Taxonomic Studies on Malagasy Dalbergia (Fabaceae). IV.

A New Species from Central and Southern Madagascar

and a Narrowed Circumscription for D. emirnensis
Nivohenintsoa Rakotonirina
Mention de Biologie et Écologie Végétales, Université d’Antananarivo,
Antananarivo 101, Madagascar. nivo.rakotonirina@mobot.mg

Peter B. Phillipson
Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, U.S.A.; Institut de
Systématique, Évolution, et Biodiversité (ISYEB), Muséum National d’Histoire Naturelle, Centre
National de la Recherche Scientifique, Sorbonne Université, École Pratique des Hautes Études,
Université des Antilles, C.P. 39, 57 rue Cuvier, 75005 Paris, France. peter.phillipson@mobot.org

Simon Crameri
Institute of Integrative Biology, ETH Zurich, Zurich, Switzerland; Missouri Botanical Garden,
4344 Shaw Blvd., St. Louis, Missouri 63110, U.S.A. sfcrameri@gmail.com

Nicholas Wilding
Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, U.S.A.; Institut de
Systématique, Évolution, et Biodiversité (ISYEB), Muséum National d’Histoire Naturelle, Centre
National de la Recherche Scientifique, Sorbonne Université, École Pratique des Hautes Études,
Université des Antilles, C.P. 39, 57 rue Cuvier, 75005 Paris, France. nwilding@mobot.org

Porter P. Lowry II
Missouri Botanical Garden, 4344 Shaw Blvd., St. Louis, Missouri 63110, U.S.A.; Institut de
Systématique, Évolution, et Biodiversité (ISYEB), Muséum National d’Histoire Naturelle, Centre
National de la Recherche Scientifique, Sorbonne Université, École Pratique des Hautes Études,
Université des Antilles, C.P. 39, 57 rue Cuvier, 75005 Paris, France. pete.lowry@mobot.org

Bakolimalala Rakouth
Mention de Biologie et Écologie Végétales, Université d’Antananarivo,
Antananarivo 101, Madagascar. ba.rakouth@gmail.com

Richardson Razakamalala
Missouri Botanical Garden, B.P. 3391, Antananarivo 101, Madagascar.
richard.razakamalala@mobot.mg

Abstract. Dalbergia emirnensis Benth. was previ­
ously treated as comprising two entities, the typical
variety from the Central Highlands of Madagascar and
D. emirnensis var. decaryi Bosser & R. Rabev. from the
southern part of the country at lower altitudes. Addi­
tional collections and analyses demonstrate that, as
currently delimited, these taxa exhibit considerable,
discontinuous variability in morphology, habitat, and
distribution, a finding supported by a recent phyloge­
nomic study. We show that the material previously in­
cluded in the typical variety of D. emirnensis repre­
sents two morphologically and geographically distinct
entities, one that occurs only in the Central Highlands
of Madagascar and includes the type specimen, and
another that is morphologically and genetically more
akin to D. emirnensis var. decaryi, found in the South­
ern Highlands of Madagascar. We describe the latter
as a new species, D. nemoralis Rakoton., Phillipson &
Crameri, in which we include D. emirnensis var. de-
caryi, recognizing it as a distinct subspecies. We also

Version of record first published online on 00 Month 2023.

doi: 10.3417/2023790 Novon 31: 73–87.

