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Enlarging the Madagascan genus Fenerivia
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INTRODUCTION
The genus Polyalthia Blume (Annonaceae) was first estab-
lished by Blume (1830) following the transfer of species names
from Unona L. f. and Guatteria Ruiz & Pav. Although only five
species were initially recognised in the genus, over 300 species
names have subsequently been published within the genus (In-
ternational Plant Names Index, IPNI: http://www.ipni.org/), of
which approximately 150 names are currently accepted (Global
Index of Annonaceae Names: http://herbarium.botanik.univie.
ac.at/annonaceae/listTax.php). Polyalthia is therefore currently
one of the largest genera in Annonaceae.
Polyalthia has been described as a “default genus which is
defined only by the absence of conspicuous synapomorphies”
(Johnson & Murray, 1999), and this has resulted in it effec-
tively becoming a “dustbin” genus for many Palaeotropical
Annonaceae taxa of uncertain affinities. The inevitable mor-
phological heterogeneity resulting from this prompted several
authors to suggest that the genus is likely to be paraphyletic or
polyphyletic (Rogstad & Le Thomas, 1989; Van Setten & Koek-
Noorman, 1992; Doyle & Le Thomas, 1994, 1996; Doyle & al.,
2000). The polyphyletic status of the genus has since been
confirmed by Mols & al. (2004a, b), who used combined matK,
rbcL, and trnL-F sequence data to show that Polyalthia species
occur in at least five distinct clades. These clades furthermore
included (or were ambiguously related to) species belonging
to several smaller genera (Enicosanthum Becc., Haplostichan-
thus F. Muell., Marsypopetalum Scheff.), implying that these
genera also need to be incorporated in taxonomic reassessments
of Polyalthia. The unresolved phylogenetic status of the genus
Polyalthia is argu ably the most sig nif icant cu rrent impediment
to Annonaceae systematics research.
Hooker & Thomson (1872) recognised 25 species of Poly-
althia, which they classified in three sections, sect. Polyalthia
(as “Eupolyalthia”), sect. Monoon (Miq.) Hook. f. & Thomson,
and sect. Trivalvaria (Miq.) Hook. f. & Thomson. The latter two
sectional names were based on genera established by Miquel
(1865). Another section, sect. Afropolyalthia Engl. & Diels,
was subsequently added to the genus by Engler & Diels (1901)
for two African species. The taxonomic scope of Polyalthia
was later reduced following: (1) the reinstatement of Trival-
varia (Miq.) Miq. as a distinct genus (Sinclair, 1951; see also
subsequent revision by Van Heusden, 1996); (2) the separa-
tion of Polyalthia sect. Afropolyalthia as the distinct genus
Greenwayodendron Verdc. (Verdcourt, 1969); (3) the transfer
of several Polyalthia species characterized by pollen octads to
Disepalum Hook. f. (Sinclair, 1955) and Enicosanthellum Bân
(Bân, 1975; species subsequently transferred to Disepalum by
Johnson, 1989); (4) the removal of the Southeast Asian “Poly-
althia hypoleuca complex” (sensu Rogstad, 1989, 1990, 1994;
Rogstad & Le Thomas, 1989) as the new genus Maasia Mols
& al. (Mols & al., 2008); and (5) the recent transfer of several
Polyalthia species to Marsypopetalum (Xue & al., 2011).
Although Polyalthia is poorly represented in continental
Africa, the genus appears to be considerably more diverse in
Madagascar. Schatz & Le Thomas (1990) recognized 18 Poly-
althia species in Madagascar, which they classified into five
in formal groups (refer red to as groups A– E). Species in groups
B and C are characterized by sulcate pollen with a columellar
exine (Schatz & Le Thomas, 1990), typical of the “malmeoid”
Phylogenetic affinities of Polyalthia species (Annonaceae) with
columellar-sulcate pollen: Enlarging the Madagascan endemic
genus Fenerivia
Richard M.K. Saunders,* Yvonne C.F. Su* & Bine Xue
School of Biological Sciences, The University of Hong Kong, Pokfulam Road, Hong Kong, China
Author for correspondence: Richard M.K. Saunders, saunders@hkucc.hku.hk
* Both authors contributed equally to this article.
Abstract
The genus Polyalthia (Annonaceae) has been demonstrated to be polyphylet ic, with the Asian specie s with columellar-
sulcate pollen recently segregated to form a new genus, Maasia. The phylogenetic affinities of the other Polyalthia species
with colu mell ar-sulc at e pollen, which occur in Madagascar, are investiga te d using ma xi mu m parsimony, ma xi mu m likeli ho od
and Bayesian phylogenetic analyses of three chloroplast DNA markers (trnL-F, matK, rbcL). The species are shown to form
a well-supported clade which warrants taxonomic recognition as a distinct genus. The name Fenerivia is adopted, consider-
ably expanding the application of this formerly monotypic name, with nine new nomenclatural combinations validated. The
genus is hypothesised to have undergone a homeotic mutation in which the outermost whorl of stamens develop as inner petals
(presumed andropetals), and the calyx is reduced to a vestigial flange.
Keywords
Annonaceae; Fenerivia; homeosis; Maasia; Madagascar; Malagasy; molecular phylogenetics; Polyalthia
Supplementary material
The alignment is available in the Supplementary Data section of the online version of this article
(http://www.ingentaconnect.com/content/iapt/tax).
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Enlarging the Madagascan genus Fenerivia
clade (Doyle & Le Thomas, 1996). In contrast, species in
groups A, D, and E are characterized by inaperturate pollen
with a non-columellar exine (Schatz & Le Thomas, 1990),
typical of many members of the “miliusoid” clade (Doyle & Le
Thomas, 1996). It is noteworthy, however, that some of the mil-
iusoid taxa previously regarded as inaperturate, including Tri-
valvaria (Walker, 1971) and Marsypopetalum s.l. (Mols & al.,
2004a), are now known to be disulculate (Xue & al., 2011), and
that such pollen may be widespread within the mil iu soid clade
(Waha & Hesse, 1988; Waha & Morawetz, 1988; Chaowasku
& al., 2008). The contrasting pollen characteristics of the dif-
ferent Madagascan representatives of Polyalthia suggest that
they are unlikely to be congeneric and that those in groups B
and C may share a closer phylogenetic affinity with Maasia,
which comprises species formerly included in the Polyalthia
hypoleuca complex (Mols & al., 2008). The present research
clarifies the phylogenetic and taxonomic relationships between
Maasia and the Madagascan Polyalthia species with columel-
lar-sulcate pollen, and is based on extensive sampling of spe-
cies from group B (P. angustielliptica Schatz & Le Thomas,
P. chapelieri Baill., P. emarginata Diels, P. madagascarien-
sis Cavaco & Keraudren) and group C (P. capuronii Cavaco
& Keraudren, P. ghesquiereana Cavaco & Keruadren, P. het-
eropetala (Diels) Gesq., P. humbertii Cavaco & Keraudren,
P. oligosperma (Danguy) Diels, P. richardiana Baill.).
Polyalthia heteropetala, a member of group C, was origi-
nally described as the sole representative of the genus Fenerivia
Diels (Diels, 1925). This taxonomically isolated position was
proposed because of the aberrant floral morphology evident in
the only known flowering specimen: unlike the flowers of the
great majority of Annonaceae species, which have a perianth
of nine organs in three whorls (three sepals, three outer petals,
three inner petals), Polyalthia (Fenerivia) heteropetala has a
perianth of 12 organs. The homology of the latter organs has
been the source of some discussion (Diels, 1925; Ghesquière,
1939; Schatz & Le Thomas, 1990; Deroin, 2007), but was inter-
preted by Saunders (2010) to be the result of a disruption to the
homeotic control of organ identity during floral development.
