arthrophyi
Arthrophyi
Arthroph}
Arthroph}
Arthroph}
Arthroph}
Arthroph}
Arthroph}
Arthroph}
Arthroph}
Arthrophyi
Arthrophyi
Arthroph}
Arthroph}
Arthroph}
Arthrophy
Arthrophyi
A
rthrophy
Arthrophy
Arthrophy
Arthrophy
Arthrophy
Arthrophy
htm angustifolium Ridl.
lum
ashtonii Philipson
'him balansae (Baill.)
Philipson
lum biformc Philipson
him
borneense Baker
lum
cenabrei
Merr.
lum collinum Philipson
lum crassum Philipson
lum
diversifolium Blume
lum engganoense Philipson
lum ferrugineum Craib
hint havilandii
Ridl.
him jackianum
(G.Don) Frodin
lumjavanicum
Blume
lum kjellbergii Philipson
'him lucens
Craib
lum macranthum Philipson
lum macrocarpum Philipson & Bui
him
meliifolium Craib
lum
montanum
Ridl.
lum otopyrenum (Baill.) Philipson
lum
pacificum Philipson
lum papyraccum Philipson
lum
prolifcrum
Philipson
lum
pulgarense
Elmer
lum ruhiginosum Ridl.
him
rufosepalum Ridl.
lum stonei A. L.
Lim
lum vicillardii (Baill.) Philipson
Cuphocarpus briquetianus Bernardi
Cuphocarpus commersonii Bernardi
Cuphocarpus
humbertianus
Bernardi
Cuphocarpus
leandrianus
Bernardi
Gastonia
crassa
(Hcmsl.)
F. Friedmann
Gastonia cutispongia Lam.
Gastonia duplicata Thouars ex Baill.
Gastonia
elegans
(W. Bull) Frodin
D2.
Polyscias
alternijolia
(Maingay ex Ridl.) Lowry &
G. M. Plunkett
D2.
Polvscias
angustijolia
(Ridl.) Lowry & G. M. Plunkett
D4.
Polvscias ashtonii (Philipson) Lowry & G. M. Plunkett
D36b.
Polyscias vicillardii (Baill.) Lowry & G. M. Plunkett
subsp.
balansae
(Baill.) Lowry & G.
M.
Plunkett
D7.
Polyscias biformc (Philipson) Lowry & G.
M.
Plunkett
Dl.
Polyscias aherniana (Philipson) Lowry & G.
M.
Plunkett
D9.
Polyscias
cenabrei
(Merr.) Lowry & G. M. Plunkett
D10.
Polyscias collina (Philipson) Lowry & G. M. Plunkett
D.l
1
Polyscias crassa (Philipson) Lowry & G. M. Plunkett
D13.
Polyscias diversifolia (Blume) Lowry & G. M. Plunkett
D15.
Polyscias engganoense (Philipson) Lowry &
G. M. Plunkett
D33.
Polyscias
siamensis
Lowry & G. M. Plunkett
D16.
Polyscias havilandii (Ridl.) Lowry & G. M. Plunkett
D17.
Polyscias
jackiana
(G. Don) Lowry & G. M. Plunkett
1)14.
Polyscias elliptica (Blume) Lowry & G. M. Plunkett
D18.
Polyscias kjellbergii (Philipson) Lowry & G. M.
Plunkett
D19.
Polyscias lucens (Craib) Lowry & G. M. Plunkett
D21.
Polyscias macranthum (Philipson) Lowry & G. M.
Plunkett
D22.
Polyscias macrocarpa (Philipson) Lowry & G. M.
Plunkett
D23.
Polyscias
meliifolia
(Craib) Lowry & G. M. Plunkett
D24.
Polyscias montana (Ridl.) Lowry & G. M. Plunkett
D25.
Polyscias otopyrena (Baill.) Lowry & G. M. Plunkett
D26.
Polyscias pacifica (Philipson) Lowry & G. M. Plunkett
D27.
Polysciaspapyracea
(Philipson) Lowry & G. M.
Plunkett
D28.
Polysciasprolifera
(Philipson) Lowry & G. M. Plunkett
D29.
Polyscias pulgarense (Elmer) Lowry & G.
M.
Plunkett
D31.
Polyscias rubiginosa (Ridl.) Lowry & G. M. Plunkett
D32.
Polyscias rufosepalum (Ridl.) Lowry & G. M. Plunkett
D35.
Polyscias stonei (Ridl.) Lowry & G. M. Plunkett
D36b.
Polyscias vicillardii (Baill.) Lowry & G. M. Plunkett
subsp. vicillardii
C7.
Polyscias briquetiana (Bernardi) Lowry & G. M. Plunkett
CI
1. Polyscias
compacta
Lowry & G. M. Plunkett
C19.
Polyscias humbertiana (Bernardi) Lowry & G. M.
Plunkett
C22.
Polyscias leandriana (Bernardi) Lowry & G. M.
Plunkett
Dll.
Polyscias crassa (Philipson) Lowry & G.
M.
Plunkett
B5.
Polyscias cutispongia (Lam.) Baker
C14.
Polyscias duplicata (Thouars ex Baill.) Lowry & G. M.
Plunkett
B8.
Polyscias maraisiana Lowry & G. M. Plunkett
Re\

P.P. Lowry II & G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae) 61
Gastonia lionnetii
F.
Friedmann
Gastonia rodriguesiana Marais
Gastoniasechellarum
(Baker) Harms
Gastonia serratifolia
(Miq.) Philipson
Gastonia spectabilis (Harms) Philipson
Munroidendron
racemosum (C. N.
Forbes)
Sherff
Reynoldsia grayana Christoph.
Reynoldsia
lanutoensis
Hoehr.
Reynoldsia marchionensis F. Br.
Reynoldsiapleiosperma
A. Gray
Reynoldsia
sandwicensis
A. Gray
Reynoldsia
tahitensis
Nadeaud
Reynoldsia tauensis A. C.
Sm.
& B. C. Stone
Reynoldsia verrucosa Seem.
Tetraplasandra flynnii Lowry &
K.
R. Wood
Tetraplasandra gymnocarpa (Hillebr.) Sherff
Tetraplasandra
hawaiensis
A. Gray
Tetraplasandra
kavaiensis
(H. Mann) Sherff
Tetraplasandra oahuensis (A. Gray) Harms
Tetraplasandra
waialealae
Rock
Tetraplasandra
waimeae
Wawra
J12.
Polyscias lionnetii (F. Friedmann) Lowry & G. M.
Plunkett
Bl4.
Polyscias rodriguesiana (Marais) Lowry & G. M.
Plunkett
J3.
Polyscias
seyeheltarum
(Baker) Lowry & G. M. Plunkett
F16.
Polyscias serratifolia (Miq.) Lowry & G. M. Plunkett
Fl
7. Polyscias spectabilis (Harms) Lowry & G. M. Plunkett
F14.
Polyscias racemosa (Forbes) Lowry & G. M. Plunkett
F9.
Polyscias lanutoensis
(Hochr.)
Lowry & G. M. Plunkett
F9.
Polyscias lanutoensis (Hochr.) Lowry & G. M. Plunkett
F10.
Polyscias marchionensis (F. Brown) Lowry & G. M.
Plunkett
F13.
Polyscias pleiosperma (A. Gray) Lowry & G. M.
Plunkett
F15.
Polyscias sandwicensis (A. Gray) Lowry & G. M.
Plunkett
F18.
Polyscias verrucosa (Seem.) Lowry & G. M. Plunkett
F9.
Polyscias lanutoensis (Hochr.) Lowry & G. M. Plunkett
Fl8.
Polyscias verrucosa (Seem.) Lowry & G. M. Plunkett
F5.
Polyscias flynnii (Lowry &
K..
R. Wood) Lowry & G. M.
Plunkett
F6.
Polyscias
gymnocarpa
(Hillebr.) Lowry & G. M. Plunkett
F7.
Polyscias
hawaiiensis
(A. Gray) Lowry & G. M. Plunkett
F8.
Polyscias kavaiensis (H. Mann) Lowry & G. M. Plunkett
Fl
1.
Polyscias oahuensis (A. Gray) Lowry & G. M. Plunkett
F19.
Polyscias waialealae (Rock) Lowry & G. M. Plunkett
F20.
Polyscias waimeae (Wawra) Lowry & G. M. Plunkett
Infrageneric classification
Polyscias J. R. Forst. & G. Forst., Char. Gen. PL: 32. 1775. — Type:
P.
pinnata J. R.
Forst. & G. Forst. [= Polyscias Scutellaria (N. L.
Burm.)
Fosberg].
A. Polyscias subg. Polyscias
Nothopanax Miq. in Bonplandia 4: 139. 1865.
[=
Polyscias fruticosa L.].
Botmierella
R. Vig. in Bull.
Soc.
Bot. France 52: 314. 1905.
tiense Nadeaud
[=
Polyscias tahitiensis (Nadeaud) Harms].
Type: N.fruticosus (L.) Miq.
Type: B. tahi-
Mcmbers
of
this
subgenus form a geographically and morphologically coherent group
that corresponds to Polyscias sensu stricto or Polyscias sect. Polyscias as defined by
Philipson
(1978,
1979, 1995). They are characterized by having petioles with an elon-
gated sheathing base whose margins are often scarious or membranous and sometimes
alate.
Dried specimens of most species in this subgenus also have a characteristic pun-
gent odor reminiscent of aniseed or cumin (a feature that is not, however, restricted to
the group). Viguier
(1905)
included at least one species belonging to Polyscias subg.

