2002 - Relationships and Generic Circumscriptions in The Psychotria Complex (Rubiaceae

Relationships and Generic Circumscriptions in the Psychotria Complex (Rubiaceae,
Psychotrieae)
Author(s): Lennart Andersson
Source: Systematics and Geography of Plants , 2002, Vol. 72, No. 1/2 (2002), pp. 167-202
Published by: Botanic Garden Meise
Stable URL: https://www.jstor.org/stable/3668767
REFERENCES
Linked references are available on JSTOR for this article:
https://www.jstor.org/stable/3668767?seq=1&cid=pdf-
reference#references_tab_contents
You may need to log in to JSTOR to access the linked references.
JSTOR is a not-for-profit service that helps scholars, researchers, and students discover, use, and build upon a wide
range of content in a trusted digital archive. We use information technology and tools to increase productivity and
facilitate new forms of scholarship. For more information about JSTOR, please contact support@jstor.org.
Your use of the JSTOR archive indicates your acceptance of the Terms & Conditions of Use, available at
https://about.jstor.org/terms
is collaborating with JSTOR to digitize, preserve and extend access to Systematics and
Geography of Plants
This content downloaded from

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
All use subject to https://about.jstor.org/terms
Syst. Geogr Pl. 72: 167-202 (2002)
"Ik"
Relationships and generic circumscriptions in the
Psychotria complex (Rubiaceae, Psychotrieae)
Lennart Andersson
Botanical Institute, Goteborg University,

P.O. Box 461, SE-405 30 Goteborg, Sweden
Abstract. - Relationships among 111 species of the Psychotria complex (Rubiaceae tribe
Psychotrieae) were estimated by cladistic analysis of nucleotide sequence variation in the rpsl6
intron (cpDNA). The Psychotria complex is characterised morphologically by caducous stipules
and (usually) testa with a reddish, ethanol soluble pigment. It was found to be comprised of two
major clades, one of which is Psychotria in a strict sense. Psychotria s. str. is characterised by usu-
ally having pyrenes without preformed germination slits, a plane or shallowly furrowed adaxial
surface and usually numerous distinct ridges on the abaxial side. Most species occur in Africa and
the Neotropics, but the genus extends eastwards to the Philippines and Australia. The second sub-
clade comprises species from areas around the Indian Ocean and in the Pacific. Members of this
subclade mostly differ morphologically from Psychotria s. str by pyrenes with distinct marginal
preformed germination slits. This subclade includes the tuberiferous, myrmecophile epiphytes,
which, it is suggested, comprise a single genus, Hydnophytum. It also includes the genera
Amaracarpus, Calycosia, Dolianthus, and Straussia. These genera are not supported as mono-
phyletic by sequence data, but are tentatively recognized based on morphological evidence.
Calycosia should be given a broader circumscription than presently and may be characterised by
entire endosperm and pollen grains with usually thin exine and small lumina. The name Straussia
has traditionally been applied only to Hawaiian species, but is applicable to a more widely
distributed group. Its major distinguishing feature is a more or less ruminate endosperm.
Key words: Amaracarpus, Calycosia, cpDNA, Cremocarpon, Gentianales, Grumilea, Hydnophy-

tinae, Hydnophytum, Mapouria, Myrmecodia, rpsl6 intron, Straussia, Streblosa.
Abbreviations: PGS, preformed germination slit; SCP, ethanol-soluble seed coat pigment.
1 Introduction
The tribe Psychotrieae comprises raphidiate genera with ovaries having a single basally attached
ovule in each locule, drupaceous fruits, and seeds with firm, non-oily endosperm and a propor-
tionally small embryo. Thus characterised, it excludes a number of genera that were included by
Robbrecht (1988, 1994). Analyses of sequence data (Andersson & Rova 1999, Bremer & Manen
2000) have shown that Gaertnera Lam. and Pagamea Aubl. are closely interrelated, but that their
exact relationships to Psychotrieae are uncertain. They have, therefore, been referred to a separate
Subject to copyright. All rights reserved. ? 2002 National Botanic Garden of Belgium
Permission for use must always be obtained from the National Botanic Garden of Belgium. ISSN 1374-7886

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Syst. Geogr. Pl. 72 (2002)
tribe, Gaertnerae, as proposed by Bremekamp (1966). Capuron (unpubl., cited from Piesschaert
2001) included Hymenocnemis Hook.f. in Gaertnera, based on morphological evidence. The gene-
ra Lasianthus Jack, Ronabea Aubl., Saldinia A.Rich. and Trichostachys Hook.f. have been shown
(Piesschaert et al. 2000a) to belong in group not closely related to Psychotrieae. The tribal name
Lasiantheae (Bremer & Manen 2000) applies to this group. Piesschaert et al. showed that
Colletoecema E.M.A.Petit should also be removed from Psychotrieae. It takes an isolated and unre-
solved posistion at the base of the Rubioideae. Sequence data (Andersson & Rova 1999, Bremer &
Manen 2000) have further shown that Declieuxia Kunth does not belong in Psychotrieae. It has later
(Piesschaert et al. 2000b) been transferred to an emended tribe Coccocypseleae. Finally, based on
morphological evidence, Piesschaert (2001) removed a number of genera that have not been
sequenced: Aphanocarpus Steyerm., Coryphothamnus Steyerm., Fergusonia Hook.f., Metabolos
Bl. (Allaeophania Thw.), Pagameopsis Steyerm., and Sarcopygme Setch. & Christoph.
Even though a number of genera have thus been removed, the Psychotrieae remain the the most
species-rich tribe of Rubiaceae, including some 36 genera and well over 2000 species. Generic
delimitations are notoriously difficult in this complex and many, partly contradictory solutions have
been proposed. Some of the problems are related to the fact that the group is morphologically uni-
form, particularly in floral morphology, and that potentially useful characters are inconspicuous and
poorly correlated. Other problems are related to the size and pantropical distribution of the group.
Except for Piesschaert's (2001) study of fruit and seed characters, and the study of Nepokroeff et
al. (1999) based on ITS and rbcL sequence data, no attempt has been made in modem times to
study a sizeable sample of the group throughout its whole range of distribution.
Based on evidence from DNA sequence variation (e.g., Andersson & Rova 1999, Nepokroeff et
al. 1999, Andersson 2001) it appears that the tribe Psychotrieae comprises two major clades.
Andersson (2001) termed these the "Psychotria complex" and the "Palicourea complex", respec-
tively. He suggested that they are usually diagnosable by members of the Psychotria complex
having stipules shed by the formation of an abscission layer, whereas those of the Palicourea com-
plex have non-caducous (but often marcescent) stipules. According to Andersson (2001, 2002) the
Palicourea complex comprises Carapichea Aubl., Chassalia Comm. ex Juss., Margaritopsis Sauv.
(s. lat., including Chazaliella E.M.A.Petit & Verdc., Chytropsia Bremek. and Readea Gillespie),
Geophila Don, Notopleura (Benth.) Bremek., Palicourea Aubl. (s. lat., including most of
Psychotria subgen. Heteropsychotria Steyerm.), and Rudgea Salisb. Although it has not been
sequenced, possession of non-caducous stipules suggests that also Trigonopyren Bremek. belongs
in the Palicourea complex. The Psychotria complex comprises the subtribe Hydnophytinae (sensu
Huxley & Jebb 1991), Psychotria (in the strict sense of Petit 1964), and numerous Pacific species
presently mostly referred to Psychotria.
Many of the problems concerning generic delimitations in the Psychotria complex are related to
circumscription of Psychotria itself. This genus has come to comprise an assemblage of species that
do not possess any striking peculiarities beyond those that characterise the tribe Psychotrieae
(Nepokroeff et al. 1999). Not surprisingly, sequence data have repeatedly (e.g., Andersson & Rova
1999, Nepokroeff et al. 1999, Andersson 2001) suggested that Psychotria is broadly paraphyletic
as presently circumscribed.
Petit (1964) took a first step towards fixing the application of the name Psychotria when typi-
fying P asiatica L. by a specimen in the Linnaean herbarium. He left the question open how this
name should be applied to a recognized species but this question was later settled by Davis et al.
(2001), who showed that the Linnaean type is conspecific with the southern Chinese population of
P. rubra Lour.
168