06_Novon 31_790.indd 73 4/24/23 10:33 AM

 74
show that D. campenonii Drake should be treated as a
synonym of D. emirnensis. Each of the accepted taxa is
provided with a full description, a comprehensive list­
ing of specimens examined, details of its geographic
distribution and habitat, and a provisional risk of ex­
tinction assessment using the IUCN Red List catego­
ries and criteria; a line drawing is also provided for
D. nemoralis subsp. nemoralis.
Résumé. Deux entités étaient reconnues auparavant
au sein de Dalbergia emirnensis Benth., la variété typi­
que venant des Hautes Terres centrales de Madagascar
et D. emirnensis var. decaryi Bosser & R. Rabev. de la
partie sud de l’île, à des altitudes plus basses. Des co­
llectes nouvelles et des analyses supplémentaires mon­
trent que, tels que délimités actuellement, ces taxons
présentent une variabilité considérable et discontinue
dans leurs morphologies, leurs habitats et leurs aires de
répartition, une constatation qui est soutenue par une
étude phylogénomique récente. Nous montrons que le
matériel précédemment inclus dans la variété typique
de D. emirnensis représente deux entités morphologi­
quement et géographiquement distinctes, l’une distri­
buée seulement dans les Hautes Terres centrales qui
correspond au type, et l’autre qui est morphologique­
ment et génétiquement plus apparentée à D. emirnensis
var. decaryi, et que l’on trouve dans les Hautes Terres
du Sud de Madagascar. Nous décrivons cette dernière
en tant qu’une nouvelle espèce, D. nemoralis Rakoton.,
Phillipson & Crameri dans laquelle nous reconnais­
sons D. emirnensis var. decaryi comme une sous-espèce
distincte. Par ailleurs, nous montrons que D. campeno-
nii Drake est mieux placé en synonymie sous D. emir-
nensis. Chacun des taxons acceptés est accompagné
d’une description complète, d’une liste compréhensive
de spécimens examinés, des détails sur sa répartition
géographique, son habitat, et une évaluation provisoire
du risque d’extinction selon les catégories et les cri­
tères de la Liste rouge de l’UICN; un dessin à trait est
fourni pour D. nemoralis subsp. nemoralis.
Key words: Circumscription, Dalbergia emirnensis,
Fabaceae, IUCN Red List, Madagascar, new species,
taxonomy.
The pantropical genus Dalbergia L. f. (Fabaceae,
subfamily Papilionoideae) was described by Linnaeus
filius in 1782 (“1781”) based on D. lanceolaria L. f.
from Sri Lanka. Its centers of diversity are in Africa,
the Americas, and Asia, and the genus comprises ca.
275 currently recognized species, with approximately
42% of them occurring in Africa (Cowell et al., 2022;
POWO, 2022). Members of Dalbergia occur in diverse
habitats, including moist and dry tropical forests, sa­
vannas, coastal dunes, and rocky outcrops, and the
genus encompasses a diversity of life forms including
06_Novon 31_790.indd 74
Novon
shrubs, trees, and lianas (de Carvalho, 1997; Bosser &
Rabevohitra, 2002; Klitgaard & Lavin, 2005).
Dalbergia species are widely recognized for their
much sought-after wood, generally known as rosewood,
although this term is also applied to ornamental timber
species from other genera (UNODC, 2016). Rosewood,
so named because of the fragrant smell of the wood of
certain species when cut (Normand, 1988; Klitgaard
& Lavin, 2005; Saha et al., 2013), is highly valued and
has a wide range of uses, including for furniture, boats,
musical instruments, and construction. Its timber is
an important commodity in domestic and international
trade (Barrett et al., 2010; Bergamino, 2013). In Mad­
agascar, the term rosewood (or Bois de rose) is reserved
for the most prized Dalbergia wood, i.e., that obtained
from species with a dark purplish red, patterned heart­
wood that darkens after cutting (Normand, 1988), while
the term palisander (or palissandre) is used to refer to
wood that is typically as durable as rosewood but with
heartwood that is lighter or brownish colored (Mason et
al., 2016). Rosewood and palisander have been har­
vested extensively in Madagascar since the 19th cen­
tury for uses in high-quality flooring, carvings, and fur­
niture (Deschamps, 1961; Schuurman & Lowry, 2009;
Waeber et al., 2019). These beautiful and durable woods
are symbols of wealth and high social status in Mada­
gascar and have long been used to decorate the houses
of the rich (Rakotondramanga, 2002). International de­
mand has been primarily for rosewood (Mason et al.,
2016), whereas in Madagascar, palisander has mostly
served the needs of the local market (Rasamoelina,
2001). The taxonomic identity of the Dalbergia species
that have been most heavily exploited in Madagascar
is largely unknown (Mason et al., 2016), although the
highest quality rosewood reportedly comes from the east­
ern part of the island (Normand, 1988; Ratsimbazafy et
al., 2016; Crameri et al., 2022a) and notably from the
southern half of the Masoala Peninsula, where precious
woods were exploited extensively in the 1960s (Schuur­
man & Lowry, 2009). Minimum quantitative estimates
(Ratsimbazafy et al., 2016) indicated that 152,437 tons
of rosewood (corresponding to about 350,430 trees)
were harvested in the period between 1991 and 2014,
of which 104,655 tons (ca. 240,586 trees) were ex­
ported through 2014, at least 61% of which were from
illegally exploited sources. Cut stumps and small piles
of harvested Dalbergia wood are still routinely encoun­
tered within protected areas (Madagascar Catalogue,
2023), including Masoala National Park, which was
estimated to be the origin of 70% of harvested timber
in the region (Patel, 2007).
Trees exploited for their precious woods in Mada­
gascar have been regarded to be of potential commer­
cial value if they reach a height of at least 20 m or a di­
ameter at breast height (DBH) of at least 20 cm (Schatz
4/24/23 10:33 AM
 Volume 31
2023
et al., 2021). Currently, more than 40 described Mala­
gasy Dalbergia species are known to meet these crite­
ria and are either reported or suspected to produce
wood of commercial value, but until recently very little
was known regarding which species reach an exploit­
able size or whether any of these are under immediate
threat of extinction. The Madagascar Precious Woods
Consortium was established in 2016 to align efforts in
the fields of taxonomy, wood anatomy, DNA sequence
analysis, wood spectroscopic and spectrometric char­
acterization, and conservation. The primary objective
of the consortium is to generate information and pro­
duce reference material and a set of identification tools
that will allow the Malagasy government to implement
improved management of this valuable resource. To this
end, field botanists and taxonomists have worked to
clarify the delimitation of the Malagasy species of Dal-
bergia, which was found to be problematic and in need
of revision (Madagascar Catalogue, 2023).
Five species from Madagascar were described in Dal-
bergia during the first half of the 19th century, but they
were later transferred to Mundulea DC. ex Miq. Ten
years later, D. emirnensis Benth. was the first true spe­
cies of Dalbergia from Madagascar to be formally pub­
lished (Bentham, 1860). Additional species endemic
to the island were published subsequently by several
botanists, including Vatke (1880), who described three
new species; Baillon (1884), who described 13 new
species; Baker (1884, 1886, 1889), who published six
more species; and Drake (1903), who revised the genus
for Madagascar and recognized a total of 25 species, of
which six were new. Following the important increase
in botanical exploration of Madagascar in the early part
of the 20th century, René Viguier further revised the
genus before his death in 1931, describing 11 new spe­
cies in a manuscript that was published posthumously
(Viguier, 1951), but placing others in synonymy and
ultimately recognizing 25 species in a full treatment
of the genus for Madagascar that was never published,
but is available in the archives at the Muséum National
d’Histoire Naturelle in Paris. Several decades later,
the combined efforts of Jean Bosser and Raymond Ra­
bevohitra resulted in the recognition of 48 species
(Bosser & Rabevohitra, 1996, 2002, 2005), 25 of which
were described as new. All of these species are endemic
to Madagascar with the exception of D. bracteolata
Baker, which also occurs in eastern Africa. Our recent
integrative approach, which draws on the abundance
of new field collections now available and insights pro­
vided by phylogenomic analysis of 2396 nuclear regions
(Crameri et al., 2022b), has highlighted problems with
the circumscription of a number of the species recog­
nized by Bosser and Rabevohitra (Crameri, 2020). This
has led to an ongoing effort to develop a revised species-­
level taxonomy for the genus that also incorporates new
06_Novon 31_790.indd 75