62 P.P.
Lowry
II & G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae)
Polyscias in his new genus
Tieghemopanax
because of the presence of
a
bicarpellate
gynoecium
(P.
macgillivrayi) and
tentatively
referred three others there as well (see
discussion below under subgenus Tieghemopanax). As circumscribed here, the native
ranges of species in Polyscias subg. Polyscias collectively extend from Java across
Malesia to tropical Australia, Micronesia, and Melanesia, S to Norfolk
Isl.
and across
Polynesia to Tahiti, the eastern limit of the entire genus.
Several taxa belonging to Polyscias subg. Polyscias, especially P.
cumingiana,
P.
fruticosa,
P.
guilfoylei and
P.
Scutellaria,
are widely cultivated throughout the trop-
ics,
often as living hedgerows. While these taxa are well represented in herbaria, their
native ranges are unknown and few if any collections appear to represent material col-
lected from native habitat. Moreover, although the cultivated taxa can almost always
be distinguished from one another without difficulty, each exhibits significant varia-
tion, especially in leaf morphology and color (e.g., numerous variegated and
dissected
forms exist), which has
led
to the publication of many names. Taxonomic limits within
these entities remain controversial and opinions have varied among recent authors
regarding how best to circumscribe them. Philipson (1978, 1979) treated species
broadly whereas several of
the
names he placed in synonymy were accepted as distinct
taxa by Frodin & Govacrts (2003), including P. balfouriana (Andre) L. H. Bailey.
P.
filicifolia (C. Moore & E. Fourn.) L. H. Bailey,
P.
obtusifolia Frodin,
P.
pinnata J. R.
Forst. & G. Forst. and
P.
sorongensis
Gibbs.
Lowry (1989) adopted Philipson's ap-
proach and we have done so here as well, preferring to defer a more detailed consider-
ation of species limits until additional data, especially from molecular analyses, be-
come available.
Taxa included:
Al.
Polyscias corticata Gibbs in J. Linn.
Soc.,
Bot. 39:
149.
1909.
— Fiji.
A2.
Polyscias cumingiana
(K.
Presl)
Fern.-Vill.,
Nov. App.: 102.
1880.
— Native
range unknown, widely cultivated in the Pacific and elsewhere. — Frodin &
Govacrts
(2003) treat Polyscias sorongensis Gibbs as distinct from
P.
cumin-
giana,
whereas Philipson (1979, 1995) and Lowry
(1989)
include it therein as
a synonym. We prefer to follow this broad definition until a more detailed study
can
be
conducted of this complex group.
A3.
Polyscias
fruticosa (L.) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3(8): 45.
1894.
— Native range unknown, widely cultivated in the Pacific and else-
where.
A4.
Polyscias guilfoylei (Bull) L. H. Bailey in Rhodora 18: 153. 1916. — Native
range unknown, widely cultivated in the Pacific and elsewhere.
A5.
Polysciasjavanica
Koord.
& Valeton, Bijdr. 7: 13. 1900. — Java, Lesser Sunda
Isl.
A6.
Polyscias macgillivrayi
(Seem.)
Harms in Engl. & Prantl, Nat. Pflanzenfam.
3(8):
45.
1894.
— Micronesia, Solomon
Isl.,
Queensland, New Guinea.
= Polyscias grandifolia Volkens in Bot. Jahrb. Syst. 31: 471. 1901, syn.
nov.
A7.
Polyscias multijuga (A. Gray) Harms in Engl. & Prantl, Nat. Pflanzenfam.
3(8):
45.
1894.
— Vanuatu, Fiji, Wallis & Futuna, Norfolk Isl.
A8.
Polyscias reineckei Harms in Bot. Jahrb. Syst. 25: 663.
1898.
— Samoa.

P.P. Lowry II & G.M. Plunkett,
Recircumscription
of Polyscias (Araliaceae) 63
A9.
Polyscias
samoensis
(A. Gray) Harms in Engl. & Prantl, Nat.
Pflanzcnfam.
3(8):
45.
1894.
— Samoa, Vanuatu.
A10.
Polyscias Scutellaria (N.L.
Burm.)
Fosberg in Univ. Hawaii
Occ.
Papers
46:
9.
1948 [= Crassula Scutellaria
Burm.f.,
Fl.
Indica:
78. 1768.] — Native range
unknown, widely cultivated in the Pacific and elsewhere. — Frodin & Govaerts
(2003) regard Polyscias pinnata J. R. Forst. & G.
Forst,
the generi-type, as
distinct from
P.
Scutellaria, but we opt to treat the latter broadly, following
Philipson (1979, 1995) and Lowry (1989), at least until more detailed studies
can be conducted to elucidate
relationships
and improve the definition of enti-
ties within this complex assemblage.
All.
Polyscias subcapitata
Kaneh.
in Bot. Mag. (Tokyo) 46: 673.
1932.
— Caroline
[si.
A12.
Polyscias tahitiensis (Nadeaud) Harms in Engl. &
K.
A. E. Prantl, Nat.
Pfian-
zenfam.
3(8): 45. 1894.
P.
reflexa J. W. Moore in Bernice P. Bishop Mus. Bull. 102: 35. 1933
[s
Bon-
nierella
reflexa
(J. W. Moore) J. W. Moore in
Occas.
Pap. Bernice Pauahi Bish-
op Mus. 10: 6.
1934].
—Tahiti,
Raiatea.
A13.
Polyscias verticillata Stone in J. Arnold Arbor. 47: 272. 1966.
—
Solomon
Islands, New Guinea.
B.
Polyscias subg.
Grotenfendia
(Seem.) R. Vig. in Bull.
Soc.
Bot. France 52:
30E 1905.
=
Grotenfendia
Seem,
in J. Bot. 2: 247. 1864.
— Lectotype
(here
designated): Polyscias repanda (DC.) Baker.
Gastonia
Comm.
ex Lam., Encycl. 2: 610. 1788.
—Type:
G cutispongia Lam.
[= Polyscias cutispongia (Lam.) Baker].
Botryopanax Miq. in Ann. Mus. Bot. Lugduno-Batavi 1: 5. 1863. — Type:
B.
borbonica Miq. [= Polyscias
paniculata
(DC.) Baker].
Polyscias subg. Grotenfendia, lectotypified here by the Reunion Island endemic
P.
repanda, corresponds to the Mascarene Islands subclade of the Indian Ocean Basin
cladc
in the updated phylogeny of Plunkett & Lowry (2010). It comprises all
15
mem-
bers of the Polyscias sensu
lato
group occurring in the Mascarenes, including taxa with
articulated pedicels historically placed in Polyscias and others lacking an articulation
that have long been referred to Gastonia (which is typified by another Reunion en-
demic,
P.
cutispongia). The oldest generic name that applies to this
clade
is Gastonia,
but when it is treated as a subgenus, as we do here, Viguier's combination based on
Grotenfendia has
nomenclatural
priority.
As indicated in our companion paper (Plunkett & Lowry
2010),
Gastonia as rede-
fined by Philipson (1970) is highly polyphyletic, comprising taxa
belonging
to four
distinct subclades within Polyscias sensu lato. As circumscribed here, Polyscias subg.
Grotenfendia bears little resemblance to Gastonia as historically treated (e.g., by
Harms
1894-97)
or as redefined by Philipson (1970,
1979,
1995)
and adopted by many
recent authors (e.g.,
Bernardi
1971,1980; Friedmann 1986; Frodin & Govaerts 2003).
Instead, it represents a well supported and geographically coherent group that appears
to have diversified after reaching the Mascarenes via long-distance dispersal from
Madagascar (Mauritius, the oldest island in the archipelago, dates from ca. 10 my;

64 P.P. Lowry II &
G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae)
Schliiter
2006). All members of the subgenus have radiating style arms, a feature that
is not unique to species from the
Mascarenes
(see Philipson 1970), but which may
nevertheless be
of taxonomic
value in combination with geography.
Taxa included:
Bl.
Polyscias
aemiliguineae
Bernardi in
Candollea
29: 153.
1974.
—
Reunion.
B2.
Polyscias bernieri
(Baill.
ex Drake) R. Vig. in Bull.
Soc.
Bot. France 52: 303.
1905.
—Reunion.
B3.
Polyscias borbonica Marais in
Kew
Bull. 39:
814.
1984.
— Reunion.
B4.
Polyscias coriacea Marais in Kew Bull. 39:
812.
1984.
— Reunion.
B5.
Polyscias
cutispongia (Lam.)
Baker,
Fl.
Mauritius Seych.:
127.
1877 [=
Gasto-
nia
cutispongia
Lam.].
— Reunion.
B6.
Polyscias
dichroostachya
Baker, Fl. Mauritius Seych.: 127. 1877. — Mauri-
tius.
B7.
Polyscias gracilis Marais in Kew Bull.
39:811.
1984.
— Mauritius.
B8.
Polyscias maraisiana Lowry & G. M. Plunkett,
nom.
nov. =
Gastonia
elegans
(W. Bull) Frodin, in D.
Frodin
& R. Govaerts, World Checklist Bibliog. Ar-
aliaceae: 166. 2003
[= Terminalia
elegans W. Bull, Cat. 14: 8.
1866,
non Poly-
scias elegans (C. Moore & F
Muell.)
Harms. — Mauritius.
= Gastonia mauritiana Marais, non Polyscias
mauritiana
Marais.
B9.
Polyscias
mauritiana
Marais in Kew Bull. 39:
812.
1984. — Mauritius.
B10.
Polyscias neraudiana (Drake) R. Vig. in Bull. Soc. Bot. France 52: 301.
1905.
— Mauritius.
Bll.
Polysciaspaniculata
(DC.) Baker, Fl. Mauritius Seych.: 127. 1877, — Mauri-
tius.
B12.
Polyscias repanda (DC.) Baker, Fl. Mauritius Seych.: 128.
1877.
— Reunion.
B13.
Polyscias rivalsii Bernardi in Candollea 29: 158. 1974. — Reunion.
B14.
Polyscias rodriguesiana (Marais) Lowry & G. M. Plunkett, comb. nov.
=
Gas-
tonia rodriguesiana Marais in Kew Bull. 39: 809. 1984.
—
Rodrigues Island.
B15.
Polyscias
sessiliflora
Marais in Kew
Bull.
39: 814. 1984. — La Reunion.
C.
Polyscias subg. Maralia (Thouars) Lowry & G. M. Plunkett,
comb,
et stat.
nov.
=
Maralia Thouars, Gen. Nov. Madagasc: 13. 1806. — Type: M.
mada-
gascariensis DC. [= Polyscias maralia
(Roem.
& Schult.) Bernardi, non
P.
ma-
dagascariensis (Seem.) Harms].
= Oligoscias
Seem,
in J. Bot. 3: 179.
1865.
—Type:
O.
madagascariensis Seem.
[=
Polyscias madagascariensis (Seem.) Harms].
Polyscias subg. Maralia is circumscribed here to include all members of
the
Malagasy
subclade
within the Indian Ocean Basin
clade
of Polyscias sensu
lato
(Plunkett &
Lowry 2010). Of
the
35 currently recognized species in this group, all except three are
endemic to Madagascar: Polyscias felicis is restricted to the Comoro Islands,
P.
stuhl-
mannii
is found only in northeastern Tanzania and adjacent Kenya, and
P.
duplicata is
widespread in Madagascar and also occurs in the Comoros. The fact that each of these
species occupies a separate, derived position within the Malagasy subclade suggests
J