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
L. Andersson, The Psychotria complex (Rubiaceae)
Petit (1964, 1966) studied African Psychotrieae and suggested that Psychotria should be
restricted to comprise only species having pyrenes without preformed germination slits (PGSs) and
a reddish, ethanol-soluble seed coat pigment (SCP). In doing so, he synonymised Camptopus
Hook.f. and Grumilea Gaertn. under Psychotria. He also suggested that Mapouria Aubl. was a
synonym, at least as far as this name had been applied to African species. He further referred to
Psychotria all such African species as had previously been placed in Cephaelis Sw. or Uragoga
Baill. Without listing particular ones, Petit referred all African species possessing PGSs and lack-
ing SCP either to Chassalia or to Chazaliella. The latter name was not validly published until
eleven years later (Verdcourt 1975). Studies based on sequence data (e.g., Nepokroeff et al. 1999,
Andersson 2001) have later given support to Petit's circumscription by showing that Chassalia and
Chazaliella belong in the Palicourea complex.
Davis & Bridson (pers. comm.), based on the observation that P. asiatica has pyrenes with pre-
formed germination slits (Davis et al. 2001) argued that Psychotria s. str. should be restricted to
comprise species possessing this feature. They did not propose explicitly any other species for
inclusion, however, nor did they consider what to do with the species lacking PGSs. It could be
deduced from the discussion of Davis et al. (2001), however, that species without PGSs would have
to be transferred to Psychotrophum P.Browne. Psychotria is conserved in relation to
Psychotrophum, but Psychotrophum could be used for a genus circumscribed such that it excludes
the type of Psychotria.
Working on Neotropical species, Steyermark (1972) included in Psychotria not only species
lacking PGSs and possessing SCP (characters he did not explicitly discuss), but also all such
Psychotrieae that lacked glandular stipule appendages (these were referred to Rudgea), which were
not herbaceous (most of those were referred to Geophila), and which did not have colourful flowers
with gibbous corolla (and were therefore referred to Palicourea). He subdivided Psychotria into
two subgenera, P. subgen. Psychotria and P. subgen. Heteropsychotria Steyerm. Steyermarks's sub-
genus Psychotria corresponds very closely to Psychotria in Petit's strict sense, but Steyermark used
other characters, namely caducous stipules, conspicuous stipule colleters and a brownish to reddish
drying colour. The subgenus Heteropsychotria possesses none of the characters used by Petit to cir-
cumscribe Psychotria. The type species (P. deflexa) as well as the majority of species placed there
are nested in the same clade as the species referred by Steyermark to Palicourea (Nepokroeff et al.
1999, Andersson 2001). Other species referred by Steyermark to subgen. Heteropsychotria have
been referred to Margaritopsis (Andersson 2001) or Carapichea (Andersson 2002). Thus, all spe-
cies referred by Steyermark to subgen. Heteropsychotria seem to belong in the Palicourea complex.
Curiously, Richard (1834), MUller (1881), Schumann (1891) and Bremekamp (1934) applied
the name Psychotria such that it closely corresponds to Heteropsychotria in Steyermark's sense,
whereas they used Mapouria to cover the same concept for which Steyermark used Psychotria sub-
gen. Psychotria. Bremekamp (1961) also emphasized ruminate endosperm as a feature diagnostic
of Mapouria. In consequence, he later (1963) segregated species with caducous stipules but with-
out "true" rumination as a separate genus, Apomuria Bremek. In Apomuria he included mainly
species from Madagascar and the Comores, but said that the genus should also include numerous
species from continental Africa. Thus, he suggested implicitly that Psychotria as conceived by Petit
is comprised of two natural groups.
Subdivision of Psychotria is treated differently by different authors and how this is done is of
course largely dependent on how the genus is circumscribed. The question of subdivision of
Psychotria is further complicated by the fact that no attempt at a subdivision has endeavoured to
treat the genus throughout its geographical range.
169