Rakotonirina et al. 75
Malagasy Dalbergia (Fabaceae). IV.
information on morphology and eco-geography (Mada­
gascar Catalogue, 2023). Taxonomic work conducted
by Wilding et al. (2021a, 2021b) and Crameri et al.
(2022a) has already resulted in the description of four
additional new species, the resurrection of one species
from synonymy, and emended descriptions for two pre­
viously published species. In the present contribution,
we provide a taxonomic reevaluation of D. emirnensis,
as circumscribed most recently by Bosser and Rabevo­
hitra (1996, 2002). Manuscripts on other groups are in
press or are currently being prepared for publication.
These will include descriptions for an additional eight
new species of potential commercial value (Wilding et
al., 2023) and also the recognition of five infraspecific
taxa as distinct species and the resurrection of two spe­
cies previously placed in synonymy (for more informa­
tion, refer to the online Madagascar Catalogue, 2023).
Dalbergia emirnensis is one of the many species in
Madagascar whose delimitation has proven to be prob­
lematic. Two varieties were recognized by Bosser and
Rabevohitra (1996): D. emirnensis var. emirnensis and
D. emirnensis var. decaryi Bosser & R. Rabev., with the
former stated to occur in the Central Highlands of the
island (sensu Everson et al., 2020), extending as far
south as Betroka in the Anosy Region in the far south­
east, and the latter extending across the south of the
country and along the west coast as far north as Moron­
dava (Menabe Region; Bosser & Rabevohitra, 2002).
Phylogenomic analyses by Crameri (2020) have cast
doubt on the current circumscription of the species
and its two varieties. In that study, 683 individuals,
including representatives of all 48 species of Malagasy
Dalbergia accepted by Bosser and Rabevohitra (2002,
2005), were analyzed, and D. emirnensis (s.l.) was re­
covered as polyphyletic. Crameri (2020) also showed
that material of typical D. emirnensis var. emirnensis
and D. campenonii Drake from the northern part of the
Central Highlands (Analamanga and Bongolava Re­
gions) forms a single well-supported lineage, while the
type of D. emirnensis var. decaryi and other collections
referred to this entity from a broad zone across south­
ern Madagascar are part of a separate clade that in­
cludes D. neoperrieri Bosser & R. Rabev., a species
from western Madagascar. The latter clade belongs to a
well-supported group (the Greveana clade), while the
separately evolving lineage containing D. emirnensis
var. emirnensis and D. campenonii showed an uncer­
tain phylogenomic placement relative to the Greveana
and Peltieri clades, respectively. Material from the
southern part of the Central Highlands of Madagascar,
attributed by Bosser and Rabevohitra (1996, 2002)
to D. emirnensis var. emirnensis, was not sufficiently
represented in the initial phylogenomic study but was
included in a subsequent analysis by Crameri et al.
(unpublished data), together with additional material of
4/24/23 10:33 AM
 76
the other related species. This expanded analysis con­
firmed that this material belongs to the same clade as
D. emirnensis var. decaryi and D. neoperrieri, rather
than to the lineage comprising typical Central High­
lands material of D. emirnensis var. emirnensis. These
results have prompted the taxonomic reevaluation of
D. emirnensis presented here.
Taxonomic Reevaluation of Dalbergia emirnensis
Our examination of material of Dalbergia emirnensis
s.l. in various herbaria with important holdings from
Madagascar (G, K, MO, P, TAN, TEF, ZT; acronyms
according to Thiers, 2022) has revealed numerous mor­
phological differences between the two described vari­
eties of the species, most notably in the size and shape
of their fruits, but also in terms of vegetative morphol­
ogy (see Table 1 for a full comparison). The fruits of
typical D. emirnensis var. emirnensis are approximately
twice the width of those of D. emirnensis var. decaryi.
Bosser and Rabevohitra never saw mature fruits of D.
emirnensis var. emirnensis in collections from around
the type locality in the northern parts of the Central
Highlands of Madagascar, as the only specimens avail­
able from this area at the time were either sterile or
in flower. They evidently assumed that fruits from the
northern subpopulations, including the type locality of
D. emirnensis, would be as small as those observed on
specimens from the southern subpopulations and sim­
ilar also to the material they separated as D. emirnensis
var. decaryi. This incorrect assumption likely led them
to recognize these taxa as varieties of a single species
rather than two separate species. In addition, the lack
of an isotype of D. emirnensis at P possibly contributed
to these authors overlooking the morphological simi­
larity between typical D. emirnensis var. emirnensis and
D. campenonii. Examination of fruiting and flowering
material of D. campenonii and typical D. emirnensis var.
emirnensis, as well as the consideration of ecological
information and the genetic data from Crameri (2020)
and subsequent analyses, leave no doubt that these two
names refer to a single species, which is restricted to
the Central Highlands.
Careful observation of available herbarium material
has also revealed significant morphological differences
between specimens attributed by Bosser and Rabevo­
hitra (1996, 2002) to Dalbergia emirnensis var. emirn-
ensis from the northern versus the southern parts of
the Central Highlands, with respect to leaflet shape in
addition to fruit size. The atypical smaller-fruited ma­
terial from the southern parts, which was included by
Bosser and Rabevohitra (2002) in D. emirnensis var.
emirnensis, is morphologically more akin to D. emirn-
ensis var. decaryi since both entities share leaflets sim­
06_Novon 31_790.indd 76
Novon
ilar in shape (narrowly elliptic or elliptic-oblong vs.
obovate to elliptic) and number ([five to] seven to 11 vs.
six to 10), as well as russet-pubescent indument on the
ovary, and both have small, obovate fruits (although
those of the latter are distinctly narrower). Based on
these findings and the insights from phylogenomic
analyses, we restrict D. emirnensis to the northern sub­
population previously assigned to D. emirnensis var.
emirnensis and describe a new species, D. nemoralis
Rakoton., Phillipson & Crameri, for the southern sub­
populations of D. emirnensis var. emirnensis combined
with the material formerly treated as D. emirnensis var.
decaryi. Within D. nemoralis, we recognize two closely
related subspecies that occupy distinct geographical
zones and are for the most part easily distinguished
morphologically, although they co-occur in a narrow
contact zone, from Sakaraha and Zombitsy to the north­
west, through Ampandrandava in the center, to Anad­
abolava in the southeast of the zone (Fig. 1). Dalbergia
nemoralis subsp. nemoralis is known predominantly
from woodland and mosaics of wooded grassland and
bushland, mostly on lateritic soils formed over igneous
or metamorphic rocks in the Central and Southern
Highlands of Madagascar at higher elevations (mostly
above 600 m). By contrast, D. nemoralis subsp. decaryi
(Bosser & R. Rabev.) Rakoton., N. Wilding, Phillipson
& Crameri, which is based on D. emirnensis var. de-
caryi, occurs further south in intact and degraded dry,
spiny thicket on limestone, fersiallitic, or sandy soils
over southern Madagascar’s tertiary limestones and un­
consolidated sands, generally at lower elevations (mostly
below 600 m; Fig. 1).
Based on morphology alone, some specimens from
the contact zone are difficult to identify to subspecies
(e.g., Du Puy et al. M994, Labat et al. 2115, and SNGF
1942). We suspect that these may be the result of gene
flow between the two subspecies, a hypothesis that is
supported by molecular data, which show intermediate
phylogenomic positions and higher levels of heterozy­
gosity in specimens from near the inferred contact zone
(Crameri, unpublished data). These specimens include
the first two mentioned above as well as four others
(Bernard 2454, Razafimamonjy 8, Service Forestier
(Capuron) 11695, and Seyrig 447), which we identified
at the species rather than subspecies level, following
our integrative approach.
We retain Dalbergia neoperrieri as an eco-geograph­
ically and morphologically distinct species despite its
close relationship to D. nemoralis (see Table 1). Dal-
bergia neoperrieri differs from D. nemoralis most no­
ticeably by its leaflet shape and the shape and size of
its fruit (see Table 1). Moreover, D. neoperrieri occurs
in dry forest and woodland on calcareous substrates
at low elevations in western Madagascar and has been
4/24/23 10:33 AM
 Volume 31 Rakotonirina et al. 77
2023 Malagasy Dalbergia (Fabaceae). IV.