P.P. Lowry II & G.M. Plunkett,
Recircumscription
of Polyscias (Araliaceae) 65
that three independent dispersal events took place, one from Madagascar to mainland
Africa giving rise to
P.
stuhlmannii,
and two to the Comoros that resulted in the pres-
ence there of the other taxa.
One member of
this
subgenus, Polyscias duplicata, has always been included in the
polyphyletic genus Gastonia primarily because its pedicels lack an articulation below
the ovary. However, a second inarticulate species described by
Bernardi
(1971) as P.
carolorum
— which is not closely related to
P.
duplicata in the phylogeny of Plunkett
& Lowry
(2010)
— was curiously never associated with Gastonia despite exhibiting
its principal defining character. Moreover, Tennant (1960) described
P.
stuhlmannii
var. inarticulata Tennant, which provides another example of the loss of an articulation
within the
subgenus.
Plunkett & Lowry
(2010)
give additional information on the tax-
onomic
value of
this
and other characters traditionally used to define genera in Polys-
cias sensu
lato,
and Plunkett
ct
al.
(2004b) examine the value of morphological fea-
tures historically used to circumscribe infrafamilial taxa within Araliaceae.
Polyscias
subg.
Maralia includes four montane species described several decades
ago in Cuphocarpus, a long-recognized segregate genus defined by having flowers
with a unicarpellate
gynoecium.
The fifth taxon traditionally placed in this genus, C.
aculeatus
(the
generi-typc),
represents a distinct lineage (see below) in which a reduc-
tion to a single carpel has taken place, a parallelism that has likewise evolved in the
taxa historically included in Arthrophyllum.
In
his studies of Polyscias from Madagascar and the Comoros, Bernardi
(1971,
1980)
recognized a total of 28 species that we include here in subg. Maralia, one of
which
(P.
tennantii)
is placed in synonymy below. Extensive exploration in Madagas-
car over the last several decades has more than doubled the number of available collec-
tions,
yielding numerous previously
undescribed
species (for example see Callmander
ct al. 2009). Our ongoing studies based on both historical material and these more
recent gatherings indicate that at least 80 taxa remain to be described in Polyscias
subg. Maralia, which will bring the total to at least
115
species, making it the largest
group within the genus and also the most morphologically diverse.
Taxa
CI.
C2.
C3.
C4.
C5.
C6.
C7.
C8.
C9.
included:
Polyscias amplifolia (Baker) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3(8):
44,
1894.
— Madagascar.
Polyscias anacardium Bernardi in Candollea 26: 50.
1971.
— Madagascar.
Polyscias
andrearum
Bernardi in Candollea 26: 50.
1971.
— Madagascar.
Polyscias ariadnes Bernardi in Candollea 26: 29.
1971.
— Madagascar.
Polyscias
aubrevillei
(Bernardi) Bernardi in Candollea 26: 26.
1971.
— Mada-
gascar.
Polyscias baretiana Bernardi in Candollea 29: 147. 1974. — Madagascar.
Polyscias
briquetiana
(Bernardi) Lowry & G. M. Plunkett, comb. nov. = Cu-
phocarpus briquetianus Bernardi in Bull.
Soc.
Bot. Suisse 76: 354. 1966.
—
Madagascar.
Polyscias carolorum Bernardi in Candollea 26: 43.
1971.
— Madagascar.
Polyscias chapelieri (Drake) Harms ex R. Vig. in Bull. Soc. Bot. France 52:
303.
1905.
—Madagascar.

66 P.P. Lowry II &
G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae)
Polyscias tennantii
Bernardi in Candollea 26: 65.
1971,
syn. nov.
CIO.
Polyscias
cissiflora
(Baker) Harms in Engl. & Prantl, Nat.
Pflanzenfam.
3(8):
44.
1894.
— Madagascar.
Cll.
Polyscias
compacta
Lowry & G. M. Plunkett,
nom.
nov. =
Cuphocarpus
com-
mersonii Bernardi in Bull.
Soc.
Bot. Suisse 76: 356.
1966,
non Polyscias
com-
mersonii (Drake) R. Vig. [=
P.
paniculata (DC.) Baker]. — Madagascar.
CI2.
Polyscias confertifolia (Baker) Harms in Engl. & Prantl, Nat. Pflanzenfam.
3(8):
44.
1894.
— Madagascar.
C13.
Polyscias
cussonioides
(Drake) Bernardi in Candollea 26: 46.
1971.
— Mada-
gascar.
C14.
Polyscias duplicata (Thouars
ex Baill.)
Lowry & G. M. Plunkett, comb. nov.
= Gastonia duplicata Thouars ex
Baill.,
Adansonia 12:
166.
1878.
— Madaga-
scar, Comoro Islands.
C15.
Polyscias felicis Bernardi in
Candollea
26: 40.
1971.
— Comoro Islands.
C16.
Polyscias fraxinifolia (Baker) R. Vig. in Bull. Soc. Bot. France 52: 303.
1905.
—
Madagascar.
C17. Polyscias
gruschvitzkii
Bernardi in Candollea 26: 53.
1971.
— Madagascar.
C18.
Polyscias heineana Bernardi in Bull. Soc. Bot. Suisse 76: 364.
1966.
— Mada-
gascar.
C19.
Polyscias
humbertiana
(Bernardi) Lowry & G. M. Plunkett, comb. nov.
=
Cu-
phocarpus humbertianus Bernardi in Bull. Soc. Bot. Suisse 76: 358. 1966. —
Madagascar.
C20.
Polyscias
lancifolia
(Drake) Harms
ex
R. Vig., Bull. Soc. Bot. France 52: 304.
1905.
— Madagascar.
C21.
Polyscias lantzii (Drake) Harms ex R. Vig. in Bull. Soc. Bot. France 52: 304.
1905.
—Madagascar.
C22.
Polyscias leandriana (Bernardi) Lowry & G. M. Plunkett, comb. nov. = Cu-
phocarpus leandrianus Bernardi in Bull. Soc. Bot. Suisse 76: 359. 1966. —
Madagascar.
C23.
Polyscias
madagascariensis
(Seem.) Harms in Engl. & Prantl, Nat. Pflanzen-
fam. 3(8): 44.
1894.
— Madagascar.
C24.
Polyscias maralia
(Rocm.
& Schult.) Bernardi in Candollea 26: 34.
1971.
—
Madagascar.
C25.
Polyscias
mullibracteata
(Baker) Harms in Engl. & Prantl, Nat. Pflanzenfam.
3(8):
44. 1894. —Madagascar.
C26.
Polyscias muraltiana Bernardi in Bull. Soc. Bot. Suisse 76: 366. 1966. —
Madagascar.
C27. Polyscias myrsine Bernard in Candollea 26: 54.
1971.
— Madagascar.
C28.
Polyscias nossibensis (Drake) Harms in Bot. Jahrb. Syst. 26: 247.
1899.
-
Madagascar.
C29.
Polyscias ornifolia (Baker) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3(8):
44.
1894.
— Madagascar.
C30.
Polyscias
rainaliorum
Bernardi in Bull. Soc. Bot. Suisse 76: 367.
1966.
Madagascar.

P.P.
Lowry
II &
G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae) 67
C31.
Polyscias
stuhlmannii
Harms in Bot. Jahrb. Syst. 26: 244. 1899. — Kenya,
Tanzania.
C32.
Polyscias tafondroensis (Drake) Harms ex R. Vig. in Bull.
Soc.
Bot. France 52:
303.
1905.
—Madagascar.
C33.
Polyscias terminalia Bernardi in Candollea 26: 57.
1971.
— Madagascar.
C34.
Polyscias tripinnata Harms in
Engl.
& Prantl, Nat.
Pflanzenfam.
3(8): 44.
1894.
— Madagascar.
C35.
Polyscias
zanthoxyloides
(Baker) Harms in Engl. & Prantl, Nat. Pflanzenfam.
3(8):
44.
1894.
— Madagascar.
D.
Polyscias subg. Arthrophyllum
(Blume)
Lowry & G. M. Plunkett,
comb,
et
stat. nov. = Arthrophyllum Blume in Bijdr.: 878. 1826. — Type: A. diversifo-
lium
Blume
[=
Polyscias diversifolia (Blume) Lowry & G. M. Plunkett].
Mormoraphis Jack ex Wall,
Numer.
List [Wallich] no.
4931. 1831.
— Type:
M. sumatrana Jack ex Wall. [= Polyscias diversifolia (Blume) Lowry & G. M.
Plunkett].
Kissodendron
Seem,
in J. Bot. 3: 201.
1865 =
Polyscias sect. Kissodendron
(Seem.) Philipson in
Blumea
24:
170.
1978.
— Type: K.
australianum
(F. Mu-
ell.)
Seem.
[=
Polyscias australiana (F.
Muell.)
Philipson].
Eremopanax
Baill.
in Adansonia 12: 158. 1878. — Type: E.
otopyrenum
Baill.
[=
Polyscias otopyrena (Baill.) Lowry & G. M. Plunkett].
Nesodoxa Calest. in Webbia
1:
100.
1905.
—
Type:
N. vieillardii (Baill.) Fedde
[=
Polyscias vieillardii (Baill.) Lowry & G. M. Plunkett subsp. vieillardii].
Irvingia F. Muell. in
Fragm.
5:17. 1865,
nom.
illeg.
Shirleyopanax
Domin
in Biblioth. Bot. 89: 484. 1928, pro syn.
Polyscias subg. Arthrophyllum is currently the largest group within the genus, com-
prising a total of
37
species (one of which is raised here from the rank of variety) and
one subspecies. The subgenus ranges from the Andaman and Nicobar Islands and
southeast Asia (Peninsular Malaysia, Thailand and Laos) across Malesia (including the
Philippines, New Guinea and the Bismarck Archipelago) and south through northeast-
ern Australia to New Caledonia. Polyscias subg. Arthrophyllum, as circumscribed here,
corresponds to the Austro-Malesian
clade
of Polyscias sensu
lato
(Plunkett & Lowry
2010)
and incorporates taxa traditionally placed in Arthrophyllum (including the New
Caledonian segregate Eremopanax Baill.) and Kissodendron, along with two species
described in Pentapanax that were transferred to Polyscias by Philipson
(1977a).
Many members of
this
group are characterized by the presence of rusty brown stel-
late or farinose indument, especially on vegetative shoots, young foliage and develop-
ing inflorescences. A majority of
the
taxa (i.e. all those previously assigned to Arthro-
phyllum) have flowers with a unicarpellate gynoecium, a feature that clearly appears to
be derived within the Austro-Malesian clade and is intimately correlated with a distinc-
tive inflorescence structure in which two to four foliaceous, often compound bracts are
borne opposite one another or in whorls, contrasting with the spiral phyllotaxy of the
leaves.
While species limits are reasonably clear among the taxa historically assigned to