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Syst. Geogr Pl. 72 (2002)
Petit (1964, 1966), who circumscribed Psychotria narrowly, subdivided the species of the
African continent into two subgenera, P. subgen. Psychotria and P. subgen. Tetramerae (Hiern)
E.M.A.Petit. He referred all such species that possess bacterial leaf galls to subgenus Tetramerae,
while all species lacking such galls were placed in subgen. Psychotria. It was suggested by
Hamilton (1989) and Taylor (1996) that P microdon (DC.) Urb. and a few related Neotropical
species might also belong in subgen. Tetramerae. Recent molecular studies (Nepokroeff et al. 1999,
Andersson 2001) have confirmed that these species are misplaced in P subgen. Psychotria, but
show that they belong in the Palicourea complex. Andersson (2001) included them in Margari-
topsis s. lat.
Petit (1964) subdivided subgen. Psychotria into five sections (P. sect. Bracteatae Hiern, P sect.
Confertiflorae Hiern., P. sect. Flaviflorae E.M.A.Petit, R sect. Holostipulatae K.Schum., and
R sect. Paniculatae Hiern), leaving twelve species unassigned as "especes de position particulibre"
and another eight species unclassified because they were incompletely known. He did not (Petit
1966) attempt to subdivide formally the subgenus Tetramerae, although an informal grouping was
presented.
Steyermark (1972), who circumscribed Psychotria broadly, subdivided Neotropical species into
two subgenera, P. subgen. Psychotria and P. subgen. Heteropsychotria. As already discussed, sub-
gen. Heteropsychotria belongs in the Palicourea complex. He did not further subdivide the sub-
genus Psychotria.
Except for Piesschaert's (2001) carpological study, generic delimitations among Asian,
Australasian and Pacific Psychotrieae have not been subject to much discussion in recent years, but
a number of genera have been recognised from these areas. Huxley & Jebb (1991) and Robbrecht
(1994) recognized the tuberiferous, myrmecophile epiphytes as a separate subtribe, Hydno-
phytinae, to which five genera were referred: Anthorhiza Huxley & Jebb, Hydnophytum Jack.,
Myrmecodia Jack, Myrmephytum Becc., and Squamellaria Becc. Analyses of sequence data (e.g.,
Andersson & Rova 1999, Nepokroeff et al. 1999) have clearly shown that the segregation of
Hydnophytinae leaves the rest of Psychotrieae paraphyletic.
Hawaiian Psychotrieae have often (e.g., Gray 1858, Hooker 1873, Schumann 1891) been
referred to a separate genus, Straussia (DC.) A.Gray. In recent times, however, Straussia has gene-
rally been regarded as a synonym of Psychotria (e.g., Fosberg 1964, Sohmer 1977, 1978; Darwin
1979). Based on pyrene morphology, Piesschaert (2001) argued that Straussia should be reinstated
as a separate genus.
Some species of Polynesian Psychotrieae have been referred to the genus Calycosia A. Gray.
Although the genus is generally recognized by recent authors (e.g., Darwin 1979, Smith & Darwin
1988, Piesschaert 2001) opinions vary among authors as to its circumscription.
The aim of this paper is to examine relationships within the Psychotria complex based on sequence
variation in one locus of plastid DNA, the rpsl6 intron. Based on the cladogram I will discuss dis-
tribution of such morphological characters that may be diagnostic of natural groups. Finally, I will
discuss the taxonomic implications of this cladogram and propose a generic classification.
2 Material and methods
2.1 Taxon sample

The species dealt with in this study were selected from some 250 species of Psychotrieae for which the sequence
the rpsl6 intron has been determined. These sequences were all analysed together in a preliminary analysis wher
170

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
non-Psychotrieae were used as outgroup. The original sample was taken with two ambitions in mind, first to include at least
two species of each of the Psychotrieae genera recognised by Robbrecht (1988, 1994), second to include a fair sample of
Psychotria species from each of the biogeographic regions (as defined by Mattick in Melchior 1964) from which the genus
has been recorded. In the preliminary analysis, the Psychotrieae form two strongly supported groups, one, the Psychotria
complex, including the type species of Psychotria (P. asiatica), the other one, the Palicourea complex including the type
species of Palicourea (P. guianensis). For the present analysis all taxa were selected that were found in the preliminary
analysis to belong in the Psychotria complex, to which was added an outgroup made up of nine representatives of the
Palicourea complex.
In order to see if a combination of rpsl6 intron data and ITS data are likely to yield stronger support and better resolu-
tion, the rpsl6 intron matrix was trimmed down to those 26 taxa for which ITS sequences are available and the ITS
sequences were added. In these combined data the Chassalia species of the outgroup of the first analysis was replaced by
another Chassalia species, for which a more complete ITS sequence was available.
All species used in this study are listed in tab. 1, where GenBank accession numbers are given together with data about
vouchers and origin of the material.
2.2 Laboratory procedures

Total genomic DNA was extracted from leaf tissue dried in silica gel or taken from herbarium specimens. Laboratory pro-
cedures for extraction, PCR amplification and sequencing were the same as those described by Andersson & Rova (1999)
and Andersson & Andersson (2000). The primers used for sequencing of the rpsl6 intron were rpsF, rpsR2 (Oxelman et al.
1997), rpsMFP, rpsMRP (Persson 2000) and rpsF2 (Andersson 2001). The ITS region was sequenced using the primers
ITS1, ITS2, ITS3 and ITS4 of White et al. (1990).
Table 1. List of taxa used in cladistic analyses.

Sequences that are new to this study are marked by an asterisk. Notes: ') = Psychotria ligularis (Rudge) Steyerm.; 2) culti-
vated at the G6teborg Botanic Garden; 3) cultivated at the Oxford Botanic Garden; 4) cultivated at the Royal Botanic Gar-
dens, Kew; 5) cultivated at the Royal Botanic Garden, Edinburgh, accession no. 19780778A; 6) as R rubra; 7) cultivated at
the National Botanic Garden, Meise, accession number 62-6512; s) cultivated at the Lyon Arboretum; 9) cultivated at the
National Tropical Pacific Garden, Hawaii;'o) cultivated at the National Botanic Garden Meise, accession number 53-6779;
") cultivated at the National Botanic Garden Meise, accession number 62-0513; 12) cultivated at the National Botanic
Garden Meise, accession number 82-0401; '3) cultivated in the Botanical Garden of the University of Vienna, accession
MK940911-3/1; 4) P. kirkii var. mucronata (Hiern) Verdc.;5) P. kirkii var. nairobiensis (Bremek.) Verdc.; '6) cultivated
at the National Botanic Garden Meise, accession number 07-3791; 7) P. kirkii var. tarambassica (Bremek.) Verdc.; '") culti-
vated at the National Botanical Garden, Meise, accession number 75-0204. 9) = P. kirkii var. volkensii (K. Schum.) Verdc.
Taxon Origin Voucher NCBI acc no.

rps16 intro ITS
Amaracarpus grandifolius Valeton New Guinea Takeuchi 8926 (L) AF410678* -

A. kochiiValeton Irian Jaya Davis 680 (K) AF410679* -
Calycosia lageniformis (Gillespie) Fiji Rova & Gustavsson 2461 (GB) AF410680* -
A.C. Sm.
C. macrocyatha Fosberg Fiji Rova & Gustavsson 2486 (GB) AF410671* -

Carapichea guianensis Aubl.' French Guiana Andersson et al. 1989 (GB) AF147567 AF149390*
Chassalia subcordatifolia (De Wild.) Gabon Andersson & Nilsson 2314 (GB) AF369840 AF410683*
F. Piesschaert
C. subochreata (De Wild) Tanzania Frimodt-Moller 2 (C) AF369841