Molecular phylogenetic data is used in the present research to
reassess putative homologies in the floral structure of P. het-
eropetala and related species with columellar-sulcate pollen.
MATERIALS AND METHODS
Taxon sampling. —
The data matrix comprised a total of
123 species of Annonaceae representing all major clades in
the family. The dataset was based on three chloroplast DNA
regions: trnL-F, matK, and rbcL. Novel sequences were gener-
ated for the additional species from Madagascar, including all
Madagascan species of Polyalthia belonging to groups B and
C (sensu Schatz & Le Thomas, 1990), together with additional
Maasia species. The samples, localities, and GenBank acces-
sion numbers are given in the Appendix.
DNA extraction, amplification, and sequencing. —
Total
genomic DNA was extracted from herbarium leaf samples us-
ing a modified cetyl trimethyl ammonium bromide (CTAB)
method (Doyle & Doyle, 1987; Erkens & al., 2008; Su & al.,
2008), with DNA purified using the Wizard PCR Preps DNA
Purification System (Promega, Madison, Wisconsin, U.S.A.),
and eluted with 50 µl preheated TE (Tris-EDTA) buffer.
The following primers were used for amplification: trnL-
F (trnL intron and trnL-trnF intergenic spacer)—primers
trnLF-12F/433R, trnLF-147F/508R, trnLF-413F/724R, and
trnLF-597F/960R (Su & al., 2008); matK gene—primers
matK-13F/515R , matK-424F/788R, and matK-449F/824R (Su
& al., 2008); and rbcL gene—primers rbcL-7F/429R, rbcL-
127F/734R , rbcL-656F/1100R, and rbcL-984F/1381R (Su & al.,
2008).
All PCR reactions were performed in a total volume of
50 µl using 1× PCR buffer, ca. 10 ng DNA, 3 mM MgCl
2
,
0.2 mM dNTP, 0.3 µM of each primer, 2 U Taq DNA poly-
merase (Promega), and 0.5 µg/µl BSA (Promega). The PCR
conditions consisted of 3 min at 94°C, followed by 35 cycles
of 45 s at 94°C denaturation; 30 s at 49°C annealing for trnL-F,
52°C–55°C annealing for matK and rbcL; and 2 min extension
at 72°C; and an additional final extension of 7 min at 74°C. Suc-
cessfully amplified products were purified with the QIAquick
PCR purification kit (Qiagen, Valencia, California, U.S.A.)
following the manufacturer’s instructions.
Sequencing reactions were carried out for both forward and
reverse strands using the same PCR primers, and the BigDye
Terminator v.3.1 cycle sequencing kit (Applied Biosystems,
Foster City, California, U.S.A.). Purified sequencing reactions
were run commercially on an Applied Biosystems 3730XL
DNA Analyzer.
Phylogenetic analyses. —
Sequences were edited and
assembled in SeqMan Pro using DNAStar Lasergene 8.0
(DNAStar, Madison, Wisconsin, U.S.A.), and aligned manually
using BioEdit v.7.0.9 (Hall, 1999) and Se-Al v.2.0a11 (Ram-
baut, 1996). The combined three-region dataset consisted of
3186 aligned characters (trnL-F: 1054 bp; matK: 789 bp; rbcL:
1343 bp) following the removal of 65 bp ambiguously aligned
positions in trnL-F.
Maximum parsimony (MP) analyses of the three com-
bined datasets were conducted using PAUP* v.4.0b10 (Swof-
ford, 2003). All characters were weighted equally and indels
were treated as missing data. The most pa rsimonious trees were
obtained with heuristic searches of 1000 replicates of random-
addition-sequence, tree bisection-reconnection (TBR) branch
swapping, and saving 10 trees from each random sequence
addition. Bootstrap support (BS) values for the nodes were
estimated using 1000 random-addition-sequence replicates
with TBR branch swapping, and 10 trees saved per replicate.
Maximum likelihood (ML) analyses were conducted for
the three-gene dataset using GARLI v.0.95 that implements
a full heuristic tree search (Zwickl, 2006). The best-fitting
nucleotide substitution model was determined by the Akaike
In formation Criterion (AIC) using the MrModelTest v.2.3 (Ny-
lander, 2004). The analyses were performed using a random
starting tree and the ML bootstrap (BS) values were estimated
from 200 bootstrap replicates in GARLI.
Partitioned Bayesian analyses were performed on the
three-gene combined dataset using MrBayes v.3.1.2 (Ronquist
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& Huelsenbeck, 2003). Separate models of molecular evolu-
tion for individual gene partitions were determined by Mr-
ModelTest v.2.3 (Nylander, 2004). Four Metropolis-coupled
Markov Chain Monte Carlo (MCMC) runs, each with five
heated chains and one cold chain, were run for 5 million
generations with trees sampled every 100 generations. The
final average standard deviation of the split frequencies was
< 0.01, indicating that the sampling of the posterior distribu-
tion was adequate. The convergence of the MCMC runs was
also compared and checked using the online program AWTY
(Nylander & al., 2008). The output AWTY plot displaying
the posterior probabilities of splits between runs was linear,
indicating that the runs had reached convergence. Effective
sampling size (ESS) and burn-in were estimated using Tracer
v.1.5 (Rambaut & Drummond, 2009); all parameters had ESS
values above 200 and the first 10% of the sampled trees was
discarded. The remaining trees were used to generate a 50%
major ity-rule consensus tree and calculate the posterior prob-
abilities (PP). A more distant outgroup, Anaxagorea silvatica,
was used to root the tree.
RES ULTS
Of the 3186 included characters, 376 bp (11.8%) were
variable but parsimony-uninformative and 438 (13.7%) were
parsimony-informative. Parsimony analysis of the combined
regions yielded 2310 most parsimonious trees of 1463 steps
(consistency index, CI = 0.645; retention index, RI = 0.784;
rescaled retention index, RC = 0.506). For the combined ML
analyses, GTR + I + G was selected as the best-fitting model.
Model parameter values were estimated as follows: base fre-
quencies: A = 0.2885, C = 0.1946, G = 0.2125, T = 0.3043; rate
matrix: A–C = 1.3785, A–G = 2.8460, A–T = 0.6622, C–G
= 0.8091, C–T = 3.4410, G–T = 1.0; proportion of invariable
sites (I) = 0.4708; and gamma shape parameter = 0.9649. The
likelihood score of the optimal ML tree, −InL, was 14,622.2148.
For the mixed-model Bayesian analyses, the best-fit model of
sequence evolution for each gene partition was selected by AIC,
as follows: GTR + G for trnL-F and matK, and GTR + I + G for
rbcL data.