68 P.P.
Lowry
II & G.M. Plunkett,
Ftecircumscription
of Polyscias (Araliaceae)
Eremopanax
and
Kissodendron,
the same can not be said for the Malesian taxa that
have unicarpellate gynoecia and have traditionally been included in
Arthrophyllum.
As
mentioned in our companion paper (Plunkett & Lowry 2010), the paucity of herbarium
material of many species, some of which have particularly large, complex
inflores-
cences, prevented Philipson
(1977b,
1979)
from completing a revision of
the
Malesian
taxa to his full satisfaction. A limited amount of material has become available over the
last
few
decades (e.g.,
Lim
1986),
but further field work will still be required before a
comprehensive taxonomic treatment can be undertaken.
Taxa included:
Dl.
Polyscias aherniana (Philipson) Lowry & G. M. Plunkett, comb. nov. = Ar-
throphyllum ahernianum
Merr.
in Philipp. J. Sci., C
1
(Suppl.):
109.
1906
[= Ar-
throphyllum
bomeense
Baker in Bull. Misc. in
Inform.
Kew
1896: 23. 1896,
non Polyscias
borneense
Philipson],
— N. Borneo, Philippines, N. Maluku.
D2.
Polyscias
alternifolia
(Maingay ex
Ridl.)
Lowry & G. M. Plunkett, comb. nov.
= Arthrophyllum alternifolium Maingay
ex
Ridl. in
Fl.
Mai.
Pen.
1:
886.
1922.
— Peninsular Malaysia.
D3.
Polyscias
ungustifolia
(Ridl.) Lowry & G. M. Plunkett, comb. nov.
=
Arthro-
phyllum
angustifolium
Ridl.
in
J. Fed.
Mai.
St. Mus. 10: 136.
1920.
—
Peninsu-
lar Malaysia.
D4.
Polyscias ashtonii (Philipson) Lowry & G. M. Plunkett, comb. nov. = Arthro-
phyllum ashtonii Philipson in Gard. Bull. Sing. 30: 303.
1977.
— Borneo.
D5.
Polyscias australiana (F.
Muell.)
Philipson in
Blumea
24: 171. 1978.
—Aus-
tralia.
D6.
Polyscias bellendenkerensis (F. M. Bailey) Philipson in Austrobaileya 1: 24.
1977.
— Queensland.
D7.
Polyscias biforme (Philipson) Lowry & G. M. Plunkett, comb. nov. = Arthro-
phyllum biforme Philipson in Bull. Mus. Natl. Hist. Nat. (Paris), ser. 4, Sect. B,
Adansonia, 5: 237. 1983.
=
Arthrophyllum
diversifolium (Diiniker)
Philipson,
non
Blume
(1826)
nee
Polyscias diversifolia
(Blume)
Lowry & G. M. Plunkett.
— New Caledonia.
D8.
Polyscias
bipinnata
(Gibbs) Philipson in Blumea 24:
170.
1978.
— New Gui-
nea.
D9.
Polyscias cenabrei
(Merr.)
Lowry & G. M. Plunkett, comb. nov.
=
Arthrophyl-
lum
cenabrei
Merr. in Philip. J. Sci. 20: 417. 1922. — Philippines.
D10.
Polyscias collina (Philipson) Lowry & G. M. Plunkett, comb. nov. = Arthro-
phyllum collinum Philipson in Gard. Bull. Sing. 30: 305. 1977.
—Borneo.
Dll.
Polyscias revoluta (Philipson) Lowry & G. M. Plunkett,
nom.
nov.
=
Arthro-
phyllum
crassum
Philipson in Gard. Bull. Sing. 30: 305. 1977, non Polyscias
crassa (Hemsl.)
Lowry & G. M. Plunkett. — Borneo.
D12.
Polyscias
disperma
(F.
Muell.)
Lowry & G. M. Plunkett,
comb,
et stat. nov.
= Kissodendron australianum (F. Muell.) Seem. var. dispermum F. Muell. in
Descr. Notes Papuan
PI.
5: 88. 1877.
[=
Polyscias australiana (F. Muell.)
Philipson var. disperma (F. Muell.) Philipson]. — New Guinea. — Recent ex-
amination of material of this taxon, heretofore consistently treated as a variety

P.P. Lowry II & G.M. Plunkett,
Recircumscription
of Polyscias (Araliaceae) 69
of
P.
australiana, shows that it
represents
a morphologically and geographi-
cally well delimited entity that warrants recognition at the
species
level.
D13.
Polyscias
diversifolia (Blume)
Lowry & G. M. Plunkett, comb. nov. = Arthro-
phyllum
diversifolium Blume in Bijdr., 879.
1826.
— Java
1)14.
Polyscias elliptica (Blume) Lowry & G. M. Plunkett. comb. nov.
=
Arthrophyl-
lum
ellipticum
Blume in Bijdr., 879. 1826
[= Arthrophyllum javanicum
Blume
in Bijdr., 879.
1826,
non Polyscias
javanica
Koord.
&
Valeton].
— Java.
D15.
Polyscias engganoense (Philipson) Lowry & G. M. Plunkett, comb. nov.
= Arthrophyllum
engganoense
Philipson in Gard. Bull. Sing. 30: 305. 1977.
— Sumatra.
1)16.
Polyscias havilandii
(Ridl.)
Lowry & G. M. Plunkett. comb. nov. = Arthro-
phyllum
havilandii
Ridl. in
Kew
Bull. 1933: 494. 1933. — Borneo.
D17. Polyscias jackiana (G. Don) Lowry & G. M. Plunkett, comb. nov.
=
Hedera
jackiana G. Don in
Gen.
Hist. 3: 394. 1834 [= Arthrophyllum
jackianum
(G
Don) Frodin]. -Andaman &
Nicobar
Isl.
to western and central
Malesia.
D18.
Polyscias kjellbergii (Philipson) Lowry & G. M. Plunkett, comb. nov.
=
Arthrophyllum kjellbergii Philipson in Gard. Bull. Sing. 30: 309. 1977. -
Sulawesi.
1)19.
Polyscias lucens (Craib) Lowry & G. M. Plunkett, comb. nov. = Arthrophyl-
lum
lucens
Craib in Bull. Misc. Inform. Kew
1930:
423. 1930.
—Thailand.
1)20.
Polyscias mackeei Lowry & G. M.
Plunkett.
nom.
nov. = Eremopanax grandi-
folia Guillaumin
in Bull. Mus. Hist. Nat. (Paris) 33: 272. 1927
[=
Arthrophyl-
lum
grandifolium
(Guillaumin) Philipson], non Polyscias grandifolia Volkens.
— New Caledonia.
1)21.
Polyscias macranthum (Philipson) Lowry & G. M. Plunkett, comb. nov.
=
Arthrophyllum macranthum Philipson in Bull. Br. Mus. Nat. Hist., Bot.
1:18.
1951.
— New Guinea.
1)22.
Polyscias macrocarpa (Philipson & Bui) Lowry & G. M. Plunkett. comb. nov.
= Arthrophyllum macrocarpum Philipson & Bui in Adansonia, ser. 2, 17: 327.
1978.—
Laos.
D23.
Polyscias
meliifolia
(Craib) Lowry & G. M. Plunkett. comb. nov. = Arthro-
phyllum
meliifolium
Craib in Bull. Misc. Inform. Kew
1930:
424. 1930. -
Thailand.
D24.
Polyscias montana (Ridl.) Lowry & G. M. Plunkett, comb. nov. = Arthrophyl-
lum
montanum
Ridl. in J. Fed.
Mai.
St. Mus. 4: 24.
1909.
- Malaysia.
D25.
Polyscias otopyrena
(Baill.)
Lowry & G. M. Plunkett, comb. nov. = Eremo-
panax otopyrena Baill. in Adansonia 12: 158. 1878.
[=
Arthrophyllum oto-
pyrenum (Baill.) Philipson].
—
New Caledonia.
Eremopanax angustata Baill.
in
Adansonia 12: 159.
1878,
emend.
Baum.-Bod.,
Ber. Schweiz Bot. Ges. 64: 133. 1954
[=
Arthrophyllum
angustatum
(Baill.)
Philipson],
syn. nov.
Eremopanax angustata Baill. f. angusticarpa Baum.-Bod. in Ber. Schweiz Bot.
Ges.
64: 134.
1954.
syn. nov.
Eremopanax angustata Baill. f elliptica Baum.-Bod. in Ber. Schweiz Bot. Ges.
64:
133.
1954,
syn. nov.

70 P.P. Lowry II &
G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae)
= Eremopanax
angustata
Baill.
f. intermedia Baum.-Bod. in Ber. Schweiz Bot.
Ges.
64: 134. 1954, syn. nov.
Eremopanax angustata Baill. f. oblonga Baum.-Bod. in Ber. Schweiz Bot. Ges.
64:
133. 1954, syn. nov.
= Eremopanax canalensis E. G. Baker in A. B. Rendle et
al.
in J. Linn.
Soc.,
Bot.
45:
323. 1921.
[=
Eremopanax angustata Baill. f. canalensis (E. G. Baker)
Baum.-Bod.],
syn. nov.
Eremopanax schlechteri Harms in Bot. Jahrb. Syst. 39: 217. 1906. [= Arthro-
phyllum schlechteri (Harms) Philipson], syn. nov.
Eremopanax schlechteri Harms f. gracilis Baum.-Bod. in Ber. Schweiz Bot.
Ges.
64: 133. 1954, syn. nov.
D26.
Polysciaspacifica
(Philipson) Lowry & G. M. Plunkett, comb. nov. = Arthro-
phyllum
pacificum
Philipson in Gard. Bull. Sing. 30: 306.
1977.
— Maluku,
Bismarck Archipelago.
D27. Polyscias papyracea (Philipson) Lowry & G. M. Plunkett, comb. nov.
=
Ar-
throphyllum
papyraceum Philipson in Gard. Bull. Sing. 30: 308. 1977. —
Sumatra.
D28.
Polysciasprolifera
(Philipson) Lowry & G. M. Plunkett, comb. nov.
=Arthro-
phyllumproliferum
Philipson in Gard, Bull. Sing. 30: 302.
1977.
— Papua New
Guinea.
D29.
Polyscias pulgarense (Elmer) Lowry & G. M. Plunkett, comb. nov.
=
Arthro-
phyllum pulgarense Elmer in
Leafl.
Philip. Bot. 7:
2551.
1915.
— Philippines.
D30.
Polyscias royenii Philipson in
Blumea
24:
170.
1978.
—
W.
New Guinea.
D31.
Polyscias
mbiginosa (Ridl.)
Lowry & G. M. Plunkett, comb. nov.
=
Arthro-
phyllum
rubiginosum
Ridl. in
Kew
Bull.
1946:
40.
1946.
— Borneo.
D32.
Polyscias rufosepala (Ridl.) Lowry & G. M. Plunkett, comb. nov.
=
Arthro-
phyllum rufosepalum
Rild.
in Kew Bull. 1946: 40.
1946.
— Borneo.
D33.
Polyscias
schultzei
Harms in Bot. Jahrb. Syst. 56: 410. 1921. —
Makulu
to
New Guinea.
D34.
Polyscias stonei (A. L.
Lim)
Lowry & G. M. Plunkett, comb. nov. = Arthro-
phyllum stonei A. L. Lim in Malaysian Forester 43: 263.
1980.
— Peninsular
Malaysia.
D35.
Polyscias
thailandica
Lowry & G. M. Plunkett,
nom.
nov. = Arthrophyllum
ferrugineum Craib in Bull. Misc. Inform. Kew 1930: 423. 1930, non Polyscias
ferruginea (Hiern) Harms — Thailand.
D36.
Polyscias vieillardii (Baill.) Lowry & G. M. Plunkett, comb. nov. = Eremopa-
nax vieillardii Baill. in Adansonia 12: 161. 1878
[=
Arthrophyllum vieillardii
(Baill.) Philipson
=
Nesodoxa vieillardii (Baill.) Fedde].
a. subsp. vieillardii — New Caledonia.
b.
subsp. balansae (Baill.) Lowry & G. M. Plunkett,
comb,
et stat. nov. = Eremopa-
nax balansae Baill. in Adansonia 12: 160. 1878.
{^Arthrophyllum
balansae
(Baum.-Bod.) Philipson].
—New
Caledonia.
Eremopanax hederoides Baum.-Bod. in Ber. Schweiz Bot. Ges. 64: 131. 1954.
[=
Arthrophyllum hederoides (Baum.-Bod.) Philipson], syn. nov.