Table 1. Comparison of key morphological characters between Dalbergia emirnensis Benth., D. nemoralis Rakoton., Phil­
lipson & Crameri subspecies, and D. neoperrieri Bosser & R. Rabev.

D. nemoralis D. nemoralis
D. emirnensis subsp. nemoralis subsp. decaryi D. neoperrieri
Number of leaflets (5 to)7 to 11(to 12) (5 to)7 to 11 6 to 10 7 to 11(13)
Petiole length 1.1–2.1 cm 1.1–1.7 cm 0.5–1.1 cm 1.6–2.6 cm
Leaflet shape ovate to elliptic; narrowly elliptic or obovate to elliptic; lanceolate or oblong;
apex acute, elliptic-oblong; apex apex rounded or apex acuminate
mucronulate apiculate to shortly emarginate
acuminate
Lower leaflet surface sparse indument gold-colored velutinous white glaucous bloom pubescent
pubescence indument mainly and fine appressed
along the midrib indument
Paniculate units within 5.9–10 cm 3.8–6.7 cm 11.5–18 cm 8.4–15.5 cm
inflorescence, length
Seeds per fruit; outline 1 to 2; obovate 1 to 2; elliptic, 1 to 3; elliptic to sub- 1 to 2; subrhombic or
shape and width of or elliptic, 0.9–1.3 cm circular, 0.7–0.8 cm elliptic, 2–2.8 cm
single-seeded fruits 1.7–2 cm

recorded mostly from the Melaky Region, with disjunct
occurrences (including the type collection) in the Boeny
Region and scattered localities further south in the
Menabe and Atsimo-Andrefana Regions (Fig. 1). As
such, these two species have adjacent ranges but differ
with respect to their habitat preferences, and although
they share many common features, they are nonetheless
morphologically distinct (see Table 1). Certain rather
poor specimens that we include in D. neoperrieri (Abra-
ham 56, Service Forestier 12701 and 21701) from the
Menabe and Atsimo-Andrefana Regions were identi­
fied by Bosser and Rabevohitra (2002) as D. emirnensis
var. decaryi. However, after careful reexamination of
these collections, we confidently refer them to D. ne-
operrieri. In all other respects, the description of the
latter species and its circumscription, as provided by
Bosser and Rabevohitra (2002), remain valid and we
consider that there is no need to publish an emended
description.
Each of the taxa treated below is provided with a full
description, a comprehensive listing of specimens ex­
amined, and details of its geographic distribution and
habitat, and Dalbergia nemoralis subsp. nemoralis is
illustrated with a line drawing. A preliminary risk of
extinction assessment is included for each taxon fol­
lowing the IUCN Red List categories and criteria (IUCN,
2012), for which calculations of the area parameters
of extent of occurrence (EOO) and area of occupancy
(AOO) were performed using GeoCAT (Bachman et
al., 2011). Because each of the species treated here
can develop into sufficiently large trees to be potential
sources of valuable wood, geo-coordinates and detailed
locality data have been withheld and public access to
this information has been restricted. Full specimen
records (with limited locality data), distribution maps,
and color photos are available on each species page in
the Catalogue of the Plants of Madagascar (Madagascar
Catalogue, 2023).

Taxonomic Treatment

Key to Dalbergia emirnensis and Related Taxa

1. Fruit ≥ 1.7 cm wide (for single-seeded fruits); leaflets often lanceolate to ovate, or sometimes elliptic-oblong.
2. Leaflet lamina ovate to elliptic, apex mucronulate; Central Highlands, in (sub)humid forests and wooded
grassland on sandy or rocky ferralitic soils, at 800–1500 m �������������������������������������������������� 1. D. emirnensis Benth.
2′. Leaflet lamina lanceolate or oblong, apex acuminate; western Madagascar, in dry deciduous forests on calcareous
soils, at 0–500 m������������������������������������������������������������������������������������������������3. D. neoperrieri Bosser & R. Rabev.
1′. Fruit ≤ 1.3 cm wide (for single-seeded fruits); leaflets elliptic to obovate.
3. Terminal leaflet lamina narrowly elliptic or elliptic-oblong, rachis and midvein with short, velvety, golden
pubescence below; inflorescences with paniculate units 3.8–6.7 cm long; at (350–)600–1600 m. . . . . . . . . . . .
���������������������������������������������������������������������������� 2a. D. nemoralis Rakoton., Phillipson & Crameri subsp. nemoralis
3′. Terminal leaflet lamina elliptic to obovate, rachis and midvein with short, sparse, white indumentum below;
inflorescences with paniculate units 11.5–18 cm long; at 30–600(–900) m . . . . . . . . . . . . . . . . . . . . . . . . . . . .
���������������������������2b. D. nemoralis subsp. decaryi (Bosser & R. Rabev.) Rakoton., N. Wilding, Phillipson & Crameri

06_Novon 31_790.indd 77 4/24/23 10:33 AM

 78 Novon

Figure 1. Map showing the approximate distribution ranges for Dalbergia L. f. species (from north to south): D. neoperrieri
Bosser & R. Rabev. (brown); D. emirnensis Benth. (orange); D. nemoralis Rakoton., Phillipson & Crameri subsp. nemoralis
(blue); D. nemoralis subsp. decaryi (Bosser & R. Rabev.) Rakoton., N. Wilding, Phillipson & Crameri (red); and contact zone of
the two subspecies of D. nemoralis (purple). Elevation classes are indicated as shades of gray.