Parsimony, maximum likelihood, and Bayesian analyses
of the combined cpDNA regions consistently recognized four
Polyalthia clades (clades I–IV in Figs 1–2). Clade IV includes
10 Polyalthia species (viz.: P. chapelieri, P. heteropetala,
P. madagascariensis, P. angustielliptica, P. oligosperma,
P. richardiana, P. humbertii, P. ghesquiereana, P. emargi-
nata, P. capuronii). This clade received very strong statisti-
cal support (MP BS = 100%, ML BS = 100%, PP = 1.00;
Fig. 1). Maasia is furthermore shown to form a well-supported
clade (MP BS = 97%, ML BS = 97%, PP = 1.00). Clade IV
shows a relatively rapid average rate of molecular evolution
(Fig. 2), with 0.1533 substitutions per site; this is approxi-
mately 6 times faster than that of clade I (0.024 substitutions
per site) and 2.5 times faster than clade II (0.060 substitutions
per site), but is similar to that of clade III (0.1383 substitutions
per site).
DISCUSSION
The results of the phylogenetic analyses presented here
(Figs 1–2) reveal that the “Polyalthia columellar-sulcate group”
(sensu Doyle & Le Thomas, 1994, 1995, 1996) is represented
by two distinct and well-supported clades: an Asian clade
with five Maasia species; and a Madagascan clade with all 10
Polyalthia species belonging to groups B and C (sensu Schatz
& Le Thomas, 1990). Significantly, the latter clade has never
been recovered in previous molecular phylogenetic analyses of
the family. The phylogenetic relationships of these two “col-
umellar-sulcate” clades are obscured, however, due to lack of
adequate resolution and support. Only one other Madagascan
Polyalthia species was included in the analysis: P. pendula
Capuron ex Schatz & Le Thomas, belonging to group E (sensu
Schatz & Le Thomas, 1990), is shown to be nested within Pol-
yalthia clade III (Fig. 1), together with P. cerasoides (from
Asia), P. stuhlmannii (from continental Africa), and P. korinti
(from Asia). Although Polyalthia species in groups A and D
and those occurring in East Africa were not included in the
present analyses, they are known to be related to species in
Polyalthia clade III based on palynological and macromor-
phological resemblances (Verdcourt, 1969; Vollesen, 1980a,
b; Schatz & Le Thomas, 1990).
It is evident that the newly identified Madagascan clade
of Polyalthia species (clade IV in Figs 1–2) is not congeneric
with Polyalthia s.str. (represented by clade II), and the species
therefore need to be reclassified. The species in Polyalthia
clade IV are furthermore not congeneric with either of the other
Polyalthia clades (I or III), nor any of the other genera which
include species formerly classified in Polyalthia.
One of the species in clade IV, P. heteropetala, was origi-
nally described by Diels (1925) as the type of the monotypic
genus Fenerivia. The name Fenerivia heteropetala was sub-
sequently transferred to Polyalthia (Ghesquière, 1939), and
although this approach was adopted in most later taxonomic
treatments (Cavaco & Keraudren, 1957, 1958; Schatz & Le
Thomas, 1990), others have maintained Fenerivia as a distinct
genus (e.g., Fries, 1959; Van Heusden, 1992; Keßler, 1993).
The recognition of Fenerivia as a distinct genus has recently
received support based on interpretation of floral anatomy
(Deroin, 2007). As the earliest generic name associated with
the species in clade IV, Fenerivia is shown here to be the cor-
rect name for all species in this clade. Nine new nomenclatu-
ral combinations are consequently validated in this paper (see
“Taxonomic treatment” section, below).
Diels (1925) originally segregated F. heteropetala from the
Madagascan Polyalthia species because of its extraordinary
floral structure, with what he interpreted as a highly reduced
calyx of three vestigial sepals, and a corolla consisting of three
broadly ovate outer petals and nine linear inner petals (Fig. 3).
Ghesquière (1939) questioned this interpretation, however,
suggesting that the “calyx” represents an extra-floral nectary,
that the “outer petals” represent the true calyx, and that the
nine “inner petals” actually represent both the inner and outer
whorls of petals (with duplication of one of the whorls). Schatz
& Le Thomas (1990) similarly doubted Diels’ interpretation of
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Enlarging the Madagascan genus Fenerivia
Polyalthia cf. glabra
Polyalthia lateriflora
Polyalthia sclerophylla
Polyalthia viridis
Polyalthia cf. longifolia
Polyalthia longifolia
Enicosanthum fuscum
Polyalthia congesta
Polyalthia rumphii
Polyalthia xanthopetala
Enicosanthum paradoxum
Enicosanthum membranifolium
Polyalthia coffeoides
Neo-uvaria acuminatissima
Neo-uvaria parallelivenia
Stelechocarpus burahol
Stelechocarpus cauliflorus
Sageraea lanceolata
Stenanona costaricensis
Stenanona panamensis
Desmopsis microcarpa
Desmopsis schippii
Sapranthus viridiflorus
Sapranthus microcarpus
Tridimeris hahniana
Meiogyne stenopetala
Fitzalania heteropetala
Meiogyne cylindrocarpa
Meiogyne virgata
Mitrephora keithii
Mitrephora alba
Mitrephora polypyrena
Popowia odoardoi
Popowia hirta
Popowia pisocarpa
Phaeanthus ebracteolatus
Phaeanthus splendens
Polyalthia borneensis
Polyalthia obliqua
Polyalthia cauliflora
Polyalthia flagellaris
Haplostichanthus longirostris
Polyalthia celebica
Polyalthia cinnamomea
Polyalthia sp.
Polyalthia subcordata I
Polyalthia subcordata II
Polyalthia longipes
Polyalthia debilis
Polyalthia suberosa
Polyalthia stenopetala
Pseuduvaria acerosa
Pseuduvaria froggattii
Pseuduvaria megalopus
Pseuduvaria siamensis
Pseuduvaria borneensis
Pseuduvaria pamattonis
Pseuduvaria galeata
Pseuduvaria setosa
Marsypopetalum pallidum
Marsypopetalum littorale
Miliusa campanulata
Miliusa longipes
Miliusa cuneata
Miliusa velutina
Miliusa horsfieldii
Miliusa lineata
Polyalthia pendula
Polyalthia stuhlmannii
Polyalthia cerasoides
Polyalthia korinti
Orophea polycarpa
Woodiellantha sp.
Orophea kerrii
Orophea celebica
Orophea enterocarpa
Alphonsea boniana
Alphonsea elliptica
Alphonsea kinabaluensis
Platymitra macrocarpa
Trivalvaria macrophylla
Monocarpia euneura
Polyalthia heteropetala
Polyalthia chapelieri
Polyalthia madagascariensis
Polyalthia angustielliptica
Polyalthia oligosperma
Polyalthia ghesquiereana
Polyalthia humbertii
Polyalthia richardiana
Polyalthia emarginata
Polyalthia capuronii
Klarobelia inundata
Pseudephedranthus fragrans
Ephedranthus sp.