P.P. Lowry II & G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae) 71
D37.
D38.
Eremopanax daenikeri Baum.-Bod. in Ber. Schweiz Bot. Ges. 64: 132.
1954.
[=
Arthrophyllum
daenikeri (Baum.-Bod.) Philipson], syn. nov.
Eremopanax glaberrima Baum.-Bod. in Ber. Schweiz Bot. Ges. 64:
131.
1954.
[= Arthrophyllum glaberrimum (Baum.-Bod.) Philipson], syn. nov.
Polyscias
willmottii
(F.
Muell.)
Philipson in Austrobaileya 1: 24. 1977. -
Queensland.
Polyscias
zippeliana
(Miq.) Valeton in Bull. Dep.
Agric.
Indes
Neerl.
10: 42.
1907.
—New
Guinea, N Queensland.
E. Polyscias
subg.
Cuphocarpus
(Decne. & Planch.) Lowry & G. M. Plunkett,
comb,
et stat. nov. = Cuphocarpus Decne. &
Planch,
in Rev. Hort.
IV,
3: 109.
1854.
— Type: C. aculeata Decne. & Planch. [= Polyscias aculeata (Decne. &
Planch.) Harms].
Polyscias subg. Cuphocarpus, which corresponds to the Cuphocarpus subclade of the
Indian Ocean Basin
clade
in our recent
phylogeny
(Plunkett & Lowry
2010),
includes
a single species,
P.
aculeata, endemic to coastal forests in eastern Madagascar. From
the
outset this taxon was recognized as a distinct genus because, unlike nearly all other
Araliaceae known at the time, its flowers have a unicarpellate
gynoecium,
a feature
also found in the species assigned to the Malesian genus Arthrophyllum (treated here
as a subgenus within Polyscias). Over the last century and a
half,
nearly all authors
have recognized Cuphocarpus as a distinct genus, the only major exception being
Harms
(1894-97),
who included it in his broadly defined Polyscias.
Four additional species were described in Cuphocarpus by Bernardi (1966), all
based on
20"1
century material collected in humid forest between 1,000 and 1,800 m
elevation in northern Madagascar. Bernardi noted that they differ from the type species
in having flowers borne in
umbellules
rather than racemules and that they have very
different
eco-geographic
requirements, but the generic placement of these four taxa
had not previously been doubted because they consistently exhibit unicarpellate flow-
ers.
Our phylogenetic studies show, however, that the montane species are nested with-
in the clade recognized here as Polyscias subg. Maralia whereas the coastal taxon is
sister to the largely African subgenus Sciadopanax (Plunkett & Lowry
2010),
indicat-
ing that the evolution of
a
single carpel occurred on two separate occasions in Mada-
gascar, and that Cuphocarpus, as defined by Bernardi
(1966,
1980), is thus polyphyl-
etic.
Taxon included:
El.
Polyscias aculeata (Decne. & Planch.) Harms in Engl. & Prantl, Nat. Pflanzen-
fam.
3(8): 44.
1894.
— Madagascar.
Cuphocarpus inermis Baker in Linn. Soc, Bot.
21:
350.
1884.
F. Polyscias subg. Tetraplasandra (A. Gray) Lowry & G.
M.
Plunkett,
comb,
et
stat. nov.
=
Tetraplasandra A. Gray, U.S.
Expl.
Exped., Phan. 1: 727. 1854.
— Type:
T.
hawaiiensis
A. Gray
[=
Polyscias
hawaiiensis
(A. Gray) Lowry &
G. M. Plunkett].

72 P.P. Lowry II & G.M. Plunkett,
Recircumscription
of Polyscias (Araliaceae)
Reynoldsia A. Gray, U.S.
Expl.
Exped., Phan. 1: 723. 1854. — Lectotypc:
R.
sandwicensis
A. Gray
[=
Polyscias
sandwicensis
(A. Gray) Lowry & G.
M.
Plunkett],
designated by Hutchinson in Genera
Fl.
PI.
2: 58.
1968.
Dipanax
Seem,
in J. Bot. 6: 130.
1868.
—
Type: D. dipyrena (H. Mann)
A.
Hel-
ler [= Polyscias kavaiense (H. Mann) Lowry & G. M. Plunkett].
Triplasandra
Seem,
in J. Bot. 6:
139.
1868.
= Tetraplasandra
sect.
Neotetrapla-
sandra Sherff in Bot.
Leafl.
8: 12. 1953. — Lectotype (here designated):
T.
oahuensis
(A. Gray) Seem. [= Polyscias
oahuensis
(A. Gray) Lowry & G.
M.
Plunkett].
Heptapleurum
sect. Pterotropia H. Mann in
Proc.
Amer.
Acad. Arts 7: 168.
1867
=
Pterotropia (H. Mann) Hillebr. in Fl. Hawaiian
[si.:
149. 1888
=
Tetra-
plasandra sect. Pterotropia
(Hillebr.)
Sherff in Bot. Leafl. 7: 10. 1952. — Lec-
toype (here designated): //.
kavaiense
H. Mann
[=
Polyscias kavaiensis
(H.
Mann) Lowry & G. M. Plunkett].
=
Tetraplasandra
sect. Eutetraplasandra Harms in Engl. & Prantl, Nat. Pflanzen-
fam.
3(8): 29.
1894,
nomen.
superfl.
Tetraplasandra
sect. Nothotetraplasandra Harms in Engl. & Prantl, Nat.
Pflan-
zenfam.
3(8): 29. 1894. — Type: T.
meiandra
(Hillebr.) Harms [= Polyscias
oahuensis A. Gray) Lowry & G. M. Plunkett].
= Peekeliopanax Harms in
Notizbl.
Bot. Gart. Berlin-Dahlem 9: 478. 1926. —
Type: P.
spectahilis
Harms
[=
Polyscias spectabilis (Harms) Lowry & G. M.
Plunkett].
= Munroidendron Sherff
in
Bot. Leafl.
7:21.
1952.
—
Type:
M.
racemosum
(C. N.
Forbes) Sherff
[=
Polyscias racemosa (C. N. Forbes) Lowry & G. M. Plun-
kett].
Polyscias subg.
Tetraplasandra
is circumscribed to include all of the species tradition-
ally assigned to three central Pacific genera long regarded as closely related to one
another
(Munroidendron, Reynoldsia and Tetraplasandra), along with two Malesian
species assigned by Philipson
(1970,
1979,
1995)
to Gastonia and four others he placed
in Polyscias sect. Eupteron (Philipson 1978, 1979, 1995). This subgenus, which cor-
responds to the New Guinea-Polynesia
clade
in our phylogeny
of
Polyscias sensu
lato
(Plunkett & Lowry 2010), includes a total of 20 species that collectively range from
Sumatra across Malesia to the Solomon Islands, and through southern Polynesia (Sa-
moa, the Society Islands, and the Marquesas) to Hawaii.
Morphologically members
of
Polyscias subg. Tetraplasandra are rather heteroge-
neous,
as reflected by the fact that they have historically been placed in five genera.
Philipson
(1970)
provided a careful review of the features that he regarded as impor-
tant for distinguishing Gastonia (which he restricted to species ranging from Malesia
west to Madagascar and the Comoros) from those occurring in the central Pacific (of-
ten referred to as the Tetraplasandra group), although he unfortunately did not con-
sider any of the other Malesian species we include here in the subgenus (viz.
P.
bel-
ensis, P. borneensis, P. florosa and
P.
philipsonii).
Nonetheless, even without taking
account of the latter taxa, Philipson (1970) acknowledged that establishing clear ge-

P.P.
Lowry
II & G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae) 73
neric limits was
problematical
and could only be accomplished by using a combination
of
characters.
Costcllo
& Motley (2007) examined relationships among the
laxa
comprising the
Tetraplasandra group and showed that all of the species from Hawaii form a
clade
resulting from radiation following long-distance dispersal to the archipelago. Their
findings also indicate that the taxa traditionally placed in
Reynoldsia
belong to two
clades, one uniting the four species from southern Polynesia and another comprising
two Hawaiian taxa, R.
sandwicensis
and the species usually placed in the segregate
genus Munroidendron, whose long, pendant, racemose inflorescences give it a very
distinctive appearance. Another curious member of this group is
P.
gymnocarpa, whose
ovary appears to be fully superior, a derived condition in a family with otherwise ex-
clusively inferior or rarely partially inferior ovaries (Costello & Motley 2004).
Several Hawaiian members of Polyscias subg. Tetraplasandra exhibit polymery.
Polyscias
racemosa,
formerly treated as Munroidendron, has 12 to 20 petals, usually
10
or
15
stamens, and an ovary with as many as
15
carpels. Another species,
P.
sand-
wicensis, is typical of
the
taxa previously placed in Reynoldsia in having 8 to
12
petals
and stamens, and up to 24 carpels. Similarly, several species traditionally placed in
Tetraplasandra exhibit polymery, such as
P.
hawaiiensis,
with 5-8 petals and up to 4
times as many stamens, along with a 7- to
13-carpellate
ovary. Two other taxa previ-
ously included in Tetraplasandra, P. waialealae and P. waimeae, both endemic to
Kauai, have flowers with up to 40 and 65 stamens, respectively, and produce copious
nectar, suggesting possible bird pollination (Lowry
1990).
Philipson (1970, 1979, 1995) defined Gastonia serratifolia (Miq.) Philipson (treat-
ed here as Polyscias serratifolia) to encompass collections from throughout Malesia,
including material from Palawan in the southwestern Philippines (described as Tetra-
plasandra philipinensis Merr.) that may represent
a
distinct taxon
(J.
Wen pers.
comm.).
Philipson (1970, 1979) also noted that
P.
spectabilis
can reach 45 meters in height,
making it the tallest Araliaceae. Lowry (1987,
1990)
placed the taxa originally de-
scribed as Tetraplasandra bisattenuata and Triplisandra. lydgatei in synonymy under
the variable Hawaiian species recognized here as
P.
oahuensis, but recent field inves-
tigations indicate that both are distinct and deserve recognition at the species level
(Motley 2005; D.
Lorence
pers. comm.). Careful examination of collections histori-
cally treated as Reynoldsia shows that the two taxa from Samoa recognized by Frodin
& Govaerts (2003) as
R.
grayana and
R.
tauensis represent the same entity as R.
lanu-
toensis (treated here as
P.
lanutoensis), prompting us to include them in synonymy.
Similarly, Frodin & Govaerts (2003) accepted both R.
tahitiensis
and R. verrucosa
from the Society Islands, whereas we recognize only one species and therefore place
the former in synonymy.
Taxa included:
Fl.
Polyscias belensis Philipson in Bull. Brit. Mus. Nat.
Hist..
Bot. 1: 13. 1951.
— New Guinea.
F2.
Polyscias bisattenuata (Sherff) Lowry & G. M. Plunkett, comb. nov. = Tetra-
plasandra bisattenuata Sherff in Bot. Leafl. 6: 26. 1952. — Hawaii.