Robyns
Cremocarpon [?] sp. "1106" Madagascar Davis 1106 (K) AF410684*
Dolianthus vaccinioides C.H.Wright New Guinea Hopkins 680 (L) AF410685*
Geophila repens (L.) I.M. Johnst. Ecuador Balslev et al. 97211 (AAU) AF369846 AF410686*
Hydnophytum formicarium Jack unknown2 Andersson 2216 (GB) AF001339 AF034912
H. grandiflorum Becc. Fiji Rova & Gustavsson 2445 (GB) AF369847 -
H. moseleyanum Becc. New Guinea3 without voucher AF410687* AF034913
Hymenocoleus hirsutus (Benth.) Robbr. Congo-Kinshasa Lejoly 81/330 (BR) AF369848 AF072018
"Mapouria"anakfinensis (K. Schum.) Madagascar Davis 1001 (K) AF410688* -
Bremek.
"M." macrochlamys Bremek. Madagascar Davis 1005 (K) AF410689* -

Margaritopsis acuifolia C. Wright Cuba Ekman 16009 (S) AF001340 -
Myrmecodia horrida Huxley & Jebb New Guinea3 without voucher AF410690* -
M. tuberosa Jack unknown2 Andersson 2217 (GB) AF000950 AF149313*
171

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Table 1. Continuation.

rpsl6 intro ITS
Myrmephytum selebicum (Becc.) Becc. unknown3 without voucher AF410691* AF034916

Notopleura guadalupensis (DC.) Puerto Rico Howard 16814 (S) AF147562 AF149384*
C.M.Taylor
Palicourea guianensis Aubl. French Guiana Andersson et al. 2015 (GB) AF001345 AF149328*
Psychotria albicaulis Scott & Elliott Ivory Coast Leeuwenberg 3034 (BR) AF410692*
PF anceps Kunth Guyana Taylor 12078 (MO) AF147544*
P ankasensis J.B.Hall Ghana4 Chase 2265 (K) AF001347
PF asiatica L. Hong Kong5 without voucher AF369854 AF0720356
PF auxopoda E.M.A.Petit Congo-Kinshasa Liengda ILBi 11 (BR) AF410693* -
P avilensis Steyerm. Venezuela Jaimes 97201 (VEN) AF410694* -
P. aff. bangueyensis Merr. Borneo Beaman 9711 (L) AF410695* -
P biaurita (Hutch. & Dalz.) Verdc. Liberia Jacques-Georges 27468 (BR) AF410682* -
P brachyantha Hiern Ghana Jongkind & Abbiw 1619 (BR) AF410696* -
P brachyanthoides De Wild. Ivory Coast Cremers 1135 (BR) AF410697* -
P brackenridgei A. Gray Fiji Rova & Gustavsson 2458 (GB) AF410698* -
P. brevicalyx Fosberg Fiji Rova & Gustavsson 2432 (GB) AF410699* -
P brevipuberula E.M.A.Petit Congo-Kinshasa Leonard et al. 1851 (BR) AF4
F brownei Spreng. Hispaniola Stahl & Lindstr6m 149 (GB) AF369856 -
PF butayei De Wild. Angola (?) Raimundo et al. 5 (BR) AF410701* -
PF calva Hiern unknown7 without voucher AF410702* -
P capensis Vatke South Africa Chase 2268 (K) AF005390 AF149370*
PF carthagenensis Jacq. French Guiana Andersson et al. 1987 (GB) AF410703* -
PF chrysantha Merrill & Perry New Guinea Pullen 381 (L) AF410704* -
PF cinerea De Wild. Tanzania Bidgood et al. 2768 (BR) AF410705* -
PF coelosperma F.M.Bailey Australia, Hyland 6005 (L) AF410706* -
Queensland
F comperei E.M.A.Petit Congo Champluvier 5331 (BR) AF410707* -
PF coptosperma (Baill.) Guillaumin New Caledonia MacKee 17073 (L) AF410708* -
PF confertiloba A.C.Sm. Fiji Rova & Gustavsson 2469 (GB) AF369857 -
R cryptogrammata E.M.A.Petit Tanzania Bidgood et al. 487 (BR) AF410709* -
PF cupularis (M011. Arg.) Standl. Guyana Jansen-Jacobs et al. 4372 (GB) AF410710* -
PF daphnoides A.Cunn. ex Hook. Australia, Lebler & Durrington s.n. AF369858 -
Queensland 28.xi.1972 (AAU)
P djumaensis De Wild. Guinea Adam 27506 (BR) AF410711* -
P eminiana (Kuntze) E.M.A.Petit Tanzania Bidgood et al. 2634 (BR) AF410712* -
P. aff. ernestii K.Krause Bolivia Persson & Gustafsson 236 (GB) AF147559* AF149381*
PF erythrocarpa Standl. Mexico' Lyon Arboretum 82.02.43001 AF410713* -
(GB)
P. faucicola K.Schum. Kenya Polhill & Robertson (BR) AF410714* -
PF flava Standl. Mexico4 Chase 2914 (K) AF001350 -
R gabonica Hiern Cameroon Cable 1326 (K) AF410715* -
F grandiflora Mann Hawaii Skottsberg 1019 (GB) AF410716* -
F grandis Sw. Cuba Rova et al. 2267 (GB) AF410717* -
P gyrulosa Stapf Borneo Beaman et al. 9230 (S) AF410718* -
P hathewayii Fosb. Hawaii Takeuchi 2313 (GB) AF410719* -
FP heterosticta E.M.A.Petit Burundi Reekmans 7461 (BR) AF410720* -
P holtzii (K. Schum.) E.M.A.Petit Kenya Napper 1369 (BR) AF410721* -
PF horizontalis Sw. Venezuela Jaimes 97203 (VEN) AF410722* -
Panama Wright s.n. (STRI) - AF072047
PF humilis Hiern Nigeria Brenan 8392 (BR) AF410723*
R insularum A.Gray Samoa9 Lorence 8026 (PTBG) AF410724* AF149388*
F iringensis E.M.A.Petit Tanzania Lovett & Thomas 2458 (BR) AF410725*
R iteophylla Stapf Borneo Axelius 303 (S) AF410726*
R kaduana (Cham. & Schltdl.) Fosberg Hawaii" Lyon Arboretum 67.
PF kimuenzae De Wild. Congo-Kinshasa Van Hove 76 (BR) AF410727*
PF kirkii Hiern unknown'o without voucher AF410728*
unknown unknown - AF072038"
172

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC

rps16 intro ITS
PF leucopoda E.M.A.Petit Kenya Polhill & Robertson 4831 (BR) AF410730*