Oxandra espintana
Pseudomalmea diclina
Ruizodendron ovale
Mosannona costaricensis
Maasia discolor
Maasia glauca
Maasia ovalifolia
Maasia multinervis
Maasia sumatrana
Cremastosperma brevipes
Cremastosperma cauliflorum
Pseudoxandra lucida
Onychopetalum periquino
Bocageopsis multiflora
Unonopsis stipitata
Malmea dielsiana
Piptostigma mortehani
Polyceratocarpus pellegrinii
Mwasumbia alba
Greenwayodendron oliveri
Annickia chlorantha
Annickia pilosa
Cananga odorata
Cyathocalyx martabanicus
Cleistopholis glauca
Tetrameranthus duckei
Meiocarpidium lepidotum
Anaxagorea silvatica
61/66/1.00
65/58/0.99
100/100/1.00
59/64/0.85
63/58/0.55
90/86/1.00
96/93/1.00
100/100/1.00
77/83/1.00
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58/–/1.00 96/94/1.00
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98/98/1.00
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83/86/1.00
64/74/0.93
100/100/1.00
71/70/1.00
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91/91/1.00
56/69/0.99
63/64/1.00
88/86/1.00
88/91/1.00
55/65/0.84
100/100/1.00
69/71/0.83
–/–/0.83
–/–/0.98
–/–/0.67
99/98/1.00
97/95/1.00
–/–/0.63
100/100/1.00
96/93/1.00
100/100/1.00
99/96/1.00
100/96/1.00
61/56/0.99
87/92/0.99
65/75/0.95
100/100/1.00
62/70/0.88
99/99/1.00
79/65/1.00
97/92/1.00
100/99/1.00
94/86/1.00
–/–/0.91
63/66/0.99
92/95/1.00
–/–/0.62
54/52/0.95
100/100/1.00
80/86/1.00
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77/68/1.00
88/94/1.00
92/89/1.00
95/96/1.00
99/98/1.00
97/97/1.00
100/99/1.00
94/92/1.00
63/70/0.95
100/100/1.00
96/98/1.00
88/90/1.00
100/100/1.00
69/67/0.91
100/100/1.00
86/81/1.00
77/93/1.00
100/100/1.00
–/100/–
100/100/1.00
97/98/1.00
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–/–/0.72
100/98/1.00
90/78/1.00
50/76/0.98
IV
III
II
I
Miliusoid cladeMalmeoid clade
Fig. 1.
Consensus tree resulting from
partitioned Bayesian analysis of rep-
resentatives of the basal grade and
short-branch clade of Annonaceae
based on three combined cpDNA
markers (trnL-F, matK, rbcL). Poly-
althia is shown to be polyphyletic
(clades I–IV, described in text), with
clade IV representing Fenerivia s.l.
MP bootstrap/ML bootstrap/Bayes-
ian posterior probability values are
indicated beside branches. Thick
lines represent branches with poste-
rior probability (PP) >0.95.
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Enlarging the Madagascan genus Fenerivia
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floral structure, but suggested that the calyx (sensu Diels) was
merely an artefact resulting from dehydration of the receptacle.
Deroin (2007) published a detailed anatomical study of
F. heteropetala and argued that Diels’ original interpretation
was correct. He demonstrated that the nine linear petals are of
two types: three bracteopetals, homologous with the inner pet-
als of other Annonaceae species; and six andropetals, derived
from stamens. This was interpreted by Saunders (2010) to be
the result of a substantial disruption of the homeotic control of
floral organ identity during development, with a centrifugal
shift so that the outermost whorl of six stamens develop as inner
petals, the inner petals develop as outer petals, the outer petals
develop to be sepal-like, and the sepals fail to develop sig-
nificantly. Only one f lowering specimen of F. heteropetala is
known (the holotype, H. Perrier de la Bâthie 4942, P) and there-
fore it is not known whether the homeotic mutation occurred
only in this individual (perhaps encouraging de la Bâthie to
collect the sample) or whether the mutation has become fixed
as a characteristic of the species as a whole.
The highly reduced calyx of three vestigial sepals (sensu
Diels) is apparent in all species in the Fenerivia clade, and is
visible in both living plants (Fig. 4A, E) and dried herbarium
specimens, and is noticeably persistent in fruiting material
(Fig. 4B, F). This feature is therefore a valuable character
for distinguishing Fenerivia from other Annonaceae genera
and is a clear synapomorphy for the genus. Deroin (2007)
demonstrated that the flange in F. heteropetala has a va scular
system that is consistent with its interpretation as a calyx,
hence confirming Diels’ view of the flange as a reduced ca-
lyx. Other species transferred here to Fenerivia superficially
have the standard number of petals and sepals for the family
(Fig. 4C, E; Cavaco & Keraudren, 1958) but also possess
the outermost “calyx” flange. Although the homology of the
flange in F. heteropetala and other Fenerivia species has yet
to be unequivocally demonstrated by examining the vascular
supply in the latter, it seems likely that the structures are
equivalent. If this interpretation is correct, the linear petals
of species such as F. angustielliptica (Fig. 4C–E) would pos-
sibly represent two distinct whorls, of mixed bracteopetal
and andropetal origin, consistent with the hypothesis of a
disruption to the homeotic control of organ identity during
floral development; unequivocal interpretation would require
detailed anatomical research to reveal the vascular supply of
the petals, however.
Fig. 2.
Phylogram resulting from par-
titioned Bayesian analysis of three
combined cpDNA markers (trnL-F, matK,
rbcL), showing relative branch lengths.
Polyalthia is represented by clades I–IV
(described in text), with clade IV repre-
senting Fenerivia s.l.
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On morphological grounds Maasia seems to be the clos-
est relative of Fenerivia, although the relationships revealed
by molecular data remain obscure due to lack of informative
characters. Both genera share a number of characters, including
inconspicuous leaf venation, axillary flowers, ovaries generally
with a single ovule, seeds with a rib-like raphe and spiniform
endosperm rumination, and monosulcate pollen grains with
psilate-perforate ornamentation (Rogstad, 1989; Rogstad & Le
Thomas, 1989; Schatz & Le Thomas, 1990; Mols & al., 2008).
In addition to the presence of the calyx flange as described
above, Fenerivia differs from Maasia in lacking a glaucous
appearance on the abaxial surface of the leaves and in having
significantly larger petals.
The polyphyletic genus Polyalthia has been significantly
reduced in scope in recent years as a result of molecular phy-
logenetic analyses, with the transfer of six species to Maasia
(Mols & al., 2008) and five species to Marsypopetalum (Xue
& al., 2011). The transfer of nine Madagascan Polyalthia spe-
cies to Fenerivia, as proposed here, further reduces the size of
Polyalthia, although it does not render the genus monophyletic.
Further research remains to address the status of Haplostichan-
thus species (associated with the “true” Polyalthia clade II,
inclusive of the type species, P. subcordata) and also the
Polyalthia species associated with Enicosanthum and Miliusa
(clades I and III, respectively).
TAXONOMIC TREATMENT
Fenerivia Diels in Notizbl. Bot. Gart. Berlin-Dahlem 9: 355.
15 Jun 1925 – Type: Fenerivia heteropetala Diels.
Small to medium-sized trees. Leaves with inconspicuous
venation, glabrous, not glaucous abaxially. Inf lorescences ax-
illary, reduced to a solitary flower; calyx highly reduced and
vestigial, present as basal flange (sometimes 3-lobed); petals in
3 whorls; outer petals (presumed bracteopetals) 3 per flower,
ovate, sepal-like; middle petals (presumed bracteopetals) 3
per flower, narrowly elliptic to linear; inner petals (presumed
andropetals) generally 3 per flower (6 per flower in F. het-
eropetala), narrowly elliptic to linear. Stamens numerous per
flower, dehiscence extrorse; connectives truncate. Carpels up
to 45 per flower; ovule solitary, basal. Fruits with persistent
calyx flange; monocarps stipitate, ellipsoid, with single seed.