74 P.P. Lowry II & G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae)
F3.
Polyscias
borneensis
Philipson in J. Bot. 78: 118. 1940. — Borneo, Philip-
pines.
F4.
Polyscias florosa Philipson in
Blumea
24: 170.
1978.
— Philippines.
F5.
Polyscias flynnii (Lowry &
K.
R. Wood) Lowry & G. M. Plunkett, comb. nov.
= Tetraplasandra flynnii Lowry &
K.
R. Wood in Novon 10: 40. 2000. —
Hawaii.
F6.
Polysciasgymnocarpa
(Hillebr.) Lowry & G. M. Plunkett, comb. nov.
s
Ptero-
tropia gymnocarpa Hillebr. in
Fl.
Haw.
Isl.,
150.
1888
[=
Tetraplasandra
gym-
nocarpa (Hillebr.)
Sherffj.
— Hawaii.
F7.
Polyscias hawaiensis (A. Gray) Lowry & G. M. Plunkett, comb. nov. = Tetra-
plasandra
hawaiensis
A. Gray in U.S.
Expl.
Exped., Phan. 1: 728. 1854. —
Hawaii.
F8.
Polyscias kavaiensis (H. Mann) Lowry & G. M. Plunkett, comb. nov. = Hep-
tapleurum kavaiense H. Mann in
Proc.
Amer.
Acad. Arts 7: 168. 1867
[=
Tetra-
plasandra kavaiensis (H. Mann) Sherffj. — Hawaii.
F9.
Polyscias
lanutoensis
(Hochr.) Lowry & G. M. Plunkett, comb. nov.
=
Rey-
noldsia lanutoensis Hochr. in Candollea 2: 482. 1925. — Samoa.
= Reynoldsia grayana Christoph. in Bull. Bernice
P.
Bishop Mus. 128: 161. 1935,
syn. nov.
= Reynoldsia tauensis A. C. Smith & B. C. Stone in J. Arnold Arbor. 49: 465.
1968,
syn. nov.
F10.
Polyscias lydgatei (Hillebr.) Lowry & G. M. Plunkett, comb. nov. = Triplasan-
dra lydgatei Hillebr., Fl. Hawaiian
Isl.:
135. 1888.
[=
Tetraplasandra lydgatei
(Hillebr.) Harms] — Hawaii.
Fll.
Polyscias marchionensis (F. Brown) Lowry & G. M. Plunkett, comb. nov.
=
Reynoldsia marchionensis F. Brown in Bull. Bernice P. Bishop Mus. 130:
209.
1935.
—
Marquesas Isl.
F12.
Polyscias oahuensis (A. Gray) Lowry & G. M. Plunkett, comb. nov.
=
Gasto-
nia oahuensis A. Gray in U. S. Expl. Exped., Phan. 1: 726. 1854
[=
Tetrapla-
sandra oahuensis (A. Gray) Harms]. — Hawaii.
F13.
Polyscias
philipsonii Bernardi
in Ber. Schweiz. Bot. Ges. 76: 377. 1966. —
New Guinea.
F14.
Polysciaspleiosperma
(A. Gray) Lowry & G. M. Plunkett, comb. nov.
=
Reyn-
oldsia pleiosperma A. Gray in U. S. Expl. Exped., Phan. 1: 725. 1854. —
Samoa.
F15.
Polyscias racemosa (Forbes) Lowry & G. M. Plunkett, comb. nov. = Tetrapla-
sandra racemosa Forbes in
Occ.
Papers Bernice P. Bishop Mus. 6: 4. 1917.
[= Munroidendron racemosum (Forbes) Sherff]. — Hawaii.
F16.
Polyscias
sandwicensis
(A. Gray) Lowry & G. M. Plunkett, comb. nov.
=
Reyn-
oldsia sandwicensis A. Gray in U. S. Expl. Exped., Phan. 1: 724. 1854. —
Hawaii.
F17. Polyscias
serratifolia
(Miq.) Lowry & G. M. Plunkett, comb. nov. = Arthro-
phyllum serratifolium Miq. in Fl. Ned.
Ind.,
Eerste Bijv.: 341.
1861 [=
Gastonia
serratifolia (Miq.) Philipson]. — Malesia to Solomon Isl.
F18.
Polyscias spectabilis (Harms) Lowry & G. M. Plunkett, comb. nov.
=
Peeke-

P.P. Lowry II & G.M. Plunkett,
Recircumscription
of Polyscias (Araliaceae) 75
liopanax spectabilis Harms in Notizbl. Bot. Gart.
Berlin-Dahlem
9: 478.
1926
[=
Gastonia spectabilis (Harms) Philipson]. — New Guinea to Solomon
Isl.
F19.
Polyscias verrucosa (Seem.) Lowry & G. M. Plunkett, comb. nov.
=
Reynold-
sia verrucosa
Seem,
in J. Bot. 2: 245. 1864. — Society Isl.
Reynoldsia tahitiensis Nadeaud in
Enum.
PI.
Tahiti: 63.
1873,
syn. nov.
F20.
Polyscias
waialealae
(Rock) Lowry & G. M. Plunkett, comb. nov.
=
Tetrapla-
sandra
waialealae Rock in Coll. Hawaii
Publ.
Bull.
1:
10.
1911.
— Hawaii.
F21.
Polyscias
waimeae
(Wawra) Lowry & G. M. Plunkett, comb. nov. = Tetrapla-
sandra waimeae Wawra in Flora 56: 158. 1873. — Hawaii.
G. Polyscias subg. Eupteron
(Miq.)
Lowry & G. M. Plunkett,
comb,
et stat. nov.
=
Eupteron Miq. in Bonplandia 4:
139.
1856.=
Polyscias sect. Eupteron (Miq.)
Philipson in
Blumea
24: 170. 1978. — Type: E. nodosum
(Blume)
Miq.
[=
Po-
lyscias nodosa (Blume) Seem.].
As circumscribed here, Polyscias subg. Eupteron includes a single species,
P.
nodosa,
which is widely distributed across Malesia and extends into the Solomon
Islands,
often
colonizing open, disturbed habitats. It can easily be recognized by its large pinnately
compound leaves up to perhaps 3 meters in length and bearing 35 to more than
50 leaflets, as well as its large paniculate inflorescences with the flowers borne in ca-
pitula.
The generic name Eupteron, proposed by Miquel in
1856
based on
P.
nodosa and a
second species,
P.
acuminata from India and Sri Lanka, has been adopted by only a
few authors during the last 150 years, most notably Hutchinson (1967). Philipson
(1978,
1979, 1995) treated Eupteron as a section within Polyscias, expanding its cir-
cumscription to include several additional Malesian species with flowers arranged in
umbellules or capitula, placing primary emphasis to define the group on the presence
of fruits whose style arms are radiating or recurved. The results of our phylogenetic
analyses show, however, that
Philipson's
broadened concept of the Eupteron group is
not monophyletic;
P.
nodosa occupies an isolated position within Polyscias sensu
lato
whereas the other taxa belong to the
clade
we recognize here as Polyscias subg. Tetra-
plasandra (Plunkett & Lowry 2010). The relationships of
P.
acuminata, which we
have not yet been able to include in our molecular analyses, are not clear but they most
likely lie either with
P.
nodosa or with the members of subgenus Tetraplasandra.
Taxon included:
Gl.
Polyscias nodosa (Blume)
Seem,
in J. Bot. 3: 181. 1865. — Malesia, Philip-
pines,
Solomon Isl.
H. Polyscias subg. Sciadopanax (Seem.) Lowry & G. M. Plunkett,
comb,
et stat.
nov.
=
Sciadopanax
Seem,
in J. Bot. 3: 73. 1865. — Type: S. boivinii Seem.
[= Polyscias boivinii (Seem.) Bernardi].
Polyscias subg. Sciadopanax comprises
13
species from Africa (including the offshore
islands of
Sao
Tome and Principe), Madagascar and the Comoro Islands, and corre-

76 P.P.
Lowry
II & G.M. Plunkett,
Recircumscription
of Polyscias (Araliaceae)
sponds to the
P.fulva
subclade of the Indian Ocean
clade
in our phylogeny (Plunkett &
Lowry 2010). Originally circumscribed by
Seemann (1865)
as a genus with just a
single species from Madagascar, it was later expanded by Viguier
(1905)
to encompass
five of the taxa recognized here, two from Madagascar and three from Africa (plus six
others now placed in synonymy), using a circumscription identical to the one adopted
below. Until recently, authors mostly treated Sciadopanax in its original monotypic
sense, i.e., to comprise only S. boivinii (e.g., Harms 1894-97; Hutchinson 1967), but
more recently Tennant
(1968)
included it in Polyscias, and while Bernardi
(1966)
ini-
tially recognized Sciadopanax he later placed it in synonymy as well
(1971,
1980),
an
approach followed by
Bamps
(1974a, 1974b, 1989) and Beentje (1994).
Members of Polyscias subg. Sciadopanax are characterized by flowers with a bi-
carpellate gynoecium and styles that are united basally to form a conical disc or sty-
lopodium bearing two short styles or sessile stigmas. The leaves of most species have
stellate, often farinose indumentum, which is also frequently found on the inflores-
cence. The subgenus has both evergreen species growing in humid habitats in Africa,
including primary forest (such as
P.
albersiana and
P.
quintasii) and more disturbed or
secondary vegetation (e.g.,
P.fulva),
as well as deciduous taxa occurring in drier habi-
tats (including
P.
farinosa and
P.
kikuyuensis in Africa,
P.
mayottensis
in the Comores,
and all three species in Madagascar). Some members of the group are very large, espe-
cially
P.
quintasii, a canopy tree that can reach 30 m in height, whereas others are much
smaller, such as the recently discovered Central African insclberg specialist
P.
aequa-
toguineensis, a shrub or small tree up to only about 3 meters
tall
(Lejoly & Lisowski
1999).
Taxa included:
HI.
Polyscias
aequatoguineensis
Lejoly & Lisowki in Bull. Jard. Bot. Natl.
Belgi-
que 67:
112.
1999.
— Equatorial Guinea (Rio Muni), Gabon.
H2.
Polyscias albersiana Harms in Bot. Jahrb. Syst. 33:
182.
1902. — Tanzania.
H3.
Polyscias baehniana (Bernardi) Bernardi in Candollea 26:
21.
1971.
— Mada-
gascar.
H4.
Polyscias boivinii (Seem.)
Bemardi
in Candollea 26: 23. 1971. — Madagas-
car.
H5.
Polyscias
farinosa (Delile) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3(8):
45.
1894.— Ethiopia.
H6.
Polyscias
fioccosa
(Drake) Bernardi in Candollea 26: 22.
1971.
— Madagas-
car.
H7.
Polyscias
fulva
(Hiern) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3(8): 45.
1894.
— Tropical Africa.
H8.
Polyscias kikuyuensis Summerh. in Bull. Misc. Inform. Kew 1926: 242.
1926.
— Kenya.
H9.
Polyscias kivuensis Bamps in Bull. Jard. Bot.
Belg.
41: 249.
1971.
— E
Dem.
Republ. Congo.
H10.
Polyscias letestui C. Norman in J. Bot. 75: 167. 1937. — Gabon, Angola.
Hll.
Polyscias mayottensis Lowry,
O.
Pascal & Labat in Adansonia, ser. 3, 21: 69.
1999.
— Comoro
Isl.
(Mayotte, Moheli).