PF linearisepala E.M.A.Petit Malawi Pope et al. 2260 (BR) AF410731* -
PF loniceroides Sieb. ex DC. Australia, N.S.W. Nordenstam & Anderberg AF410732* -
1369 (S)
Australia, N.S.W. Telford 8522 (herb. not cited) - AF072034
PF lucens Hiern unknown" without voucher AF410733"
P maingayi Hook.f. unknown12 without voucher AF001352
FP mapourioides DC. French Guiana Andersson et al. 1911 (GB) AF001353 A
P? mariana Bartl. ex DC. Mariana Islands9 Lorence 7959 (PTBG) AF147570 AF149395*
PF mariniana (Cham. & Schltdi.) Fosb. Hawaii'3 Kiehn 940911 (WU) AF001354
Hawaii Nepokroeff & Flynn 951 (WIS) - AF034904
FP megistophylla Standl. Colombia Devia 5745 (MO) AF410734* -
FP miombicola Verdc. Tanzania Bidgood et al. 596 (BR) AF410735* -
FP mucronata Hiern'" Congo-Kinshasa d'Huart 89 (BR) AF410736* -
PF nairobiensis Bremek.1s Tanzania Lippert 5908 (BR) AF410737" -
F? nervosa Sw. Cuba Rova et al. 2249 (GB) AF410738" -
Puerto Rico Taylor & Miller 10415 (MO) - AF072046
PF obovata Wall. Sumatra Boeea 6609 (S) AF410739* -
PF orophila E.M.A.Petit Kenya Bytebier & Kirika 36 (BR) AF410740* -
PF parvula A.Gray Fiji Rova & Gustavsson 2478 (GB) AF410741" -
P peduncularis (Salisb.) Steyerm. unknown'6 without voucher AF410742* -
"Africa" without voucher - AF072044
P? aff. pichisensis Standl. Bolivia Persson &

PF plantaginoidea E.M.A.Petit Congo-Kinshasa B
FP psychotrioides (DC) Roberts Congo-Kins
PF psychotrioides (Heller) Fosb. Hawaii Takeuchi 2468 (GB) AF410746* -
P. aff. pumila Hiern Malawi Blackmore & Brummitt 1480 (BR) AF410747" -
PF punctata Vatke Tanzania Van Have 14 (BR) AF369865 -
P remota Benth. Bolivia Persson & Gustafsson 258 (GB) AF147576 AF149403*
PF riparia (K.Schum. & K.Krause) Kenya Gillett 19274 (BR) AF410748* -
E.M.A.Petit
FP rubra (Lour.) Poir. Japan Aiba 4 (GB) AF410749* -

PF rupicola Schltr. New Caledonia Selling 42 (S) AF410750* -
PF sarmentosa BI. Sumatra Jacobs 8052 (C) AF410751* -
PF schweinfurthii Hiern Cameroon Breteler 2738 (BR) AF410752* -
P? serfipiles De Wild. Guinea Jacques-Georges 27519 (BR) AF410753* -
P serpens L. Hong Kong9 Flynn 1716 (PTBG) AF147577* AF149404*
PF siamica (Craib) Hutch. China, Kwangsi Steward & Cheo 1206 (S) AF410754" -
PF spithamea S. Moore Tanzania Richards 11831 (BR) AF410755* -
PF succulenta (Schweinf.) Hiern Zimbabwe Richards 2024 (BR) AF410756* -
P? tarambassica Bremek.'7 Kenya S. Paulo 1044 (BR) AF410757* -
PF tenuissima E.M.A.Petit Congo-Kinshasa Malaisse 10620 (BR) AF410758* -
PF tephrosantha A. Gray Fiji Rova & Gustavsson 2448 (GB) AF411077* -
FP trichotoma Mart. & Gal. Ecuador Stahl 2944 (GB) AF410759* -
P. turbinata A.Gray Fiji Rova & Gustavsson 2431 (GB) AF410760* -
P. unicarinata (Fosberg) A.C.Sm. Fiji Rova & Gustavsson 2430 (GB) AF410761" -
& S.Darwin
PF verschuerenii De Wild."1 unknown'" without voucher AF410762" -

PF viridis Ruiz & Pav. Bolivia Persson & Gustafsson 209 (GB) AF410763* -
PF vogeliana Benth. unknown Pierret 16 (BR) AF410764* -
PF volkensii K. Schum.'" Tanzania Carmichael 1276 (BR) AF410765" -
PF zombamontana E.M.A.Petit Zimbabwe Allard 557 (BR) AF410766* -
P. sp. "2350" Gabon Andersson & Nilsson 2350 (GB) AF410767* -
P. sp. "2489" Fiji Rova & Gustavsson 2489 (GB) AF410768* -
Rudgea hostmanniana Benth. French Guiana Andersson et al. 1954 (GB)
Squamellaria imberbis (A. Gray) Becc. Fiji A.C. Smith 8326 (S) AF003620
Streblosa aff. myriocarpa Merr. Borneo Nielsen 517 (AAU) AF003621
173

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
2.3 Alignment and indel coding

Sequences were aligned by hand such that the number of informative mutations was minimised. One inversion was replaced
by its reverse motif. In the rpsl6 data, insertions, deletions and the inversion were coded as separate characters, following
the rules specified by Andersson & Chase (2001). No indel coding was applied to ITS data. In rpsl6 data, one section
(28-133 bp) was found impossible to align unambiguously and was left out of the analysis. Another section (18-30 bp)
could not be aligned in the outgroup and was replaced by gaps there. Two sections of the ITS sequences (35-37 bp of ITS 1
and 22-27 bp of ITS2) could not be aligned and were replaced by gaps in outgroup taxa.
2.4 Morphological data

Data on selected morphological characters were compiled into a matrix (tab. 2). The characters selected are such that have
been considered important by Bremekamp (1934, 1961, 1963), Petit (1964) and Steyermark (1972). Data were taken from
literature reports and personal observations, as specified in tab. 2, and were used only to explore the possibilities to charac-
terise morphologically the clades found based on molecular data. Thus, they were not used in cladistic analyses, only for
optimisation on a predefined tree.
Because detailed morphological data were unavailable for a large proportion of the sequenced taxa, the molecular tree
was pruned prior to optimization. Thus, the tree used for optimization (fig. 5-9) included only those 55 taxa for which
detailed information on pyrene morphology was available. A hypothetical taxon, representing an imaginary outgroup was
added to the base of the tree. I used a hypothetical outgroup because presently available cladistic analyses (Andersson
unpubl, Andersson & Rova 1999, Bremer & Manen 2000, Piesschaert et al. 2000) disagree about which group is the actual
sister group of Psychotrieae. Among the possible candidates, only three tribes have comparable drupaceous fruits, viz.
Gaertnereae, Morindeae and Prismatomerideae. The hypothetical outgroup was coded such that it is polymorphic for all
states occurring among these tribes.
Table 2. Morphological characters used for optimisation (fig. 5-9).