Fenerivia angustielliptica (G.E. Schatz & Le Thomas)
R.M.K. Saunders, comb. nov. ≡ Polyalthia angustiellip-
tica G.E. Schatz & Le Thomas in Bull. Mus. Natl. Hist.
Nat., B, Adansonia 12: 120–121, fig. 1 (7–15), 3 (5–6). 1990
(“P. angusti-elliptica”).
Fenerivia capuronii (Cavaco & Keraud ren) R.M.K. Saunders,
comb. nov. ≡ Polyalthia capuronii Cavaco & Keraudren
in Bull. Jard. Bot. État Bruxelles 27: 75, t. 1, fig. 3. 1957.
Fenerivia chapelieri (Baill.) R.M.K. Saunders, comb. nov. ≡
Polyalthia chapelieri Baill. in Adansonia 8: 349. 1868 ≡
Unona chapelieri (Baill.) Baill. in Bull. Mens. Soc. Linn.
Paris 1: 338. 1882.
= Polyalthia lucens Baker in J. Linn. Soc., Bot. 21: 318–319.
1884 [synonymy according to Schatz & Le Thomas, 1990:
120].
= Polyalthia lamii Cavaco & Keraudren in Bull. Jard. Bot. État
Bruxelles 27: 76. 1957 [synonymy according to Schatz
& Le Thomas, 1990: 120].
Fenerivia emarginata (Diels) R.M.K. Saunders, comb. nov. ≡
Polyalthia emarginata Diels in Notizbl. Bot. Gar t. Berlin-
Dahlem 9: 340–341. 1925.
Fenerivia ghesquiereana (Cavaco & Keraud ren) R.M.K. Saun-
ders, comb. nov. ≡ Polyalthia ghesquiereana Cavaco
& Keraudren in Bull. Jard. Bot. État Bruxelles 27: 77, t. 1,
fig. 2. 1957.
Fenerivia heteropetala Diels in Notizbl. Bot. Gart. Berlin-
Dahlem 9: 356. 1925 ≡ Polyalthia heteropetala (Diels)
Ghesq. in Rev. Zool. Bot. Africaines 32: 142. 1939.
Fig. 3.
Fenerivia heteropetala, showing ovate “outer petals” (OP) and
li near “inner petals” (IP) of mi xed or igin (discu ss ed in text). Although
the drawing shows 10 linear petals, the type specimen and descrip-
tions of the species indicate that there are only nine. Redrawn from
Cavaco & Keraudren (1958: fig. XIV) by Vatsala Mirnaalini; repro-
duced from Saunders (2010).
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60 (5) • October 2011: 1407–1416
Fenerivia humbertii (Cavaco & Keraudren) R.M.K. Saun-
ders, comb. nov. ≡ Polyalthia humbertii Cavaco & Kera-
udren in Bull. Jard. Bot. État Bruxelles 27: 76, t. 1, fig.
1. 1957.
= Polyalthia leandrii Cavaco & Keraudren in Bull. Jard. Bot.
État Bruxelles 27: 76. 1957 [synonymy according to Schatz
& Le Thomas, 1990: 122].
Fenerivia madagascariensis (Cavaco & Keraudren)
R.M.K. Saunders, comb. nov. ≡ Polyalthia madagas-
cariensis Cavaco & Keraudren in Bull. Jard. Bot. État
Bruxelles 27: 78. 1957.
Fenerivia oligosperma (Danguy) R.M.K. Saunders, comb.
nov. ≡ Artabotrys oligospermus Danguy in Bull. Mus.
Fig. 4.
Flower and fruit morphology in Fenerivia.
A–B,
post-fertilization flower (A) and fruit (B) of Fenerivia angustielliptica, showing flange
lobes (arrowed) on receptacle.
C– F,
flowers (C–E) and fruit (F) of Fenerivia capuronii, showing flange lobes (arrowed) on receptacle. Voucher
specimens: A, B, Pon cy 1540 (MO); C, E, Chatrou 669 (TAN, WAG); D, Chatrou 667 (TAN, WAG); F, Chatrou 668 (TAN, WAG). Photographs:
A, B, © Odile Poncy; C–F, © Lars Chatrou.
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Bân, N.T. 1975. A new genus of the Annonaceae Juss.—Enicosanthel-
lum Bân. Bot. Žhurn. (Moscow & Leningrad) 60: 808–812. [in
Russian]
Blume, C.L . 1830. Flora Javae [Anonaceae]. Brussels: J. Frank.
Cavaco, A. & Keraudren, M. 1957. Notes systématiques et bio-
géographiques sur les An nonacées de Madagascar et des Comores.
Bull. Jard. Bot. État Bruxelles 27: 59–93.
Cavaco, A. & Keraudren, M. 1958. Famille Annonacées (Annona-
ceae). Pp. 1–109 in: Humbert, H. (ed.), Flore de Madagascar et
des Comores. Paris: Didot & Cie.
Chaowasku, T., Mols, J. & Van der Ham, R. 2008. Pollen morphol-
ogy of Miliusa and relatives (Annonaceae). Grana 47: 175–184.
Deroin, T. 2007. Floral vascular pattern of the endemic Malagasy genus
Fenerivia Diels (Annonaceae). Adansonia 29: 7–12.
Diels, L. 1925. Revisio Anonacearum madagascariensum. Notizbl. Bot.
Gart. Berlin-Dahlem 9: 334–357.
Doyle, J. & Doyle, J.L. 1987. A rapid DNA isolation method for small
quantities of fresh tissues. Phytochem. Bull. 19: 11–15.
Doyle, J.A., Bygrave, P. & Le Thomas, A. 2000. Implicat ions of mo-
lecular data for pollen evolution in Annonaceae. Pp. 259–284 in:
Harley, M.M., Morton, C.M. & Blackmore, S. (eds.), Pollen and
spores: Morphology and biology. Kew: Royal Botanic Gardens.
Doyle, J.A. & Le Thomas, A. 19 94. Cladistic analysis and pollen evolu -
tion in Annonaceae. Acta Bot. Gallica 141: 149–170.
Doyle, J.A. & Le Thomas, A. 1995. Evolution of pollen characters
and relationships of African Annonaceae: Implications of a cla-
distic analysis. Pp. 241–254 in: Le Thomas, A. & Roche, E. (eds.),
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Doyle, J.A. & Le Thomas, A. 1996. Phylogenetic analysis and chara ct er
evolution in Annonaceae. Bull. Mus. Natl. Hist. Nat., B, Adansonia
18: 279–334.
Engler, A. & Diels, L. 1901. Annonaceae. Pp. 1–96 in: Engler, A. (ed.),
Natl. Hist. Nat. 28: 247. 1922 ≡ Polyalthia oligosperma
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341–342. 1925.
Fenerivia richardiana (Baill.) R.M.K. Saunders, comb. nov.