06_Novon 31_790.indd 78 4/24/23 10:33 AM

 Volume 31
2023
1. Dalbergia emirnensis Benth., J. Proc. Linn. Soc.,
Bot. 4 (Suppl.): 33. 1860. TYPE: Madagascar. Va­
kinankaratra Reg. [Prov. Antananarivo]: Habi ad
Praud Ramassina prov: Emirna, fl., W. Bojer s.n.
(holotype, K [barcode] K000379219!).
Dalbergia campenonii Drake, Hist. Phys. Madagascar 30:
181. 1902 [1903], syn. nov. TYPE: Madagascar. Centre
de Madagascar [without precise locality], P. Campenon
s.n. (lectotype, here designated, P [bc] P00060552!;
isolectotype, P [bc] P00060553!).
Trees to ca. 42 m tall, 72 cm diam., sometimes
multi-stemmed, or shrubs. Branches glabrescent, ro­
bust, pale gray in vivo (golden yellow in sicco), with
raised, white lenticels, bark dark gray-brown. Leaves
alternate, 8.5–12 cm, with (5 to)7 to 11(to 12) alternate
leaflets; petiole and rachis yellow-green in vivo (pale
gray to light brown in sicco), puberulent along the ra­
chis and the petiole; petiole 1.1–2.1 cm; stipules trian­
gular, 3.7–4.7 × 1–1.2 mm, caducous; leaflets 27–51 ×
12–22 mm, the proximal ones sometimes smaller than
the rest, the distal leaflet slightly larger, the others rel­
atively uniform; petiolules 1.5–2.5 mm, yellow-green
in vivo (pale gray in sicco), pubescent; lamina ovate to
elliptic, base cuneate to broadly obtuse, margins slightly
recurved, apex acute, mucronulate, venation simple
brochidodromous, with 7 to 9 principal secondary veins
per side; upper and lower surfaces bright midgreen in
vivo (midbrown in sicco), upper surface slightly glossy,
glabrous, except for pubescence along the midrib, lower
surface matte, with sparse indument, higher-order veins
forming a loose network, the midrib forming a channel
above, prominent below. Inflorescences terminal, a pan­
icle composed of dense, multibranched corymbiform
paniculate units, 5.9–10 cm, decreasing in size toward
the apex, the basal units subtended by well-developed,
pinnate, leaflike bracts, the apical units with reduced,
caducous, scalelike bracts, distal portions curved, sub­
spicate, axes green in vivo (light brown in sicco), pu­
bescent; peduncle 0.5–0.9 cm. Flowers subtended by a
triangular, puberulent, scalelike bract, 0.6–0.8 × 0.4–
0.7 mm; pedicel 0.1–1 mm, pubescent (pedicels of
apical flowers shorter than those of basal flowers); brac­
teoles persistent, ovate with rounded apex, partially
enclosing the base of calyx, 0.5–0.8 × 0.6–0.7 mm,
puberulent, yellow-green in vivo (brown in sicco); calyx
pale green in vivo (yellowish in sicco), puberulent, not
persisting on fruits, 2–3 mm, fused in lower 2/3, the 2
upper lobes fused to form a single rounded unit, 0.9–1
× 1–1.6 mm, apex emarginate, the 2 lateral lobes trian­
gular, 0.5–0.6 × 0.4–0.7 mm, lowest lobe triangular,
keeled, 1–1.2 × 0.9–1.1 mm; petals glabrous, white be­
coming cream-colored in vivo (cream to dark cream in
sicco); standard petal pandurate, 2–3.5 × 0.8–2.2 mm,
claw 0.5–0.8 mm, apex shallowly bilobed, notched;
wing petals 2.1–2.3 × 0.9–1.1 mm, claw 0.6–0.7 mm;
06_Novon 31_790.indd 79
Rakotonirina et al. 79
Malagasy Dalbergia (Fabaceae). IV.
keel petals 1.7–2.3 × 0.8–1.1 mm, claw 0.4–0.5 mm;
androecium glabrous, monadelphous, 2–2.3 mm; sta­
mens 8 to 10, filaments free for the upper 1/3; gynoe­
cium 1.5–2.6 mm; stipe 0.6–1.1 mm, with russet tri­
chomes; ovary 0.7–1.2 mm, pubescent, with 2 to 3
ovules; style 0.2–0.3 mm, glabrous. Fruits yellow in
vivo (dark brown in sicco), body obovate or elliptic,
(3.5–)4.6–5.1 × 1.7–2 cm when single-seeded, or
oblong-­elliptic, 6–7 × 1.8–2 cm when 2-seeded, base
acute, apex rounded, surface smooth but with a coarse
network of mostly longitudinal ribs over the seed(s),
glabrous; stipe 3–5 mm, mostly glabrescent but with
some persistent indument; style caducous. Seeds sub­
reniform, flattened, brown, smooth, ca. 7.4 × 3 mm.
Distribution and ecology. Dalbergia emirnensis oc­
curs in the northern areas of Madagascar’s Central
Highlands, within the subhumid bioclimatic zone. It
occurs across a large part of the Analamanga Region
and just crosses into the northeastern corner of the
Vakinankaratra Region; it has also been recorded in
the area where the Melaky, Bongolava, and Menabe
Regions meet, as well as in the northern part of the
Alaotra-Mangoro Region. It is found in remnants of
humid and subhumid forest and in wooded grassland,
mainly along streams and rivers, and close to forest
edges, as well as in secondary thicket in recently burned
areas. Shrubby individuals probably result from regen­
eration of numerous basal stems after cutting and/or
burning, based on observations in the area around
Ankafobe, where wildfires are very common. Dalbergia
emirnensis has been recorded on sandy and rocky fer­
ralitic soils, at 800–1500 m.
Phenology. Dalbergia emirnensis has been collected
in flower from November to February (during the be­
ginning and middle of the rainy season) and in fruit
from January to May.
Conservation status. Dalbergia emirnensis has an
estimated EOO of 34,394 km² and an estimated AOO
of 60 km² (based on 2 × 2 km grid), as calculated from
all known collections made during the past 50 years.
Older collections representing occurrences from sites
where no recent collections have been made subse­
quently are excluded since they are presumed to have
likely been extirpated, mostly from near Antananarivo
and around Lac Alaotra, where natural forest habitat
has been drastically impacted during the last five de­
cades. Two of the 15 known occurrences occur respec­
tively within two protected areas (Ankafobe Special
Reserve and Zahamena National Park). The species
is subjected to ongoing threats both inside and outside
protected areas from slash-and-burn agriculture, selec­
tive logging and wood harvesting, grazing, and frequent
4/24/23 10:33 AM
 80
human-caused wildfires, which are particularly com­
mon in the Ankafobe area. Dalbergia emirnensis exists
at seven locations with respect to the most serious
threat, slash-and-burn agriculture. Continuing decline
is projected in its AOO, habitat quality, number of lo­
cations, and number of mature individuals. Dalbergia
emirnensis is therefore preliminarily assessed as Vul­
nerable under criterion B2 [VU B2ab(ii,iii,iv,v)].
Vernacular names and uses. Hazovola (Nomban-
janahary et al. 10, 11); Voamboana (Raharimampion-
ona 417, Service Forestier 832-R-182); Sovoka (Réserves
Naturelles 6038).
The wood of Dalbergia emirnensis is used for house
construction (Nombanjanahary et al. 10, 11, Rivohari-
son 98) and as a source of firewood (Nombanjanahary
et al. 10, 11).
Etymology. The epithet “emirnensis” is derived
from the word “Emirna” (= Emyrna), an old spelling
of the name for the Imerina people and their territory,
as proclaimed by King Ralambo in the 16th century
(Ogot, 1992). This territory encompassed a large part
of the Central Highlands of Madagascar, including the
area where the type specimen was collected, and cor­
responds approximately to the present-day Antanana­
rivo Province and its four administrative regions.
Notes. Dalbergia emirnensis, as circumscribed here,
represents a morphologically and genetically coherent
species restricted to the Central Highlands. The south­
ern limit of its known distribution reaches the northern
limit of D. nemoralis. Both species are members of
the same primary clade of Malagasy Dalbergia (Super­
group II), as identified in several studies (Crameri,
2022b), but they can be distinguished from one an­
other mainly by the width of their single-seeded ma­
ture fruits (≥ 1.7 cm in D. emirnensis vs. ≤ 1.3 cm in D.
nemoralis). Moreover, D. nemoralis and D. neoperrieri
(from western Madagascar) are closely related within
the Greveana clade, whereas D. emirnensis cannot be
placed confidently within this clade (Crameri, 2020),
suggesting that it is only somewhat distantly related to
D. nemoralis and D. neoperrieri. It is possible that an­
cestral hybridization involving species in the Greveana
clade and/or another group may have contributed to
the evolution and uncertain phylogenetic placement
of D. emirnensis. Further analyses that include addi­
tional material from subpopulations in the Central
Highlands will be required to test this hypothesis and
to explore whether D. emirnensis and D. nemoralis
co-occur and whether they are reproductively isolated
from one another.
We treat Dalbergia campenonii as a synonym of
D. emirnensis. Since it is based on two syntypes, both
06_Novon 31_790.indd 80
Novon
housed at P, we designate the specimen of Campenon
s.n. bearing the barcode P00060552 as the lectotype
and the duplicate bearing the barcode P00060553 as
an isolectotype. We note also that Bosser and Rabevo­
hitra (2002) corrected the original spelling of the spe­
cies name, “D. campenoni” to D. campenonii, but they
erroneously designated the syntypes as holotype and
isotype.
Additional specimens examined. MADAGASCAR. Alaotra-­
Mangoro Reg. [Prov. Toamasina]: Andilamena, 3 Nov.
2020, ster., Nombanjanahary et al. 10, 11 (DBEV, MO, P,
TAN, ZT); same loc., 2 Nov. 2020, fl., Randrianaivo et al.
3589 (DBEV, MO, P, TAN, ZT); Manaka Est, 7 Feb. 1953, fl.,
Réserves Naturelles 6038 (P); Menaloha, 25 Feb. 1951, fl.,
Cours 4001 (P, TAN). Analamanga Reg. [Prov. Antanan-
arivo]: Ambohimanga Rova, 5 July 2019, ster., Randriana-
ivo & Bernard 3488 (DBEV, MO, P, TAN, ZT); same loc., 11
May 2021, ster., Rajaonarivelo & Ravaomanalina 14, 16, 17,
18 (DBEV, MO, TAN); Andramasina, 16 Jan. 1950, fl., Service
Forestier (Capuron) 677 (P, TEF); same loc., Dec. 1958, fl.,
Bosser 12491 (TAN [2 sheets]); same loc., 4 Jan. 1969, fl.,
Chauvet 466 (MO, P, TEF); Andranofeno Nord, 15 Dec. 2019,
fl., Wilding et al. 271 (DBEV, MO, P, TAN, ZT); Andranofeno
Sud, 15 Dec. 2019, fl., Wilding et al. 270 (DBEV, MO, P,
TAN, ZT); same loc., 2 Apr. 2022, fr., Rakotonirina 1266
(DBEV, MO, P, TAN); Angavokely, Jan. 1950, fl., Service For-
estier (Capuron) 651 (P, TEF); Ankafobe, 14 Nov. 1999, fl.,
Razafindrakoto 27 (MO, P, TAN); same loc., 24 Feb. 2005, fr.,
Lehavana et al. 285 (TAN); same loc., 5 Feb. 2014, fr., Birkin-
shaw 1990 (MO, P, TAN); same loc., 12 Oct. 2018, bud, Ran-
drianaivo 3179 (DBEV, MO, P, TAN, ZT); same loc., 26 Jan.
2020, fr., Rivoharison 98 (MO, P, TAN); Ankazobe, 10 Jan.
2013, fl., Raharimampionona & Rasolofonirina 417 (MO, P,
TAN); Tananarive (Antananarivo), rte. du Sud, fr., Bosser 9569
(MO, TAN); Manankazo, 24 May 1955, ster., Service Forestier
120-R-222 (P); Manankazo, 24 May 1955, ster., Service For-
estier 832-R-182 (P); s. loc., Campenon s.n. (P). Bongolava
Reg. [Prov. Antananarivo]: Ambaravaranala, 25 Jan. 1985,
fl., Dorr et al. 3657 (MO, P); same loc., 6 May 2009, fr.,
Razafimandimby et al. 290 (TEF). Melaky Reg. [Prov. Ma-
hajanga]: Ihorombato, 28 Nov. 2019, ster., Randrianaivo &
Andrianarivelo 3542, 3544, 3545, 3550 (DBEV, MO, P, TAN,
ZT); same loc. & date, bud, Randrianaivo & Andrianarivelo
3559, 3561, 3589 (DBEV, MO, P, TAN, ZT). Vakinankaratra
Reg. [Prov. Antananarivo]: Iharanandriana, 25 Feb. 2023,
ster., Ravololomanana & Porembski 808 (MO, P, TAN).
2. Dalbergia nemoralis Rakoton., Phillipson & Cra­
meri, sp. nov. TYPE: Madagascar. Itasy Reg. [Prov.
Antananarivo]: Fokontany d’Amboalefoka, 22
Jan. 2021, fl., N. Rakotonirina & R. Randriana-
ivo 1233 (holotype, TAN!; isotypes, DBEV!, MO-
6903000!, P [bc] P00853287!, ZT-00242665!).
Figure 2.
Diagnosis. Dalbergia nemoralis Rakoton., Phillipson &
Crameri can be distinguished from the closely related D. ne-
operrieri Bosser & R. Rabev. by its (narrowly) elliptic, elliptic-­
oblong, or obovate leaflets (vs. lanceolate or oblong in D.
neoperrieri) with an apiculate to rounded or emarginate (vs.
acuminate) apex, and by its narrower (0.7–1.3 cm vs. 2–2.8
cm wide) and frequently 2- or 3-seeded (vs. mostly 1- or occa­
sionally 2-seeded) fruits.
4/24/23 10:33 AM
 Volume 31 Rakotonirina et al. 81
2023 Malagasy Dalbergia (Fabaceae). IV.

