Botanical Journal of the Linnean Society, 2006, 152, 331–386. With 20 figures
© 2006 The Linnean Society of London, Botanical Journal of the Linnean Society, 2006, 152, 331–386 331
Blackwell Publishing LtdOxford, UKBOJBotanical Journal of the Linnean Society0024-4074The Linnean Society of London, 2006? 2006
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Original Article
REVISION OF HYMENODICTYEAE
S. G. RAZAFIMANDIMBISON and B. BREMER
*Corresponding author.
E-mail: sylvain.razafimandimbison@bergianska.se
Taxonomic revision of the tribe Hymenodictyeae
(Rubiaceae, Cinchonoideae)
SYLVAIN G. RAZAFIMANDIMBISON
1,2
* and BIRGITTA BREMER
1,2
1
The Bergius Foundation at the Royal Swedish Academy of Sciences, PO Box 50017, SE-10405,
Stockholm, Sweden
2
Department of Botany, Stockholm University, SE-10691, Stockholm, Sweden
Received October 2005; accepted for publication April 2006
The palaeotropical tribe Hymenodictyeae Razafim. & B. Bremer, belonging to the otherwise predominantly Neotro-
pical subfamily Cinchonoideae s.s. (coffee family or Rubiaceae), is revised here. The tribe as presently circumscribed
contains two genera, Hymenodictyon Wall. and Paracorynanthe Capuron ex J.-F. Leroy, and is distinct from the other
Cinchonoideae tribes with capsular fruits in having stipules bearing large, deciduous colleters on the margins, val-
vate corolla aestivation, and lenticellate capsular fruits that contain elongate, bilaterally flattened, and accrescent
placentae. We recognize 22 Hymenodictyon species, including four new species (H. antakaranensis sp. nov.,
H. epiphyticum sp. nov., H. madagascaricum sp. nov., and H. tsingy sp. nov.), and the two described species
of Paracorynanthe. Hymenodictyon is distinguished from its sister genus, Paracorynanthe, by simple or compound
spicate, racemose or thyrsoid inflorescences and corolla lobes without any appendages, rather than compound umbel-
liform inflorescences and corolla lobes prolonged by ciliate appendages. A full taxonomic treatment, keys, and dis-
tribution maps of all recognized Hymenodictyon and Paracorynanthe species are provided. Five lectotypes and one
neotype are designated. Finally, six Hymenodictyon species are illustrated for the first time. © 2006 The Linnean
Society of London, Botanical Journal of the Linnean Society, 2006, 152, 331–386.
ADDITIONAL KEYWORDS: Africa – Asia – Hymenodictyon – Madagascar – Paracorynanthe – systematics –
taxonomy.
INTRODUCTION
The tribe Hymenodictyoneae Razafim. & B. Bremer,
correctly spelled by Bridson & Verdcourt (2003: 385 &
423) as Hymenodictyeae, was described by Razafiman-
dimbison & Bremer (2001) to accommodate the two
palaeotropical genera, Hymenodictyon Wall. and Para-
corynanthe Capuron. Andersson & Persson (1991)
placed these genera in the tribe Coptosapelteae, which
was demonstrated by Razafimandimbison & Bremer
(2001) to be highly polyphyletic. Hymenodictyeae
belongs to the subfamily Cinchonoideae s.s. (Bremer
et al., 1999) of the coffee family (Rubiaceae) and has
been shown to be sister to the mostly palaeotropical
tribe Naucleeae s.l. (Razafimandimbison & Bremer,
2002). In Andersson & Antonelli (2005) the Hymeno-
dictyeae–Naucleeae clade was resolved as sister to a
predominantly Neotropical clade, which contains
Chiococceae sensu Bremer (1992), Cinchoneae sensu
Andersson & Antonelli (2005), Guettardeae, Hame-
lieae, Hillieae, Isertieae sensu Bremer & Thulin
(1998), and Rondeletieae. In Razafimandimbison &
Bremer (2001: 526), Paracorynanthe, represented by
P. antankarana Capuron ex J.-F. Leroy, and
Hymenodictyon, represented by four Hymenodictyon
species [H. decaryi Homolle, H. flaccidum Wall.,
H. floribundum (Hochst. & Steud.) B.L. Rob., and
H. parvifolium Oliv.] formed a strongly supported
monophyletic group (bootstrap support value = 100%).
The phylogenetic relationships within this Hymenod-
ictyon–Paracorynanthe clade (= Hymenodictyeae)
were unresolved. On the other hand, an unpublished
phylogenetic analysis based on external transcribed
spacers sequence data (S. Razafimandimbison and B.