≡ Polyalthia richardiana Baill. in Adansonia 8: 350. 1868
≡ Unona richardiana (Baill.) Baill. in Bull. Mens. Soc.
Linn. Paris 1: 339. 1882.
= Polyalthia dielsii Cavaco & Keraudren in Bull. Jard. Bot.
État Bruxelles 27: 77. 1957 [synonymy according to Schatz
& Le Thomas, 1990: 124].
ACKNOWLEDGEMENTS
Financial support was provided by a grant from the Hong Kong
Research Grants Council (HKU 775009M) and a grant from the
University of Hong Kong Research Committee, both awarded to
RMKS. We are grateful to: La rs Chatrou and Odile Poncy for allowi ng
permission to re produce their photographs; the directors and cur at or s
of P and WAG herbaria for access to their collections and/or for provid-
ing leaf samples for DNA extraction; three anonymous reviewers for
their comments on the manuscript; and Tanawat Chaowasku, Thierry
Deroin, and George Schatz for useful discussion.
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Appendix.
Voucher information and GenBank accession numbers for samples used in this study (*newly sequenced data).
Species, origin, voucher, GenBank accession number (in following order: trnL-F, matK, rbcL)
Alphonsea boniana Finet & Gag nep.: Vie tn am , P.J.A. Keßler 3116 (L), AY3190 77, AY518809, AY318965; Alphonsea elliptica Hook. f. & Thomson: cultivated,
Kebun Raya, Bogor [X-F-52], Van Balgooy 5141 (L), AY319078, AY518807, AY318966; Alphonsea kinabaluensis J. Sinclair: Malaysia, Ridsdale DV-S-3048
(L), AY319080, AY518811, AY318968; Anaxagorea silvatica R.E. Fr.: Brazil, P.J.M. Maas & al. 8836 (U), AY743458, AY743477, AY743439; Annickia chlo-
rantha (Oliv.) Setten & Maas: Gabon, M.S.M. Sosef 1877 (WAG), AY841671, AY841393, AY841594; Annickia pilosa (Exell.) Setten & Maas: Gabon, M.S.M.
Sosef 1803 (WAG), AY743469, AY743488, AY743450; Bocageopsis multif lora (Mart.) R.E. Fr.: Guyana, M.J. Jansen-Jacobs & al. 5789 (U), AY841678,
DQ018262, AY841600; Cananga odorata (Lam.) Hook. f. & Thomson: Costa Rica, L.W. Chatrou & al. 93 (U), AY841680, AY841394, AY841602; Cleistop-
holis glauca Engl. & Diels: Gabon, J.J. Wieringa & al. 3278 (WAG), AY841681, AY841395, AY841603; Cremastosperma brevipes (DC.) R.E. Fr.: French
Guiana, U. Scharf 76 (U), AY743573, AY743550, AY743527; Cremastosperma cauliflorum R.E. Fr.: Peru, L.W. Chatro u & al. 224 (U ), AY743565, AY743542,
AY743519; Cyathocalyx martabanicus Hook. f. & Thomson: cultivated, Kebun Raya, Bogor, J.B. Mols 11 (L), AY841683, DQ125054, AY841605; Desmopsis
microcarpa R.E. Fr.: Costa Rica, L.W. Chatrou & al. 85 (U), AY319173, AY518804, AY319059; Desmopsis schippii Standl.: Costa Rica, L.W. Chatrou & al.
94 (U), AY319174, AY518805, AY319060; Enicosanthum fuscum (King) Airy-Shaw: Thailand, P.J.A. Keßler 3222 ( L), AY319085, AY518787, AY318973;
Enicosanthum membranifolium J. Sinclair: Thailand, P.J.A. Keßler 3198 (L), AY319086, AY518788, AY318974; Enicosanthum paradoxum Becc.: Borneo,
P.J.A. Keßler & Arifin PK 2746 (L), AY319 087, AY518789, AY318975; Ephedranthus sp.: Brazil, Maas & al. 8826 (U), AY841694, AY841396, AY841616;
Fenerivia angustielliptica (G.E. Schatz & Le Thomas) R.M.K. Saunders (= Polyalthia angustielliptica G.E. Schatz & Le Thomas): Madagascar, O. Poncy
1540 (P),* JF810397, JF810373, JF810385; Fenerivia capuronii (Cavaco & Keraudren) R.M.K. Saunders (= Polyalthia capuronii Cavaco & Keraudren):
Madagascar, P.P. Lowry II 4436 (WAG),* JF810398, JF810374, JF810386; Fenerivia chapelieri (Baill.) R.M.K. Saunders (= Polyalthia chapelieri Baill.):
Madagascar, Ludovic & Rallotoarivony 221 (P),* JF810399, JF810375, JF810387; Fenerivia emarginata (Diels) R.M.K. Saunders (= Polyalthia emarginata
Diels): Madagascar, J.S. Miller 3527 (WAG),* JF810400, JF810376, JF810388; Fenerivia ghesquiereana (Cavaco & Ke raud ren) R. M.K. Saunde rs (= Polyalthia
ghesquiereana Cavaco & Keraudren): Madagascar, R. Randrianaivo 282 (WAG),* JF810401, JF810377, JF810389; Fenerivia heteropetala Diels (= Polyalthia
heteropetala (Diels) Ghesq.): Madagascar, G. Cours 2606 (P),* JF810402, JF810378, JF810390; Fenerivia humbertii (Cavaco & Keraudren) R.M.K . Saunders
(= Polyalthia humbertii Cavaco & Keraudren): Madagascar, S.T. Malcomber & A.J.M. Leeuwenberg 1173 (WAG),* JF810403, JF810379, JF810391; Fenerivia
madagascariensis (Cava co & Keraudren) R.M.K. Saunder s (= Polyalthia madagascariensis Cava co & Keraudren): Ma da ga scar, J. Rabenantoandro & A. Monja
1590 (P),* JF810404, JF810380, JF810392; Fenerivia oligosperma (Danguy) R.M.K. Saunders (= Polyalthia oligosperma Danguy): Madagascar, Ursch s.n.
(WAG),* JF810405, JF810381, JF810393; Fenerivia richardiana (Baill.) R.M.K. Saunders (= Polyalthia richardiana Baill.): Madagascar, P. Antilahimena 157
(WAG),* JF810406, JF810382, JF810394; Fitzalania heteropetala (F. Muell.) F. Muell.: Australia, T.H. Kemp 7267 (L), AY773282, AY773280, AY773281;
Greenwayodendron oliveri (Engl.) Verdc.: Ghana, C.C.H. Jongkind & al. 1795 ( WAG), AY743470, AY743489, AY743451; Haplostichanthus longirostris
(Scheff.) Heusden: New Guinea, W. Takeuchi & D. Ama 15656 (L), AY319091, AY518826, AY318979; Klarobelia inundata Chatrou: Peru, L.W. Chatrou & al.