Figure 2. Dalbergia nemoralis Rakoton., Phillipson & Crameri subsp. nemoralis. —A. Flowering branch. —B. Fruiting
branch. —C. Leaflet (left, upper surface; right, lower surface). —D. Flower. —E. Floral dissection, from left to right: adaxial
surfaces of standard petal, wing petal, keel petal; androecium (opened to show adaxial surface); gynoecium. —F. Calyx (outer
surface, split open and flattened). (A, C–F, N. Rakotonirina & R. Randrianaivo 1233; B, N. Rakotonirina & R. Randrianaivo
1232). Drawn by Roger Lala Andriamiarisoa.

06_Novon 31_790.indd 81 4/24/23 10:33 AM

 82
Trees to ca. 20 m tall, 40 cm diam., sometimes multi-­
stemmed shrubs. Branches glabrous, glabrescent, ro­
bust, yellow-green in vivo (brown with pale gray pattern
in sicco), with scattered, white lenticels, bark brown
with a whitish gray pattern. Leaves alternate, 7–13.4
cm, with (5 to )6 to 11 alternate leaflets; petiole and
rachis yellow-green or yellowish in vivo (brown or dark
brown in sicco), puberulent or velutinous along the
rachis and the petiole; petiole 0.5–1.7 cm; stipules tri­
angular, 4–7.1 × 0.9–3 mm, caducous; leaflets (11–)
16–64(–71) × 6–20 mm, the proximal ones sometimes
smaller than the rest, the distal leaflet slightly larger,
the others relatively uniform; petiolules 1.3–5(–6) mm,
yellow-green in vivo (light to dark brown in sicco), pu­
bescent, velutinous; lamina obovate to elliptic (some­
times broadly so), narrowly elliptic, or elliptic-oblong,
base cuneate, margins slightly recurved, apex rounded,
emarginate, or apiculate to shortly acuminate, venation
simple brochidodromous, with 7 to 11 principal sec­
ondary veins per side; upper and lower surfaces green
to yellow-green in vivo (brown in sicco), upper surface
matte or glossy, puberulous or glabrous, lower surface
matte, puberulent or with a glaucous bloom and fine
appressed or velutinous indument, especially along the
midrib, higher-order veins forming a loose network,
the midrib forming a channel above, prominent below.
Inflorescences terminal, a panicle composed of a se­
ries of lax, regular, well- or sparsely branched corym­
biform paniculate units, 3.8–18 cm, decreasing in size
toward the apex, the basal units subtended by well-­
developed, pinnate, leaflike bracts, the apical units
with reduced, caducous, scalelike bracts, distal por­
tions curved, subspicate, axes dark yellow in vivo (light
golden brown in sicco), axes yellow-green in vivo (brown
in sicco), pubescent; peduncle 0.4–0.5 cm. Flowers
subtended by a triangular, puberulous, scalelike bract,
0.4–0.7 × 0.3–0.4 mm; pedicel 0.1–0.8 mm, pubes­
cent (pedicels of apical flowers shorter than those of
basal flowers); bracteoles persistent, ovate with rounded
apex, partially enclosing the base of the calyx, 0.4–
0.8 × 0.3–0.6 mm, puberulous, yellow-green in vivo
(orange-­brown or light brown in sicco); calyx yellow-­
green in vivo (dark yellow-colored in sicco), puberu­
lent, not persisting on fruits, 1.8–3.6 mm, fused in
lower 2/3, the 2 upper lobes rounded, and fused to form
a single unit, 0.3–1.1 × 1–2 mm, the 2 lateral lobes
triangular, 0.4–0.8 × 0.4–0.7 mm, the lowest lobe tri­
angular, 0.9–1.7 × 0.4–0.9 mm; petals glabrous, pu­
berulent, white to cream (fide Du Puy M415), fading
to yellow (fide Du Puy M404) in vivo (dark yellow in
sicco); standard petal violin-shaped, 3–4.4 × 2.2–3 mm,
claw 0.5–1 mm, apex shallowly bilobed, notched; wing
petals 2.7–4.2 × 1.1–1.3 mm, claw 0.4–1.2 mm; keel
petals (2–)2.2–3.7 × 1–1.6 mm; androecium glabrous,
monadelphous, 2.1–3.7 mm; stamens 10, filaments free
06_Novon 31_790.indd 82
Novon
for the upper 1/3 or 1/2; gynoecium (2.2–)2.5–4 mm;
stipe 0.7–2 mm, pubescent (subspecies nemoralis) with
russet trichomes (subspecies decaryi); ovary 0.9–1.8
(–2) mm, puberulous (subspecies nemoralis) or pubes­
cent (subspecies decaryi), with 2 or 3 ovules; style
0.3–0.7 mm, glabrous. Fruits (immature) yellow-green
in vivo (brown in sicco), body elliptic to suborbicular,
1.5–2.5 × 0.7–1.3 cm when 1-seeded, elliptic-oblong,
2.2–3.6(–4.6) × 0.6–1.1 cm when 2-seeded or 2.9–3.5
(5.2) × 0.6–0.7(–0.9) cm when 3-seeded (subspecies
decaryi), base acute, apex rounded, mucronate, surface
smooth but with a coarse network of mostly longitudi­
nal ribs over the seed(s), extending almost to the edge,
glabrous; stipe 3–6 mm, mostly glabrescent but with
some persistent indument or with sparse indument at
the base; style caducous. Seeds reniform, flattened,
brown, 2.5–5 × 1.7–2.5 mm.
Conservation status. Dalbergia nemoralis has an
estimated EOO of 130,804 km² and an estimated AOO
of 132 km² (based on 2 × 2 km grid), as calculated from
all known collections made during the past 50 years.
Older collections representing occurrences from sites
where no recent collections have been made subse­
quently are excluded since they are presumed to have
likely been extirpated, mostly from Antsirabe, Am­
pandrandava, Antanimora, Bekily, and some around
Atsimo-­Andrefana, where natural forest habitat has
been impacted during this period. Seventeen of the 55
known collection sites occur within seven protected
areas (Analavelona, Andohahela, Angavo, Behara-­
Tranomaro, Complexe Vohidava-Betsimalaho, Itremo,
and Zombitsy). The species is subjected to ongoing
threats both inside and outside protected areas from
selective logging, artisanal mining, slash-and-burn ag­
riculture, fire, and grazing. While continuing decline
is projected in its AOO, habitat quality, number of lo­
cations, and number of mature individuals, because
D. nemoralis exists at 15 locations with respect to the
most serious threat, wood exploitation (pers. obs.), it is
preliminarily assessed as Least Concern.
Etymology. The epithet “nemoralis” is the Latin
word meaning “from woods or groves,” alluding to the
habitat of the species.
Notes. Two eco-geographically distinct subspecies
are recognized within Dalbergia nemoralis: one occur­
ring in the Central and Southern Highlands of Mada­
gascar at elevations mostly above 600 m, and the other
is found predominantly in the southern hills and low­
lands, generally at elevations below 600 m. However,
these two entities come into contact in a zone situated
approximately where the lower-elevation southern part
of Madagascar rises to the Southern Highlands (Fig. 1).
4/24/23 10:33 AM
 Volume 31
2023
Gene flow between the two subspecies likely occurs in
this zone, where assignment of collections to subspe­
cies may be difficult or impossible. It is also interest­
ing to note that the morphological differences between
D. nemoralis and the closely related D. neoperrieri
appear to be most pronounced when material of D.
nemoralis from the extreme south of Madagascar is
compared to typical collections of D. neoperrieri from
the Boeny Region, while differences in leaflet size and
shape (but not in fruit characters) appear less pro­
nounced when comparing material of the two species
from the Atsimo-Andrefana Region in the southwest­
ern part of the island. Additional analyses, including
samples from the Atsimo-Andrefana and Menabe Re­
gions, will be required to assess whether D. nemoralis
and D. neoperrieri co-occur, and if so, whether they are
reproductively isolated.
Specimens not assigned to subspecies. MADAGASCAR.
Androy Reg. [Prov. Toliara]: Ambia, 24–25 Jan. 1955,
fl., Service Forestier (Capuron) 11695 (P, TEF); Ampandran­
dava, 31 Dec. 1942, fl., Seyrig 447 (P [2 sheets]); Beraketa
toward Ampandrandava, 6 Mar. 2008, fr., SNGF 1942 (K,
TEF); Beraketa, 18 Mar. 1997, fr., Du Puy et al. M994 (K,
MO, P). Atsimo-Andrefana Reg. [Prov. Toliara]: Anala­
velona, 27 Jan. 2016, fl., Bernard 2454 (MO, P); Tongobory,
25 Jan. 2014, ster., Razafimamonjy 8 (P); Zombitsy, 13 Feb.
1990, fl., Labat et al. 2115 (K, MO, P).
2a. Dalbergia nemoralis subsp. nemoralis.
Trees to ca. 17 m tall, 24 cm diam., or shrubs.
Branches glabrescent, yellow-green in vivo (brown in
sicco). Leaves 7–12 cm, with (5 to)7 to 11 leaflets; pet­
iole and rachis yellow-green in vivo (brown in sicco),
velutinous along the rachis and petiole; petiole 1.1–
1.7 cm; stipules triangular, 4–7.1 × 0.9–3 mm; leaflets
(11–)16–52(–60) × 6–14(–16) mm; petiolules 1.3–5 mm,
yellow-green in vivo (light brown in sicco), velutinous;
lamina narrowly elliptic or elliptic-oblong, margins
slightly recurved, apex apiculate to shortly acuminate,
venation with 9 to 11 principal secondary veins per
side; upper and lower surfaces green to yellow-­green
in vivo (brown in sicco), upper surface slightly glossy,
puberulous, lower surface with gold-colored, velutinous
indument, especially along the midrib. Inflorescences