Characters and states. 1: stipules non-caducous (0) or caducous (1); 2 marginal PGSs absent (0) or present (1); 3: abaxial
pyrene surface without ridges (0), with 1-3 ridges (1), with 4-6 ridges (2); 4: ethanol-soluble seed coat pigment absent
(0) or present (1); 5: endosperm non-ruminate (0), with a single, median, adaxial intrusion (1), or truly ruminate (2).
Symbols for source of information: a = personal observation, b = Piesschaert 2001, c = A. Davis & D. Bridson (pers.
comm.), d = van Royen 1973. Notes: ') as S. wilsonii; 2) as P. asiatica; 3) "almost absent", 4) the codings refer to the
specimen Hallier 906, det. Valeton; another specimen is distinctly different and probably not conspecific; 5) as Psychotria
ligularis; 6) as M. nudiflora; 7) codings for this terminal is a combination of the features occurring in Gaertnereae and
Morindeae s.lat. Judging from molecular analyses (e.g., Andersson & Rova 1999, Bremer & Manen 2000) these are the
only groups likely to be the sister group of Psychotrieae and in which the fruits are drupaceous.
Taxon Source Characters

1 2 3 4 5
PACIFIC CLADE
Hydnophytum formicarium a, b 0 1 0 0 0
Myrmecodia tuberosa a, b 0 0 0 0 0
Squamellaria imberbis a, b' 0 1 0 0 0
Psychotria brackenridgei a, b 1 1 1 1 0
Calycosia lageniformis a 1 1 1 0 0
Psychotria brevicalyx a 1 1 1 0 0
P turbinata a, b 1 1 1,2 ? 0
P unicarinata a 1 1 1 0 0
P sp. "2489" a 1 1 2 0 1
Amaracarpus sp. c 1 1 1,2 ? 0
Dolianthus vaccinioides c, d 1 1 1,2 ? 0
Psychotria chrysantha b ? 1 0 1 2
P confertiloba a, b 1 1 1 1 2
P gyrulosa b ? 1 0 1 2
P hathewayii a 1 1 0 0 2
P mariana b ? 1 0 1 2
P mariniana a, b 1 1 0 1 1
Streblosa b ? 0 1,2 1 0
PSYCHOTRIA S. STR. I
Psychotria brownei a, b' 1 0 2 1 2
P carthagenensis a, b 1 0 2 1 2
P ernestii a, b 1 0 2 1 ?3
P flava b ? 0 1,2 1 2
174

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Taxon Source Characters

1 2 3 4 5
P grandis a, b 1 0 0 1 2
P horizontalis a, b 1 0 2 1 ?3
P nervosa a, b 1 0 2 1 2
P peduncularis b ? 0 2 1 2
P trichotoma a, b 1 0 2 1 2
P viridis a, b 1 0 2 1 2
P vogeliana a, b 1 0 1,2 1 2
PSYCHOTRIA S. STR. II
P calva a, b 0 0 0,1 1 0
P heterosticta a, b 0 0 0 1 2
P kimuenzae a, b 0 0 0 1 0
P punctata a, b 0 0 0 1 1
PSYCHOTRIA S. STR. III
P asiatica a, c 1 1,0 2 1 1
P auxopoda a, b 1 0 0 1 1
R capensis a, b 1 0 0 1 2
P djumaensis a, b 1 0 2 1 2
P eminiana a, b 1 0 0 1 2
P mapourioides a, b 1 0 2 1 2
P orophila a, b 1 0 2 1 2
P remota a, b 1 0 2 1 2
P succulenta a, b 1 0 0 1 2
PSYCHOTRIA S. STR. IV
P daphnoides a, b 1 0 1 1 2
P loniceroides a, b 1 0,1 1,2 1 0,2
P rupicola a, b 1 0 1 1 ?
P sarmentosa b4 ? 0 0 1 2
P tephrosantha a, b 1 1 1 1 0
OUTGROUP
Carapichea guianensis a, b5 0 0 0,1,2 0 0
Chassalia subochreata a, b 0 1 1 0 0
Geophila repens a, b 0 0 0,1,2 0 0
Hymenocoleus hirsutus a, b 0 1 0 0 0
Margaritopsis acuifolia a, b6 0 1 0 0 0
Notopleura guadalupensis a, b 0 1 0 0 0
Palicourea guianensis a, b 0 1 1 0 0
Rudgea hostmanniana a, b 0 1 1,2 0 1
hypothetical7 b ? 1 0 0,1 0,2
2.5 Cladistic procedures

All cladistic analyses were made with
gested that very large numbers of maxi
all. Instead, I used parsimony jackknifing
was made only in order to get an estimat
The jackknife analysis was made with
TBR branch swapping and saving only 1
of random addition sequence were run,
Optimisation of morphological characte
3 Results
The finally aligned rpsl6 matrix comprised 956 characters, 674 constant, 122 variable but un
informative, and 160 informative. Among informative characters 27 were codes for insertio
175

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
myrmecophiles
Amaracarpus
63
63 Calycosia n
Co 0
95 Dolianthus
Straussia
-"0"0
'
StreblosaCo5"0
63 Psychotria'D"
95
92 Psychotria
clade I o
clade II
54 Psychotria clade III nx
89 Psychotria
100 "Cremocarpon" clade IV"CO
sp. "1106"
"Mapouria" anakfinensis
68 Notopleura guadalupensis
Rudgea hostmanniana -o
Palicourea guianensis8-
Margaritopsis acuifolia o
76
Carapichea guianensis o
56 76 Hymenocoleus hirsutus -
Chassalia subochreata x
83 Geophila repens
Figure 1. Results from jackknife analysis of rpsl6 intron data: overview of major topological features of the
sensus tree. Details are shown in fig. 2-3, as indicated by brackets in this figure. Figures above branches ar
(37% deletion) support values. Boxes representing major clades and grades are drawn such that their horizon
sion corresponds to the maximum divergence among the taxa they represent.
deletions and inversions. The jackknife consensus tree is shown in fig. 1-3. The heuris
found 8704 maximally parsimonious trees and many more would probably have been f
the search been continued. The trees found were 460 steps long, with a consistency in
(without uninformative characters), a retention index of 0.91 and homoplasy index of 0
The aligned matrix of combined rpsl6 and ITS data comprised 1485 characters, 1013 co
variable but uninformative and 292 informative. Of these, 854 characters (84 informa
derived from rpsl6 and 631 (208 informative) from the ITS. Based on the full matrix, a
parsimonious tree (fig. 4) was found. This was 1018 steps long, with a consistency ind
(uninformative characters excluded) and a retention index of 0.68. Using only rpsl6 d
matrix gave 28 MPTs with a length of 206 steps, a consistency index of 0.75 and reten
of 0.90. Using only ITS data gave 48 MPTs with a length of 801 steps, a consistency in
and retention index 0.62.
Most branches are more strongly supported by combined data than by any one of the compo-
nent data sets alone. The exceptions are mainly those branches involved in conflicting topologies,
but also the one supporting P. ernestii, P. horizontalis, P nervosa and P peduncularis. Based on
ITS data this is not supported at all in the jackknife, based on rpsl6 data it is supported by 83%,
and it is supported by only 64% based on combined data.
176