205 (U ), AY743471, AY743490, AY743452; Maasia discolor (Diels) Mols & al.: New Gui nea, W. Takeuchi & D. Ama 16394 (L), AY319135, AY518872, AY3190 21;
Maasia glauca (Hassk.) Mols & al.: cultivated, Kebun Raya, Bogor [XX-D-81], J.B. Mols 20 (L), AY319137, AY518871, AY3190 23; Maasia multinervis (D iels)
Mols & al.: New Guinea, S.H. Rogstad 819 (WAG),* JF810407, JF810383, JF810395; Maasia ovalifolia (S.H. Rogstad) Mols & al.: Bor neo, P.S. Ashton 7863
(WAG),* JF810408, JF810384, JF810396; Maasia sumatrana (Miq.) Mols & al.: Borneo, unknown collector SAN 143 918 (SAN), AY319153, AY518873,
AY319039; Malmea dielsiana R.E. Fr.: Peru, L.W. Chatrou & al. 122 (U), AY231288, AY238964, AY238955; Marsypopetalum littorale (Blume) B. Xue &
R.M.K. Saunders: cultivated, Kebun Raya, Bogor [IV-H-53], Ra stini 153 (L), AY319140, AY518835, AY319026; Marsypopetalum pallidum (Blume) Backer:
Thailand, P.J.A. Keßler 3192 (L), AY319092, AY518834, AY318980; Meiocarpidium lepidotum Engl. & Diels: Gabon, F.J. Breteler 13947 (WAG), EU169776,
EU16 9687, EU169754; Meiogyne cylindrocarpa (Burck) Heusden: Borneo, C.E. Ridsdale DV-M1-1930 (L), AY319093, AY518796, AY318981; Meiogyne
stenopetala (F. Muell.) Heu sden: Aust ralia , L. Jessup 706 (K), AY319083, AY773279, AY318971; Meiogyne virgata (Blume) Miq.: Borneo, P.J.A. Keßler 2751
(L), AY319094, AY518798, AY318982; Miliusa campanulata Pierre: Tha il and, P. Chalermglin 440407-11 (TISTR), AY319096, AY5188 42, AY31898 4; Miliusa
cuneata Craib: Thailand, P. Chalermglin 440214-7 (L), AY319097, AY518844, AY318985; Miliusa horsfieldii (Benn.) Pierre: cultivated, Kebun Raya, Bogor,
J.B. Mols 1 (L), AY319098, AY518849, AY318986; Miliusa lineata (Craib) Ast: Thailand, P.J.A. Keßler 3202 (L), AY319099, AY518848, AY318987; Miliusa
longipes King: Thailand, P. Pholsena 2651 (L), AY319100, AY318988, AY318988; Miliusa velutina (Dunal) Hook. f. & Thomson: Thailand, P. Pholsena 2842
(L), AY319105, AY518847, AY318993; Mitrephora alba Ridl.: Thailand, P. Chalermglin 440304-1 (Thailand Institute of Scientific and Technological Research,
Bangkok), AY319106, AY518855, AY318994; Mitrephora keithii Ridl.: Thailand, P.J.A. Keßler 3190 (L) & D.J. Mid dleton & al . 877 (L), AY319108, AY518857,
AY318995; Mitrephora polypyrena (Blume) Miq.: cultivated, Kebun Raya, Bogor, J.B. Mols 7 (L), AY319110, AY518858, AY318997; Monocarpia euneura
Miq.: Borneo, F. Slik 2931 (L), AY319111, AY518865, AY318998; Mosannona costaricensis R.E. Fr.: Costa Rica, L.W. Chatrou & al. 90 (U), AY743496,
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60 (5) • October 2011: 1407–1416Saunders & al. •
Enlarging the Madagascan genus Fenerivia
AY743503, AY743510; Mwasumbia alba Couv reur & D.M. Johnso n: Tanzania, T.L.P. Couvreur 85 (WAG), EU747674, –, EU747680; Neo-uvaria acuminatis-
sima (Miq.) Airy-Shaw: Borneo, C.E. Ridsdale DV-SR-4671 (L), AY319112, AY518793, AY318999; Neo-uvaria parallelivenia (Boerl.) H. Okada & K. Ueda:
cultivated, Kebun Raya, Bogor, P.J.A. Keßler sub IV-H-73 (L), AY319113, AY518794, AY319000; Onychopetalum periquino (Rusby) D.M. Johnson & N.A.
Murray: Bolivia, L.W. Ch atrou & al. 425 (U), AY319179, AY518876 , AY319065; Orophea celebica Miq.: Su lawesi, P.J.A. Keßler 2953 (L), AY319117, AY518814,
AY319 004; Orophea enterocarpa Maingay ex Hook. f.: Thailand, P. Chalermglin 440403 (Thailand Institute of Scientific and Technological Research, Bang-
kok), AY319119, AY518815, AY319006; Orophea kerrii P.J.A. Kessler: Thailand, P. Ch alermglin 440416-1 (L), AY319121, AY518818, AY319008; Orophea
polycarpa A. DC.: Thailand, P.J.A. Keßler 3234 (L), AY319123, AY518819, AY319010; Oxandra espintana (Spruce ex Benth.) Baill.: Per u, L.W. Chatrou &
al . 133 (U), AY319180, DQ01826 0, AY31906 6; Phaeanthus ebracteolatus (Presl) Merr.: New Gui nea, T.M.A. Utteridge 17 (K), AY319125, AY518863, AY319012;
Phaeanthus splendens Miq.: Borneo, P.J.A. Keßler B 1564 (L), AY319126, AY518864, –; Piptostigma mortehani De Wild.: Gabon, J.J. Wieringa 2779 (U),
AY743473, AY743492, AY743454; Platymitra macrocarpa Boerl.: cultivated, Kebun Raya, Bogor [IV-G-53], H. Okada 3457 ( L), AY319127, AY518812 ,
AY319 013; Polyalthia borneensis Merr.: Bor neo, C.E . Rids dale DV- SR-7921 (L), AY319128, AY518821, AY319014; Polyalthia cauliflora Hook. f. & Thomson:
cultivated, Singapore Botanical Garden, P.J.A. Keßler 3114 (L), AY319129, AY518803, AY319015; Polyalthia celebica Miq.: cultivated, Kebun Raya, Bogor
[IV-C-97], J.B. Mols 9 (L), AY319130, AY518827, AY319016; Polyalthia cerasoides (Roxb.) Be dd .: Thailand, P. Chalermglin 440214-4 (L), AY319131, AY518854,
AY319017; Polyalthia cinnamomea (Hook. f. & Thomson) Hook. f. & Thomson: Borneo, C.E. Ridsdale DV-M1-347 (L), AY319132, AY518828, AY319018;
Polyalthia coffeoides (Thwaites ex Hook. f. & Thomson) Thwaites: Sri Lanka, R.M.C.S. Ratnayake 1/03 (HKU), EU522178, EU522233, EU522288; Polyalthia
congesta (Ridl.) J. Sinclair: Borneo, C.E. Ridsdale DV-S-5105 (L), AY319133, AY518790, AY319019; Polyalthia debilis (Pierre) Finet & Gagnep.: Thailand,
P.J.A. Keßler 3228 (L), AY319134 , AY518832, AY31902 0; Polyalthia flagellaris (Becc.) Airy-Shaw: Borneo, Duling 38 (K), AY319136, AY518824 , AY319022;