composed of sparsely branched corymbiform, panicu­
late units, 3.8–6.7 cm, axes yellow-green in vivo (brown
in sicco), pubescent; peduncle 0.4–0.5 cm. Flowers with
scalelike bract 0.5–0.7 × 0.3–0.4 mm; pedicel 0.1–0.6
mm, bracteoles 0.5–0.8 × 0.4–0.6 mm, puberulous,
orange-brown in sicco; calyx 2.6–3.6 mm, the 2 upper
lobes in a unit 0.3–1.1 × 1.6–2 mm, apex rounded, the
2 lateral lobes 0.4–0.8 × 0.4–0.7 mm, lowest lobe 0.9–
1.7 × 0.4–0.9 mm; petals glabrous, white in vivo (dark
yellow in sicco); standard petal (3–)3.5–4.4 × 2.2–3
mm, claw 0.5–1 mm; wing petals 2.7–4.2 × 1–1.3 mm,
claw 0.4–1.2 mm; keel petals (2–)2.5–3.7 × 1–1.6 mm,
06_Novon 31_790.indd 83
Rakotonirina et al. 83
Malagasy Dalbergia (Fabaceae). IV.
claw 0.8–1.2 mm; androecium 3.2–3.7 mm; filaments
of stamens free for the upper 1/2; gynoecium (2.2–)
2.5–3.7 mm; stipe with russet trichomes, 0.7–2 mm;
ovary puberulous, 0.9–1.5(–2) mm, with 2 ovules; style
0.3–0.7 mm. Fruits (immature) yellow-green in vivo
(brown in sicco), body elliptic, 2–2.5 × 0.9–1.3 cm
when single-seeded, elliptic to oblong, ca. 3.6 × 1.1
cm when 2-seeded; stipe 4–6 mm, with sparse indu­
ment; style caducous. Seeds not seen.
Distribution and ecology. Dalbergia nemoralis
subsp. nemoralis occurs in the Central and Southern
Highlands, within the subhumid and subarid biocli­
matic zones. It has been recorded in Itasy, across Vak­
inankaratra and Amoron’i Mania, to Matsiatra Ambony
Regions, and also at rather scattered localities in the
north of Anosy Region to the northern limit of Androy
Region (Fig. 1). The subspecies has been recorded from
woodlands, wooded grassland-bushland mosaics, grass­
lands and other open vegetation, and on stream banks.
It occurs mostly on lateritic soils formed over igneous
or metamorphic rocks, at (350–)600–1600 m.
Phenology. Dalbergia nemoralis subsp. nemoralis
has been collected in flower from December to March
(at the middle of the rainy season), and in fruit mainly
from January to March, and rarely in August.
Conservation status. Dalbergia nemoralis subsp.
nemoralis has an estimated EOO of 18,170.4 km² and
an estimated AOO of 28 km² (based on 2 × 2 km grid),
as calculated from all known collections made during
the past 50 years. Older collections representing oc­
currences from sites where no recent collections have
been made subsequently are excluded since they are
presumed to have likely been extirpated, mostly from
Antsirabe and Ampandrandava, where natural forest
habitat has been impacted during this period. Two of
the 11 known collection sites occur within two protected
areas (Complexe Vohidava-Betsimalaho and Itremo).
The subspecies is subjected to ongoing threats outside
and within the two protected areas mentioned above,
where effective protection is known to be inadequate,
from selective logging, artisanal mining, slash-and-burn
agriculture, and grazing. Dalbergia nemoralis subsp.
nemoralis exists at four locations with respect to the
most serious threat, wood exploitation (pers. obs.). Con­
tinuing decline is projected in its AOO, habitat quality,
number of locations, and number of mature individu­
als. Dalbergia nemoralis subsp. nemoralis is therefore
preliminarily assessed as Endangered under criterion
B2 [EN B2ab(ii,iii,iv,v)].
Vernacular names and uses. Fitoeram-boanosy
(Rako­tonirina & Randrianaivo 1233), Magnary (Rave-
4/24/23 10:33 AM
 84
lonarivo et al. 4575), Manary (Service Forestier 9565),
Tanamolahy (Réserves Naturelles 720).
The dark-brown wood of Dalbergia nemoralis subsp.
nemoralis is hard and sufficiently long lasting to be re­
garded as highly prized for construction, and has been
recorded as used for window frames (Rakotonirina &
Randrianaivo 1233). This taxon has also been noted as
toxic, without precise indication of which parts sup­
posedly exhibit toxic properties and to whom (Réserves
Naturelles 720).
Notes. Some material of Dalbergia nemoralis subsp.
nemoralis was included in the nominal variety of D.
emirnensis by Bosser and Rabevohitra (1996, 2002)
but, as indicated above, is better treated as distinct
from that species owing to its eco-geographic separa­
tion from, but closer genetic and morphological rela­
tionship with, material from southern Madagascar now
included in D. nemoralis subsp. decaryi. Specimens
of D. nemoralis subsp. nemoralis can be distinguished
from material of D. nemoralis subsp. decaryi by their
narrowly elliptic or elliptic-oblong (vs. obovate to el­
liptic) leaflets, whose lower surface has gold-colored,
velutinous indument (vs. white glaucous bloom, fine
appressed indument), and their frequently shorter, more
compact inflorescences (often less than 6 cm vs. gener­
ally more than 12 cm in length). Dalbergia nemoralis
subsp. nemoralis is distributed in woodland and wooded
grassland (vs. spiny thicket for subsp. decaryi).
Additional specimens of Dalbergia nemoralis subsp. nemor-
alis examined. MADAGASCAR. Amoron’i Mania Reg.
[Prov. Fianarantsoa]: Itremo, Feb. 1919, fl., Perrier de la
Bâthie 12462 (P [2 sheets]). Androy Reg. [Prov. Toliara]:
Ampandrandava, fr., s.d., Seyrig 23 (P); Beraketa, Mar. 1934,
fl., Humbert 14178 (P); Manavy, 11 Dec. 1953, fl., Service
Forestier 9565 (P, TEF). Anosy Reg. [Prov. Toliara]:
Anadabolava-Betsimalaho, 8 Feb. 2013, fl., Ravelonarivo et
al. 4575 (MO, P, TAN, UPS); Betroka, 17 Jan. 1954, ster.,
Service Forestier 17-R-87 (P); same loc., 2 Mar. 1966, fr., Ser-
vice Forestier 26036 (P, TEF); Vohidava-Betsimalaho, 20 Jan.
2022, Antilahimena & Masinkery 9886 (MO, P, TAN); same
loc., 20 Jan. 2022, Antilahimena & Masinkery 9991 (MO, P,
TAN). Itasy Reg. [Prov. Antananarivo]: Miarinarivo, 22
Jan. 2021, fl., fr., Rakotonirina & Randrianaivo 1232 (DBEV,
MO, P, TAN, ZT); same loc., 2 Dec. 2019, fl., ster., Randri-
anaivo & Andrianarivelo 3568, 3569 (DBEV, MO, P, TAN,
ZT). Matsiatra Ambony Reg. [Prov. Fianarantsoa]: An­
jamana, 23 Aug. 2009, fr., Andriamahay & Rakotoarisoa
2323 (K, MO, P, TAN, TEF); Iakanga, 12 Dec. 1954, fl., Ré-
serves Naturelles 720 (TEF). Vakinankaratra Reg. [Prov.
Antananarivo]: Antsirabe, Dec. 1913, fl., Perrier de la Bâthie
4271 (P).
2b. Dalbergia nemoralis subsp. decaryi (Bosser
& R. Rabev.) Rakoton., N. Wilding, Phillipson &
Crameri, comb. et stat. nov. Basionym: Dalbergia
emirnensis var. decaryi Bosser & R. Rabev., Bull.
Mus. Natl. Hist. Nat., B., Adansonia, sér. 4, 18:
06_Novon 31_790.indd 84
Novon
209. 1996. TYPE: Madagascar. Anosy Reg. [Prov.
Toliara]: sables blancs au N de Beloha, 13 Jan.
1962, fl., Service Forestier (Capuron) 20677 (ho­
lotype, P [bc] P00060671!; isotypes, K [bc]
K000379218!, MO-5086113!, P [bc] P00060672!,
TEF [bc] TEF000129!).
Trees to ca. 20 m tall, 40 cm diam., sometimes multi-
stemmed shrubs. Branches glabrous, yellow-­green in
vivo (pale gray in sicco), bark whitish gray. Leaves
7–13.4 cm, with 6 to 10 leaflets; petiole and rachis yel­
lowish in vivo (dark brown in sicco), puberulent along
the rachis and petiole; petiole 0.5–1.1 cm; stipules not
seen, leaving a visible scar; leaflets (12–)25–64(–71) ×
(7–)8–20 mm; petiolules 3–4(–6) mm, yellow-green in
vivo (dark brown in sicco), pubescent; lamina obovate
to elliptic (sometimes broadly so), margins slightly rev­
olute, apex rounded or emarginate, venation with 7 to
10 principal secondary veins per side; upper and lower
surfaces bright midgreen in vivo (gray-green to light
brown in sicco), upper surface glabrous, lower surface
with white glaucous bloom and fine appressed indu­
ment. Inflorescences composed of a series of lax, regu­
lar, well-branched corymbiform, paniculate units, 11.5–
18 cm, axes dark yellow in vivo (light golden brown in
sicco), pubescent; peduncle 0.8–1.7 cm. Flowers with
scalelike bract 0.4–0.5 × 0.3–0.4 mm; pedicel 0.6–0.8
mm, bracteoles 0.4–0.6 × 0.3–0.4 mm, puberulent,
light brown in sicco; calyx 1.8–2.5 mm, the 2 upper
lobes in a unit 0.9–1 × 1–1.6 mm, apex retuse, the 2
lateral lobes 0.7–0.8 × 0.5–0.7 mm, lowest lobe 0.9–1
× 0.6–0.8 mm; petals puberulent, white to cream (fide
Du Puy M415), fading to yellow (fide Du Puy M404) in
vivo; standard petal 3–3.2 × 2.8–2.9 mm, claw 0.5–0.8
mm; wing petals 2.8–3 × 1–1.1 mm, claw 0.7–0.8 mm;
keel petals 2.2–2.5 × 1.2–1.3 mm, claw 0.8–1.1 mm;
androecium 2.1–2.5 mm; filaments of stamens free for
the upper 1/3; gynoecium 2.9–4 mm; stipe pubescent,
1–1.7 mm; ovary pubescent, 1.5–1.8 mm, with 3 ovules;
style 0.4–0.5 mm. Fruits yellow-green in vivo (brown
in sicco), the body elliptic to suborbicular, 1.5–2.1 ×
0.7–0.8 cm when single-seeded, elliptic-oblong, 2.2–
3.4(–4.6) × 0.6–0.9 cm when 2-seeded, or 2.9–3.5(5.2)