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Based on rpsl6 (fig. 1) as well as combined data (fig. 4), the partition between ingroup and out-
group is supported in 100% of the jackknife replicates and no maximally parsimonious tree was
found where this partition is not present.
Both data sets show two major clades, which will henceforth be called the Pacific clade and
Psychotria sensu stricto. The Pacific clade is strongly supported based on both data sets. Psychotria
s. str. is moderately supported based on rpsl6 data alone and strongly supported based on combined
data.
In all analyses, all sampled myrmecophiles belong in one strongly supported subclade (fig. 2,
4). Differences between data sets with respect to internal resolution is largely due to the fact that
Hydnophytum grandiflorum and Squamellaria imberbis are not represented in the combined data
set. Otherwise the two topologies are compatible. There is strong support, however, that the three
Hydnophytum species of the rpsl6 data set do not form a monophyletic group. Two species of
Calycosia together with several Fijian species of Psychotria (in the broad sense of Smith & Darwin
1988) form a strongly supported clade based on rpsl6 data (fig. 2). None of them is represented in
the combined data set. There is ambiguous support that Psychotria grandiflora is the sister of this
Calycosia clade. In the rpsl6 analysis, two species of Amarcarpus form a moderately supported
group together with Psychotria insularum. Dolianthus vaccinioides is the sister of this group, but
support is ambiguous. Three Hawaiian species of Psychotria form one weakly supported group, and
Psychotria gyrulosa and R iteophylla form another one. There is weak support that Streblosa myri-
ocarpa is the sister of the remainder of the Pacific clade.
The rpsl6 analysis does not show any other supported relationships within the Pacific clade,
which here includes an unresolved residue comprising a number of species of Psychotria in the
broad sense of Smith & Darwin (1988) and Sohmer (1977). The Hawaiian species of Psychotria
that belong here are largely such that have been referred to the genus Straussia. Based on combined
data, relationships among the four species representing this residue are completely resolved, but
support for this resolution is weak or lacking, except that there is strong support from ITS data that
Psychotria kaduana and R mariniana belong together.
Details of Psychotria s. str. are shown in fig. 3. Although internal resolution is largely poor,
there are also some more or less strongly supported groups. In order to facilitate reference to this,
four groups have been labelled in fig. 3. As shown by fig. 1, there are two species, "Cremocarpon"
sp. "1106" and "Mapouria" anakfinensis, which do not belong to any of these groups and which
are not shown in fig. 3. The Psychotria s. str. clade includes the type species of Psychotria (P asi-
atica in clade III), Mapouria (P mapourioides in clade III; but see discussion of the name
Mapouria), Psychotrophum (P brownei in clade I), and Psychotria subgen. Tetramerae (R puncta-
ta in clade II). It also comprises two species of Cremocarpon (as this was circumscribed by
Piesschaert 2001): Cremocarpon sp. "1106" (without resolved relationships) and R rupicola (in
clade IV). Psychotria s. str. is represented by species from the Neotropics, Africa, Indomalaysia,
New Guinea, and Australasia, including Australia. Strikingly, none of the Hawaiian species come
out in this clade.
4 Discussion
4.1 Major features

As far as differences in taxon sample allow comparisons, and as far as well supported relations
are concerned, results from the rpsl6 analysis are congruent with the ITS tree of Nepokroeff e
(1999). Both trees show two major clades within the Psychotria complex (i.e. "Psychotria sens
177

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
63 Myrmecodia horrida
66 Myrmecodia tuberosa
Myrmephytum selebicum
Hydnophytum moseleyanum
Hydnophytum formicarium
99.C Hydnophytum grandifiorum

Squamellaria imberbis
Psychotria grandiflora
Calycosia lageniformis
62 - Calycosia macrocyatha
Psychotria brackenridgei
8 Psychotria brevicalyx
Psychotria turbinata
Psychotria unicarinata
Psychotria sp. "2489"
63 Amaracarpus grandifolius
Amaracarpus kochii
Psychotria insularum
Dolianthus vaccinioides
Psychotria kaduana
63 Psychotria mariniana
Psychotria psychotrioides (Heller) Fosb.
am .Psychotria gyrulosa
Psychotria iteophylla
Psychotria chrysantha
Psychotria confertiloba
Psychotria hathewayii
Psychotria mariniana
Streblosa myriocarpa 5 changes
Figure 2. Results of jackknife analysis of rpsl6 intron data (part of jackknife

(cf fig. 1). Figures at branches are jackknife (37% deletion) suppor
178

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Psychotria carthagenensis 63 Psychotria faucicola
Psychotria ernestii 63 Psychotria tarambassica*
Psychotria pichisensis L Psychotria volkensii*
87 Psychotria viridis 98 Psychotria kirkii
97Psychotria
a93 avilensis
Psychota i ae
9 Psychotria maingayi
61 - Psychotria horizontalis I Psychotria mucronata*
Psychotria megistophylla - Psychotria nairobiensis*
Psychotria brownei - Psychotria punctata
Psychotria peduncularis " Psychotria butayei
99 Psychotria biaurita 62 Psychotria miombicola
64
92 Psychotria b 63
brachyanthoides _0
-- Psychotria iringensis
Psychotria schweinfurthii Psychotria tenuissima
Psychotria serfipiles Psychotria cinerea o
63 o
Psychotria flava 63 Psychotria sp. "2350" 0
87 h =
Psychotria
Psychotria grandis
trichotoma c 87 plantaginoidea_
Psychotria Psychotria heterosticta
96 Psychotria erythrocarpa d Psychotria brachyantha
Psychotria nervosa I Psychotria cryptogrammata
Psychotria lucens Psychotria holtzii
Psychotria humilis
99 Psychotria asiatica Psychotria kimuenzae
5 Psychotria rubra Psychotria leucopoda
Psychotria bangueyensis . Psychotria linearisepala
Psychotria siamica - Psychotria pumila
Psychotria anceps Psychotria spithamea
Psychotria cupularis Psychotria verschuerenii
Psychotria mapourioides
Psychotria remota Psychotria rupicola
Psychotria capensis 71 Psychotria obovata 0
Psychotria eminiana Psychotria sarmentosa a
Psychotria riparia k 98 Psychotria parvula 0
._Psychotria
Psychotriagabonicaalbicaulis
72 Psychotria Psychotria
coelosperma o teph . 72
Psychotria auxopoda Q Psychotria coptosperma o