Polyalthia cf. glabra J. Sinclair: cultivated, Kebun Raya, Bogor [IV-H-106], Rastini 224 (L), AY319146, AY518782, AY319032; Polyalthia korinti (Dunal)
Thwaites: Sri Lanka, R.M.C.S. Ratnayake 2/03 (HKU), EU522179, EU522234, EU522289; Polyalthia laterif lora Kurz: cultivated, Kebun Raya, Bogor [XII-
B-VII-37) (L), AY319138, AY518781, AY319024; Polyalthia longifolia (Sonn.) Thwaites: Sri Lanka, R.M.C.S. Ratnayake 3/04 (HKU), EU522180, EU522235,
EU522290; Polyalthia cf. longifolia (Sonn.) Thwaites: Sulawesi, J.B. Mols 14 (L), AY319139, AY518785, AY319025; Polyalthia longipes (Miq.) Koord. &
Valeton: Borneo, C.E. Ridsdale DV-M2-11443 (L), AY319142, AY518829, AY319028; Polyalthia obliqua Hook. f. & Thomson: Borneo, Ambriansyah & Arifin
AA 1694 (L), AY319143, AY518822, AY319029; Polyalthia pendula G.E. Schatz & Le Thomas: Madagascar, Rabevohitra 2386 (K, WAG), AY319144, AY518852,
AY319030; Polyalthia rumphii (Hensch.) Merr.: Sumatra, M.M.J. van Balgooy & A.K. van Setten 5654 (L), AY319145, AY518791, AY319031; Polyalthia
sclerophylla Hook. f. & Thomson: cultivated, Kebun Raya, Bogor [XX-D-82] (L), AY319147, AY518783, AY319033; Polyalthia stenopetala (Hook. f. &
Thomson) Finet & Gagnep.: Thailand, P. Chaler mglin 440302 (Thailand Institute of Scientific and Technological Research, Bangkok), AY319148, –, AY319034;
Polyalthia stuhlmannii (Engl.) Verd.: Kenya, Luke & Robertson 1424 (K, WAG), AY319149, AY518853, AY319035; Polyalthia subcordata (Blu me) Blume
(I): Java, B. Gravendeel & al. 549 (L), AY319150, AY518830, AY319036; Polyalthia subcordata (Blume) Blume (II): Java, B. Gravendeel & al. 678 ( L),
AY319151, AY518831, AY319037; Polyalthia suberosa (Roxb.) Thwaites: India, L.W. Chatrou 480 (U) & cultivated, Botanical Gardens Kaiserslautern,
anonymous s.n., AY319152, AY518833, AY319038; Polyalthia viridis Craib: Thailand, P. Chalermglin 440214-3 (L), AY319154, AY518784, AY319040; Poly-
althia xanthopetala Merr.: Borneo, C.E. Ridsdale DV-S-5107 (L), AY319155, AY518792, AY319041; Polyalthia sp.: Borneo, C.E. Ridsdale DV-M1-12314 (L),
AY319 08 4, AY518825, AY318972; Polyceratocarpus pellegrinii Le Thomas: Ca meroon, J.J.E. de Wilde 8718 (WAG), EU747678, –, EU747684; Popowia hirta
Miq.: Borneo, P.J.A. Keßler B 1628 (L), AY319156, AY518860, AY319042; Popowia odoardoi Diels: Borneo, C.E. Ridsdale DV-SR-7422 (L), AY319157,
AY5188 61, AY319043; Popowia pisocarpa (Blume) Endl.: Jav a, M.M.J. van Balgooy & A.K. van Setten 5683 (L), AY319158, AY518862, AY319044; Pseudephe-
dranthus fragrans (R.E. Fr.) Ar isteg.: Venezuela, P.J.M. Maas & al. 6878 (U ), AY841729, –, AY841651; Pseudomalmea diclina (R.E. Fr.) Chat rou: Per u, L.W.
Chatrou & al. 211 (U), AY319128, AY841398, AY319068; Pseudoxandra lucida R.E. Fr.: Peru , L.W. Chatrou & al. 213 (U), AY319190, AY319190, AY319076;
Pseuduvaria acerosa Y.C.F. Su & R.M.K. Sau nder s: New Guinea, J.S. Womersley NGF 43899 (L), EU522181, EU522236, EU522291; Pseuduvaria borneensis
Y.C.F. Su & R.M.K. Saunders: Borneo, P.S. Ashton S 18415 (L), EU522184, EU522239, EU522294; Pseuduvaria fragrans Y.C.F. Su & al.: Thailand, T. Chao-
was ku 27 (L), EU522231, EU522286, EU522341; Pseuduvaria froggattii (F. Muell .) Jessu p: Au st ralia, A. Ford & L. Cinelli 4776 (H KU ), EU52219 6, EU522 251,
EU522306; Pseuduvaria galeata J. Sinclair: Peninsular Malaysia, J. Sinclair 10689 (L), EU522197, EU522252, EU522307; Pseuduvaria megalopus (K. Schum.)
Y.C.F. Su & J.B. Mols: New Guinea, W. Takeuchi & D. Ama 16235 (L), EU522208, EU522263, EU522318; Pseuduvaria pamattonis (Miq.) Y.C.F. Su &
R.M.K. Saunders: Philippines, A.C. Podzorski SMHI 520 (L), EU522217, EU522272, EU522327; Pseuduvaria setosa (King) J. Sincl ai r: Thailand , J.F. Maxwell
86-208 (L), EU522224, EU522279, EU522334; Ruizodendron ovale (Ruiz & Pav.) R.E. Fr.: Ecuador, P.J.M. Maas & al. 8600 (U), AY841735, –, AY841657;
Sageraea lanceolata Miq.: Borneo, C.E. Ridsdale DV-M2-1692 (L), AY319164, AY518799, AY319050; Sapranthus microcarpus (Donn. Sm.) R.E. Fr.: Hon-
duras, P.J. M. Maas & al. 8457 (U ), AY319166, AY518806, AY319052; Sapranthus viridiflorus G.E. Schatz: Costa Rica, L.W. Cha trou & al. 55 (U), AY319165,
AY743493, AY319051; Stelechocarpus burahol (Blume) Hook. f. & Thomson: cultivated, Kebun Raya, Bogor, J.B. Mols 13 (L), AY319167, AY518803, AY319053;
Stelechocarpus cauliflorus (Scheff.) R.E. Fr.: cultivated, Kebun Raya, Bogor [XV-A-196] (L), AY319168, AY518800, AY319054; Stenanona costaricensis
R.E. Fr.: Costa Rica, L.W. Chatrou & al. 67 (U), AY319183, AY518801, AY319069; Stenanona panamensis Standl.: Costa Rica, L.W. Chatrou & al. 100 (U),
AY31918 4, AY518802, AY319070; Tetrameranthus duckei R.E. Fr.: Brazil, D.W. Stevenson & al. 1002 (U), AY841736, –, AY841658; Tridimeris hahniana
Baill.: Mexico, G.E. Schatz 1198 (K), AY319169, –, AY319055; Trivalvaria macrophylla (Blume) Miq.: Java, M. Chase 1207 (K), AY319170, –, AY319056;
Unonopsis stipitata Diels: Peru, L.W. Chatrou & al. 253 (U), AY841740, AY841400, AY841662; Woodiellantha sp.: Borneo, Lugas 311 (K), AY841743, –,
AY841665.
Appendix.
Continued.