P.P.
Lowry
II & G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae) 77
H12.
Polyscias
quintasii Exell
in Cat.
Vase.
PI. S.
Tome:
195. 1944. —
Sao
Tome,
Principe.
H13.
Polyscias richardsiae
Bamps
in Bull. Jard. Bot.
Belg.
47: 260. 1977.
—Tanza-
nia.
I. Polyscias subg. Tieghemopanax (R. Vig.) Lowry & G. M. Plunkett.
comb,
et
stat. nov. = Tieghemopanax R. Vig. in Bull.
Soc.
Bot. France 52: 305. 1905.
— Lectotype:
T.
balansae
(Baill.)
R. Vig.
[=
Polyscias
balansae(Baill.)
Harms],
designated by Hutchinson in Genera
Fl.
PI.
2: 75. 1967.
Gelibia Hutch., Genera Fl.
PI.
2: 57.
1967 =
Polyscias sect.
Gelibia
(Hutch.)
Philipson in
Blumea
24:
169.
1978.
—Type:
G.
branderhorstii
(Harms) Hutch.
[=
P.
elegans
(C. Moore & F.
Mucll.)
Harms],
Montagueia
E. G. Baker in Rcndle et
al.
in
.1.
Linn.
Soc.,
Bot. 45: 291. 1921.
- Type: M.
haplostemon
E. G. Baker [=
P.
cissodendron
(C. Moore & F. Mu-
cll.) Harms].
Polyscias subg.
Tieghemopanax
corresponds to the Tieghemopanax
clade
identified in
several recent studies
(Eibl
et al.
2001;
Lowry et al. 2004; Plunkett et al. 2001, 2004b)
and confirmed in our companion paper
(Plu'
kctt
& Lowry
2010).
This group was first
recognized by
Viguier (1905)
as a new genus, which he circumscribed to include spe-
cies whose flowers have a
bicarpcllate gynoecium
and styles that are free or united
only at
the
base. He recognized 26 species, including 20 from New Caledonia (eight of
which are accepted below, either as species or subspecies, with the remainder placed
in synonymy), along with five occurring in Australia (three of which we do not include
in subgenus Tieghemopanax, i.e.,
P.
macgillivrayi,
P.
mollis and
P.
murrayi)
and one
from Madagascar
(P.
cussonioides,
which
wc
place here in subgenus
Maralia).
Viguier
(1905)
also doubtfully assigned three other species to his new
genus,
all of which we
place in Polyscias subg. Polyscias (viz.,
P.
multijuga,
P.
reincckei and
P.
samoensis).
Over the last century, Tieghemopanax was accepted by only a few authors (e.g..
Guil-
laumin
1938; Hutchinson 1967) while most others (e.g., Smith & Stone 1965; Stone
1965;
Bernardi
1971; Smith 1985; Lowry 1989; Frodin & Govaerts 2003) included it
within Polyscias.
Members of Polyscias subg. Tieghemopanax form a morphologically and geo-
graphically coherent group. In addition to sharing the diagnostic features used by
Viguier (1905), all of the taxa included here have leaves that lack a sheathing petiole
base (except in
P.
elegans, whose petiole is weakly clasping). There is also a tendency
toward having leaflets that are thick and often coriaceous (notable exceptions being
P.
hracteata
subsp. bracteata,
P.
cissodendron,
P.
crenata and
P.
schmidii). The group
is centered in New Caledonia, where ten of the
15
currently recognized species occur
(all except one endemic), with two species occurring in Fiji, one in NE Australia and
New Guinea, one endemic to Australia, and one restricted to Vanuatu. Recent field
work and herbarium studies have revealed
12
additional
species,
all endemic to New
Caledonia, which are being described in a forthcoming revision (Lowry & Plunkett in
prep.).

78 P.P.
Lowry
II &
G.M.
Plunkett, Recircumscription of Polyscias (Araliaceae)
Taxa included:
11.
Polyscias balansae
(Baill.)
Harms in Engl. & Prantl, Nat.
Pflanzenfam.
3(8):
44.
1894.
— New Caledonia.
12.
Polyscias bracteata (R. Vig.) Lowry in D. Frodin & R. Govaerts, World Check-
list Bibliogr. Araliaceae: 284. 2004. — New Caledonia.
a.
subsp.
bracteata. — New Caledonia.
b.
subsp.
subincisa
(R. Vig.) Lowry & G. M. Plunkett,
comb,
et stat. nov.
=
Tieghe-
mopanax
subincisus R. Vig. in Bull.
Soc.
Bot. France 52: 307. 1905. [= Polys-
cias subincisa (R. Vig.) Lowry]. — New Caledonia.
=
Tieghemopanax sessiliflorus
R. Vig. in Bull. Soc. Bot. France 52: 308. 1905,
syn. nov.
13.
Polyscias
cissodendron
(C. Moore & F.
Muell.)
Harms in Engl. & Prantl, Nat.
Pflanzenfam. 3(8): 45. 1894. — New Caledonia, Lord Howe Island, Vanuatu,
Santa Cruz
Isl.
14.
Polyscias crenata (Pancher & Sebert) Frodin in D. Frodin & R. Govaerts,
World Checklist Bibliogr. Araliaceae: 285. 2004. — New Caledonia.
15.
Polyscias
culminicola
A. C. Smith in Contrib. U.S. Natl. Herb. 37: 85.
1967.
-Fiji.
16.
Polyscias dioica (Vieill. ex Pancher) Harms in Engl. & Prantl, Nat. Pflanzen-
fam. 3(8): 45. 1894.
—New
Caledonia.
17.
Polyscias elegans (C. Moore & F. Muell.) Harms in Engl. & Prantl, Nat. Pflan-
zenfam. 3(8): 45.
1894.
— New Guinea, NE Australia.
18.
Polyscias joskei Gibbs in J. Linn. Soc, Bot. 39: 148.
1909.
— Fiji.
19.
Polyscias lecardii (R. Vig.) Lowry in D. Frodin & R. Govaerts, World Check-
list Bibliogr. Araliaceae: 291. 2004.
—New
Caledonia.
110.
Polyscias microbotrys (Baill.) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3(8):
45.
1894.
— New Caledonia.
111.
Polyscias pancheri (Baill.) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3(8):
45.
1894.— New Caledonia.
112.
Polyscias sambucifolia (Sieb. ex DC.) Harms in Engl. & Prantl, Nat. Pflanzen-
fam. 3(8): 45.
1894.
— E & SE Australia.
113.
Polyscias schmidii Lowry in Bull. Mus. Natl. Hist. Nat., Paris,
ser.
4, sect. B,
Adansonia
11:
140.
1989.—
Vanuatu.
114.
Polyscias scopoliae (Baill.) Lowry in D. Frodin & R. Govaerts, World Check-
list Bibliogr. Araliaceae: 298. 2004. — New Caledonia.
115.
Polyscias
weinmanniae
(Baill.) Harms in Engl. & Prantl, Nat. Pflanzenfam.
3(8):
45.
1894.
— New Caledonia.
J. Polyscias subg. Indokingia
(Hemsl.)
Lowry & G. M. Plunkett,
comb,
et stat.
nov. =
Indokingia
Hemsl. in Hooker's Icon.
PI.
29: t. 2805. 1909. — Type:
/. crassa Hemsl.
[=
Polyscias crassa (Hemsl.) Lowry & G. M. Plunkett].
Polyscias subg. Indokingia corresponds to a small group referred to in our companion
paper as the Seychelles
clade
(Plunkett & Lowry 2010), which comprises just three
species, all endemic to the Seychelles Islands. Hemsley
(1909)
described Indokingia to