× 0.6–0.7(–0.9) cm when 3-seeded; stipe 3–4 mm,
mostly glabrescent but with some persistent indument
at the base; style caducous. Seeds reniform, flattened,
brown, 2.5–5 × 1.7–2.5 mm.
Distribution and ecology. Dalbergia nemoralis
subsp. decaryi occurs in southern Madagascar, within
the subarid bioclimatic zone. It is most frequent in the
southern part of Anosy Region and across much of An­
droy Region, and also occurs at rather scattered local­
ities in the southern part of Atsimo-Andrefana Region.
The subspecies is found in intact or degraded spiny
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 Volume 31
2023
thicket, persisting in grazed areas around villages, scrub­
land, and often in association with Alluaudia procera
(Drake) Drake (Didiereaceae). Dalbergia nemoralis
subsp. decaryi has been recorded on fersiallitic, lime­
stone, or sandy soils formed over southern Madagas­
car’s tertiary limestone or unconsolidated sands, at
30–600(–900) m.
Phenology. Material of Dalbergia nemoralis subsp.
decaryi has been collected in flower from December
to February (during the beginning and middle of the
rainy season), and in fruit from February to April, and
rarely in August.
Conservation status. Dalbergia nemoralis subsp.
decaryi has an estimated EOO of 28,660 km² and an
estimated minimum AOO of 88 km² (based on a 2 ×
2 km grid), as calculated from all known collections
made during the past 50 years. Older collections rep­
resenting occurrences from sites where no recent col­
lections have been made subsequently are excluded
since they are presumed to have likely been extirpated,
mostly from Antanimora, Bekily, and some around
Atsimo-­Andrefana, where natural forest habitat has
been impacted during this period. Twelve of the 38
known collection sites occur within four protected areas
(Andohahela, Angavo, Behara-Tranomaro, and Com­
plexe Vohidava-Betsimalaho). It is subjected to on­
going threats both inside and outside protected areas
from slash-and-burn agriculture, illegal logging, artis­
anal mining, grazing, and fire, and the subspecies ex­
ists at eight locations with respect to the most serious
plausible threat, slash-and-burn agriculture. Continu­
ing decline is projected in its AOO, habitat quality, the
number of locations, and the number of mature individ­
uals. Dalbergia nemoralis subsp. decaryi is therefore
preliminarily assessed as Vulnerable under Criterion
B2 [VU B2ab(ii,iii,iv,v)].
Vernacular names and uses. Magnary (Bernard et
al. 2879, 2880); Manary (Andriamiarisoa 2649 et seq.,
Decary 4366, Humbert 1291, 12476, Ramamonjiarisoa
22, 26, Réserves Naturelles 6197, 10775, Service For-
estier 922, 3427, 9397, 9492, 25591, 25596, 73-R-189,
13-R-316); Manarinalafia (Réserves Naturelles 8198);
Manarim-bato (Service Forestier 27-R-316); Manarint­
saka (Service Forestier 15532); Manary belity (Service
Forestier 9253); Manary toloho (Réserves Naturelles
5023, 6423); Manary vatany (Service Forestier 2833);
Mangary (Du Puy M404, M415).
The dark red-brown wood of Dalbergia nemoralis
subsp. decaryi is hard and long lasting; it is used in
construction (Du Puy M404, M415, Service Forestier
25596) and for fences (Service Forestier 27-R-316), as
06_Novon 31_790.indd 85
Rakotonirina et al. 85
Malagasy Dalbergia (Fabaceae). IV.
well as for boards and hand tools (Service Forestier 9492,
13-R-316).
Notes. Material of Dalbergia nemoralis subsp. de-
caryi was assigned to D. emirnensis var. decaryi by
Bosser and Rabevohitra (1996, 2002) but, as shown
above, is better treated as a subspecies of D. nemoralis
(see notes above under D. nemoralis subsp. nemoralis).
Additional specimens of Dalbergia nemoralis subsp. decaryi
examined. MADAGASCAR. Androy Reg. [Prov. Toli-
ara]: Ambovombe, 11 Feb. 1925, fl., Decary 3521 (P, TEF);
same loc., 2 Feb. 1931, fl., Decary 8448 (MO, P); Mitsoriaka,
19 July 2021, ster., Andriamiarisoa & Bernard 2659, 2660,
2661, 2662, 2663, 2665, 2666, 2667, 2668 (DBEV, MO, P,
TAN, ZT); Ampanihy-Betioky, 21 Apr. 2004, fr., SNGF 749
(K); Ampira, 10 Mar. 1955, ster., Service Forestier 73-R-189
(P); Androy, 24 Mar. 1951, fr., Service Forestier 3427 (P, TEF);
Angavo, 20 July 1926, ster., Decary 4366 (P); same loc., 25
Jan. 1955, ster., Service Forestier 13-R-316 (P); Antanimora
(Atsimo), 23 Nov. 1977, bud, Ramamonjiarisoa 26 (P); entre
Antanimora et Ambovombe, 7 Dec. 1961, fl., Service Forestier
(Capuron) 20434 (K, MO, NY, P, PRE, TEF, WAG); entre An­
tanimora et la vallée de l’Ikonda, 24 Jan. 1963, fl., Service
Forestier 22495 (P, TEF); Bafana, 23 Nov. 1977, ster., Rama-
monjiarisoa 22 (P); Bekily, 16 Feb. 1954, fr., Service Forestier
9253 (K [2 sheets], MO, P, TEF, WAG); Belindo-Ambovombe,
17 Jan. 1955, fl., Service Forestier 15532 (MO, P, TEF); Be­
loha, 27 Jan. 1990, fl., Du Puy M415 (K [2 sheets], MO, P,
TAN); same loc., 13 Jan. 1962, fl., Service Forestier 20677 (K,
MO, P, TEF); Bereny, 24 Jan. 1966, fl., Service Forestier
25596 (K, MO, P, PRE, TEF, WAG); Tanamolora, 10 Dec.
1965, fl., Service Forestier 25591 (K, P, TEF); Vohipary, 18
Feb. 1955, ster., Service Forestier 27-R-316 (P); same loc., 19
Aug. 2001, fr., Rakotondranony 205 (K). Anosy Reg. [Prov.
Toliara]: Amboasary, 22 Nov. 1977, ster., Ramamonjiarisoa
25 (P); Ampisopiso-Mahazoarivo, 12 July 2021, ster., Bernard
& Andriamiarisoa 2879, 2880 (DBEV, MO, P, TAN, ZT);
Anadabolava, Dec. 1933, fl., Humbert 12476 (G, K, MO, P,
TEF, WAG); Andohahela, 17 Mar. 1994, fr., Randriambo-
lolona et al. 90 (G, K, MO, P, PRE, TAN, UPS, WAG); same
loc., 28 Jan. 1990, fl., McPherson 14933 (K, MO, P, TAN,
TEF); same loc., 22 Mar. 2010, fr., Ratovoson 1583 (K, MO, P,
TAN); same loc., 12 Jan. 1954, fl., Réserves Naturelles 6197
(K, MO, P, TEF, WAG); same loc., 24 Feb. 1960, fr., Réserves
Naturelles 10775 (P); Behara, 10 July 2021, ster., Andriamia-
risoa & Bernard 2649, 2650, 2651, 2652, 2653 (DBEV, MO,
P, TAN, ZT); same loc., 24 Jan. 1954, fl., Réserves Naturelles
6423 (B, BR, G, K, MO, NY, P, PRE, TEF, WAG); same loc.,
25 Jan. 1953, fl., Réserves Naturelles 5023 (K, MO, P); Bev­
ilany, 7 Dec. 1968, fl., Service Forestier 28559 (P, TEF); Iso­
mono, Dec. 1933, fl., Humbert 12917 (BR, G, K, MO, NY, P,
PRE, TEF); Morahariva, Dec. 1933, fl., Humbert 13203 (P);
Ranopiso, 24 Dec. 1956, fl., Réserves Naturelles 8198 (P,
TEF); Tsiombe, 24 Jan. 2001, fl., Phillipson 5267 (MO, P);
Vohimainty, 24 Jan. 1990, fl., Du Puy M404 (K [2 sheets],
P, TAN). Atsimo-Andrefana Reg. [Prov. Toliara]: An­
dranohinaly, 20 Mar. 1954, fr., Service Forestier 9397 (MO, P,
TEF); Androka, 7 Feb. 1990, fr., Labat et al. 2093 (K, MO,
P, TAN); Hazoroa, 3 Mar. 1951, fl., Service Forestier 2833 (P,
TAN, TEF); Rerihitiny, 26 Mar. 1954, fr., Service Forestier
9322 (K, P, TEF); same loc., 27 Mar. 1954, fr., Service For-
estier 9492 (G, P, PRE, TEF); Sakaraha, 3 Mar. 1951, fr., Ser-
vice Forestier 3379 (P, TEF); same loc., 12 Dec. 1962, fl.,
Service Forestier 22269 (K, P, TEF).
4/24/23 10:33 AM
 86
Acknowledgments. We thank the curators of the
herbaria at the Conservatoire et Jardin botaniques (G)
in Geneva, the Royal Botanic Gardens, Kew (K), and at
FOFIFA (TEF) in Ambatobe and the Parc Botanique et
Zoologique de Tsimbazaza (TAN) in Antananarivo, for
granting access to the specimens housed in their re­
spective collections. We thank several people without
whose help this work would not have been possible:
Claude-Anne Gauthier (Institut de Recherche pour le
Développement and Muséum National d’Histoire Na­
turelle [MNHN] Representative in Madagascar) and
Maïté Delmas (MNHN) for facilitating the “Bourse du
Gouvernement Français” provided to the first author,
and Roger Lala Andriamiarisoa (Missouri Botanical
Garden, Madagascar) for preparing the line drawings
of Dalbergia nemoralis. We are grateful to two anony­
mous reviewers and to the editor for helpful comments
that improved the final version of the manuscript. We
acknowledge funding from the Service de Coopération
et d’Action Culturelle (SCAC) of the French Embassy
in Madagascar (Bourse du Gouvernement Français
2020, N° Dossier Campus France n° 971938 E), the
Fondation Franklinia, the National Geographic Society
(EC-63842 R-20), and the Madagascar Precious
Woods Consortium as part of the G3D (Gestion Dura­
ble des bois précieux Dalbergia et Diospyros de Mada­
gascar) Project funded by the Délégation de l’Union
Européenne à Madagascar (DEUM). S. C.’s contribu­
tion was supported by the Swiss Federal Food Safety
and Veterinary Office (ARAMIS # 3.20.01) and a grant
from the Rübel Foundation.
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