P.P.
Lowry
II & G.M. Plunkett,
Recircumscription
of Polyscias (Araliaceae) 79
accommodate the species referred to here as P. crassa, which he curiously distin-
guished
from the unrelated (albeit also polymerous) palmate-leaved genus Tupidan-
thus, mentioning only that the new plant from the Seychelles had foliage that resem-
bled Gastonia cutispongia from Reunion. Friedmann (1986) transferred this taxon to
Gastonia
and described a new species in that genus, both of which possess inarticulate
pedicels, a feature shared with the third Seychelles species that had been included in
Gastonia by Harms nearly a century earlier (1894-97).
Many groups of plants show strong phytogeographic ties between the Seychelles
and Madagascar (Schatz
1996),
at least in part reflecting their geological history (the
two landmasses were connected until the Seychelles separated and began to move
northward with the Indian plate ca. 88 ma; Storey et
al.
1995). The results of our phy-
logenetic
studies (Plunkett & Lowry
2010)
suggest, however, that the taxa in Polyscias
subg. Indokingia are not closely related to species in either subgenus Maralia or Scia-
dopanax, the two groups present on Madagascar, but rather have evolved from a com-
mon ancestor that colonized the archipelago from the east, most likely somewhere in
Australasia.
Taxa included:
Jl.
Polyscias crassa
(Hemsl.)
Lowry & G. M. Plunkett, comb. nov. = Indokingia
crassa Hemsl. in Hooker's Icon. PL 29: t. 2805. 1906. [= Gastonia crassa
(Hemsl.) F. Friedmann]. — Seychelles.
J2.
Polyscias
lionnetii
(F.
Friedmann) Lowry & G. M. Plunkett, comb. nov. = Gas-
tonia lionnetii
F.
Friedmann in Bull. Mus. Natl. Hist. Nat., Paris, ser. 4, sect. B,
Adansonia4:
253. 1986. — Seychelles.
J3.
Polyscias
sechellamm
Baker in
Fl.
Mauritius Seych.: 128. 1877. [= Gastonia
sechellarum
(Baker) Harms]. — Seychelles.
K. Polyscias subg.
Palmervandenbroekia
(Gibbs) Lowry & G. M. Plunkett,
comb,
et stat. nov. = Palmervandenbroekia Gibbs in Fl. Arfak Mts.:
162.
1917.
= Polyscias sect. Palmervandenbroekia (Gibbs) Philipson in
Blumea
24:
171.
1978.
— Type: Palmervandenbroekia papuana Gibbs = Polyscias palmervan-
denbroekii
Bernardi,
won
Polyscias papuana (Miq.) Seem. (1965).
Polyscias subg. Palmervandenbroekia is a small, geographically restricted group of
five species occurring in mid- to high-elevation primary forest on New Guinea (the
Vogelkop Peninsula, Cyclops Mountains and the central range in West Papua and Pap-
ua, Indonesia, as well as the Western District of Papua New Guinea). Gibbs
(1917)
described the monotypic genus Palmervandenbroekia to accommodate a distinctive
new species
{P.
papuana) characterized by flowers with a bicarpellate gynoecium and
united styles forming a short, beak-like projection in fruit. Bernardi (1971) placed
Palmervandenbroekia in synonymy under Polyscias and published a new name for the
type species as the combination in the latter genus was already occupied. Philipson
(1978)
formally recognized the group as a section within Polyscias and described three
new species based on material examined in preparation for his revision of Araliaceae
for Flora Malesiana (Philipson
1978).

80 P.P.
Lowry
II & G.M. Plunkett,
Recircumscription
of Polyscias (Araliaceae)
Philipson (1995) suggested that a fifth species from the mountainous interior of
New
Guinea, described as Polyscias roemeriana Harms, might also belong to the
Palmervandenbroekia group, based on the description provided by Harms
(1921),
and
in particular on the presence of a compact inflorescence, flowers with a bicarpellate
gynoccium,
and long, united styles. Phillipson pointed out that if this placement were
correct,
P.
roemeriana would surely represent a distinct member of the Palmervanden-
broekia group by virtue of its bipinnate (vs. once-pinnate) leaves. He refrained, how-
ever, from placing
P.
roemeriana among the species he recognized, treating it instead
as an insufficiently known taxon because the holotype at Berlin had been destroyed
and he had not found any additional material. While examining the unidentified mate-
rial
of Polyscias
at the herbarium in Leiden, a collection with bipinnate leaves made in
1988 in central New Guinea was found that appeared to belong to Polyscias
subg.
Palmervandenbroekia. Careful comparison of this gathering with the protologue of
P.
roemeriana clearly shows that it belongs to this taxon. We have therefore listed
P.
ro-
emeriana below as the fifth member of the
subgenus,
and have designated this re-
cently identified collection as its neotype.
We have not yet been able to include any members
of
Polyscias subg. Palmervan-
denbroekia in our molecular
phylogenetic
studies, but there seems little doubt that the
group belongs to Polyscias sensu
lato.
Moreover, the distinctive morphology of these
species, combined with their restricted geographic range and similar ecological prefer-
ences,
strongly suggest that they form a
clade.
Until phylogenetic data
become
avail-
able and we are able to determine the position of the Palmervandenbroekia group
within Polyscias, we have opted to follow the approach used for the main clades elu-
cidated by our molecular studies (Plunkett & Lowry 2010) and recognize this distinc-
tive entity at the
subgeneric
rank.
Taxa included:
Kl.
Polysciaspalmervandenbroekii
Bernardi in
Candollca26:
16.
1971.
=
Palmer-
vandenbroekia papuana Gibbs (1917), non Polyscias
papuana (Miq.)
Seem.
(1865).
— W Papua.
K2.
Polyscias jacobsii Philipson in
Blumea
24:
171.
1978. — W Papua.
K3.
Polyscias roemeriana Harms in Bot. Jahrb. Syst.
56:411.
1921.
—
Type:
Indo-
nesia, Papua: Valentijn
Mts.,
trail between Koruppun and Angguruk,
'Menag-
mok'
forest
camp,
4"24'S,
139°37'E, 2600 m,
19.VII.
1988,
J.-M.Mangen 2131,
neotype L (0794866)!,
isoneotypc:
A (image seen). — New Guinea.
—As
the
holotype at Berlin [Southwest New Guinea, von Roemer (Exped.
Lorentz)
1239,
Bf]
was destroyed and no isotypes have been located, we have desig-
nated the only other known collection, gathered some
100
km to the cast of the
original locality, as the neotype.
K4.
Polyscias sleumeri Philipson in Blumea 24:
171.1978.
— W Papua.
K5.
Polyscias vogelkopensis Philipson in Blumea 24:
171.
1978.
— W Papua.
Taxa
incertae
sedis:
Polyscias acuminata (Wight)
Seem,
in J. Bot. 3: 181. 1865. — S. India, Sri Lanka.
- This species was originally included in the genus Eupteron along with
J

P.P. Lowry II &
G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae) 81
P.
nodosa, but its relationships can not easily be deduced from morphology and
may alternatively lie with the members of Polyscias subgenus Tetraplasandra.
The placement
of
P.
acuminata within our classification of
Polyscias
will there-
fore have to wait until we are able to obtain molecular sequence data.
Polyscias macdowallii (F.
Muell.) Domin
in Biblioth. Bot. 89: 483. 1928.
—
Queen-
sland. — This species, which was accepted by Frodin & Govaerts (2003), ap-
pears to be similar (if not identical) to
P.
mollis based on published descriptions.
However, because we have not yet been able to examine any authentic material,
we prefer to refrain from assigning
P.
macdowallii to a subgenus until morpho-
logical and/or molecular data are available that provide an indication of its
relationships.
Polyscias mollis (Benth.) Harms in Engl. & Prantl, Nat.
Pflanzenfam.
3(8): 45. 1894.
— Queensland. — The placement of
this
species remains poorly resolved and/
or weakly supported in the phylogenetic trees based on molecular data. In some
trees,
P.
mollis is part of a basal polytomy. In other trees, it is sister to a
clade
comprising P.
purpurea
(see below) and the taxa forming Polyscias subg.
Arthrophyllum,
or allied to a much larger clade uniting subgenera Maralia,
Grotenfendia, Sciadopanax, Cuphocarpus, Tetraplasandra,
lndokingia
and
Tieghemopanax. Given these disparities, it would be premature to assign
P.
mollis to any of
the
subgenera recognized here.
Polyscias
murrayi
(F. Muell.) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3(8): 45
1894.
— Queensland. — Frodin & Govaerts (2003) recognized this taxon, in-
dicating that it occurs in New South
Wales,
Queensland and Victoria. Philipson
(1979,
1995)
noted that P. murrayi is similar to P.
ledermannii
from New
Guinea, which he included in his expanded (and polyphyletic) Polyscias sect.
Eupteron.
Examination of specimens in several herbaria confirms that the
leaves, inflorescences and fruits of these two species indeed share a strong re-
semblance, supporting Philipson's idea that they are closely related. Unfortu-
nately, our phylogenetic trees provide little clue as to the placement of
P.
mur-
rayi (which falls in a polytomy at or near the base
of Polyscias
sensu
lato),
and
we have not be able to obtain samples of
P.
ledermannii for use in molecular
studies. Both species are thus left incertae sedis.
Polyscias ledermannii Harms in Bot. Jahrb. Syst. 56: 409. 1921. — New Guinea. —
As noted above, this species is morphologically very similar to
P.
murrayi and
will probably prove to be its closest relative.
Polyscias pentamera (Baker) Harms in Engl. & Prantl, Nat. Pflanzenfam. 3(8): 44.
1894.
— Madagascar. — This species appears to fall within the range of mor-
phological variation exhibited by members
of Polyscias
subg. Maralia, and we
would have no difficulty including it there were it not for the fact that it is
placed outside the corresponding clade in our molecular phylogeny (Plunkett &
Lowry
2010).
Instead,
P.
pentamera occupies an unresolved position at the base
of the large Indian Ocean Basin clade, which also includes four other clades
represented by subgenera Grotenfendia, Cuphocarpus, Sciadopanax and
Maralia, a finding that is rather difficult to reconcile with both morphology and
geography. We anticipate that additional data might confirm the inclusion of

82 P.P. Lowry II &
G.M.
Plunkett,
Recircumscription
of Polyscias (Araliaceae)
P.
pentamera
in subgenus
Maralia,
but until that time we prefer to refrain from
assigning it there.
Polyscias purpurea C. T. White in
Proc.
Roy.
Soc.
Queensland 47: 64. 1935
(publ.
1936).
—
Queensland. — The plaeement of this species tracks closely that of
P.
mollis in the phylogenetic trees based on molecular data (see above). Neither
taxon falls consistently in any single
clade,
and no clear indication of their rela-
tionships can be deduced from morphology or geography. We therefore prefer
to take a conservative approach, leaving both species unplaced, pending future
research.
Acknowledgments
We are
grateful
to Michael G. Pimenov and his colleagues for organizing the
\Th
International Apiales
Symposium, and to Claudia Erbar and Peter Leins for their invitation to contribute to this special issue
of
Plant
Diversity and Evolution. We also thank W. Wagner and W. Lack for their careful review and
constructive
critique of the
manuscript,
and several colleagues, especially P. B. Phillipson, G. E.
Schatz and J. Wen, for valuable comments and suggestions that helped us refine our thinking about
how best to address
the
classification of Polyscias sensu
lato.
The curators of the following herbaria
kindly provided access to their collections and/or provided digital images: A, G,
K,
L, LUX, MO,
NOU,
NY, P, US (acronyms as in Holmgren
ct
al.
1990). Financial support was provided by the U.S.
National Science Foundation (DEB 9981641, 0613728, 0614152 and 0743355), the National Geo-
graphic Society (5793-96), the John D. and Catherine T.
MacArfhur
Foundation, and the Andrew W.
Mellon Foundation.
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Addresses of the authors:
Dr. Porter P. Lowry II, Africa & Madagascar Department, Missouri Botanical Garden, P.O. Box
299,
St. Louis, Missouri
63166-0299
USA; and Museum National d'Histoire Naturelle, Departe-
ment Systematique
& Evolution (UMR 7205), 57 rue Cuvier, CP
39.
75231
Paris CEDEX 05,
France; e-mail: pete.lowry@mobot.org.
Prof.
Dr. Gregory M. Plunkett, Cullman Program for Molecular
Systematics,
The New York
Botanical Garden, Bronx, New York 10458-5126 USA; e-mail: gplunkett@nybg.org.