87_.
Psychotria brevipuberula I Psychotria daphnoides
62 Psychotria comperei Psychotria loniceroides
Psychotria succulenta Psychotria serpens
Psychotria djumaensis
m
Psychotria orophila
Psychotria zombamontana
SPsychotria ankasensis
Psychotria psychotrioides (DC.) Roberts
"Mapouria" macrochlamys 10 changes
Figure 3. Results of jackknife analysis of rpsl6 intron data (part o

(cf fig. 1). Brackets show extension of major (I-IV) and minor clad
Generic names that appear within quotes are such that are conside
nation exists with Psychotria. Names in group e that are followed b
(1975) to varieties of P. kirkii. Figures above branches are jack

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
stricto" in the sense of Nepokroeff et al.). Fifteen species are common to Nepokroeff's et al. sam-
ple and the rpsl6 sample and all of them appear in the same clades. Nepokroeff's et al. rbcL tree
differs in that Psychotria graciliflora Benth. is placed as the sister of all other species of the
Psychotria complex. However, this placement has no support and in the ITS tree the same species
is nested deeply in one of the two clades, viz. in "Psychotria s. str." in the present sense (fig. 1).
s99/501s 9Myrmecodia tuberosa
100/100100 Myrmephytum selebicum

Hydnophytum formicarium
63/<50ro4 Psychotria insularum

o100101100- Hydnophytum moseleyanum 0
100/(r Psychotria kaduana D

9961389 Psychotria mariniana
Psychotria mariana
97,s3i<o ~Psychotria ernesti

62/<50/<50 Psychotria horizontalis
64/83/-60 Psychotria nervosa
Psychotria peduncularis
72/-70 98/981
s9 r Psychotria mapourioides 0
991-198 Psychotria remota
55/40/
<50 Psychotria capensis
96 91/72
Psychotria kirkii
Psychotria asiatica
65/-/91 Psychotria loniceroides

99/95/97 Psychotria serpens
71/-/88 - Notopleura guadalupensis

Palicourea guianensis -o
Rudgea hostmanniana ' 8
Carapichea guianensis i o
81/91/
Hymenocoleus hirsutus .8
92n89 Chassalia subcordatifolia (
20 changes 98/64/97 Geophila repens
Figure 4. Results of the analysis of combined ITS and rpsl6 intron data: the single most parsimonious tree.
extension of outgroup and the two major clades of the ingroup. Figures at branches are jackknife (37% del
values and are given thus: support based on combined data/support based on rpsl6 intron data alone/ suppo
ITS data alone. For branches that do not appear in a jackknife consensus tree, but are not contradicted eit
support is given as "<50". When branches occur in this tree but are contradicted by a jackknife consensus
on separate data, this is implied by a hyphen.
180

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Based on ITS data Nepokroeff et al. (1999) found only 264 most parsimonious trees, as com-
pared to >8704 in the present analysis. Unsurprisingly, the strict consensus of the ITS trees is
generally more resolved than the jackknife consensus tree from the rps16 analysis. The resolution
of the ITS tree is largely unsupported by jackknife analysis, however. The maximally parsimonious
ITS trees (with 85 terminal taxa) were found to have a retention index of 0.67 whereas those of the
rpsl6 intron trees (with 120 terminals) had a retention index of 0.91. These differences may be
explained in part by the fact that the ITS data set included a taxonomically broader sample of taxa,
in which more parallelism should be expected. However, it seems to be a recurrent pattern that
homoplasy is greater in ITS data than in data from plastid gene introns (cf. for example Plunkett &
Downie 1999). Consistent with differences in homoplasy levels, support values are on the average
higher in the rpsl6 intron analysis.
The present analyses of a matrix comprised of combined ITS and rps16 data yields essentially
the same results: ITS data give fewer trees and thus a more resolved phylogeny than rpsl6 data do,
but ITS data are more homoplasious. Since ITS and rpsl6 intron data largely give support to dif-
ferent but still compatible relationships, it is to be expected that combination of data sets may yield
phylogenies that are at the same time better resolved and better supported. There seems to be little
genuine conflict between ITS and rpsl6 data.
4.2 Character distribution

Andersson (2001) suggested that the Psychotria complex is characterised by caducous stipule
the Palicourea complex by non-caducous. The present analysis shows the same pattern (fig. 5),
with considerably more detail with respect to the Psychotria complex. The present optimisa
unequivocally suggests that presence of caducous stipules is the ancestral state of the Psych
complex and that non-caducous stipules is due to secondary reversals within the group. Non
caducous stipules occur in only four places, namely among the myrmecophiles, in Amarcarpus
Psychotria clade II, and in P. rupicola. Parallel occurrences of caducous stipules are unkn
among the approximately 125 species of the Palicourea complex that have so far been sequen
(Andersson 2001 and unpubl.). Based on these facts, it may be said that whereas non-caducous
ules do not, in itself, allow any very strong conclusions about relationship, caducous stipules d
nitely suggest membership of the Psychotria complex. Furthermore, any member of the Psych
complex having non-caducous stipules should be expected to be either a tuberiferous myrme
cophile or to have bacterial nodules, these features being diagnostic of the larger groups of
Psychotria complex in which non-caducous stipules occur.
Optimisation of presence of marginal PGSs (fig. 6) shows that the state is present in
Psychotrieae ancestor and the Palicourea complex ancestor, whereas it is equivocal in the Pac
clade ancestor. Even though it is a homoplasious character, absence of marginal PGSs appear
be a synapomorphy of Psychotria s. str. Still, its diagnostic value is reduced because it is de
strably homoplasious and it cannot be used with confidence in isolation.
The optimisation shown here differs strikingly from the one shown by Nepokroeff et al. (1
where the entire Psychotria complex seems to be consistently characterised by absence of P
Nepokroeff et al. did not give any precise information about the sources of their morpholo
data, except a reference to Petit (1964), who did not examine any members of the Pacific clad
number of taxa of the Pacific clade scored by Nepokroeff et al. as lacking PGSs have been e
ined both by Piesschaert (2001) and myself and we both agree that they have distinct marg
PGSs. I conclude that Nepokroeff's et al. optimisation was based on largely inadequate morph
logical data.
181

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
stipules caducous Hydnophytum formicarium

[I ] stipules non-caducous Myrmecodia tuberosa
state unknown/equivocal I Squamellaria imberbis
Calycosia lageniformis
Psychotria brevicalyx
Psychotria turbinata -o
Amarcarpus kochii o
Psychotria chrysantha a
Psychotria gyrulosa
Psychotria mariana
Streblosa myriocarpa
Psychotria brownei
Psychotria carthagenensis
Psychotria emestii
Psychotria viridis -
Psychotria horizontalis 0%
Psychotria flava 8 0
Psychotria grandis 8
Psychotria trichotoma -
Psychotria vogeliana
Psychotria nervosa -u
Psychotria calva -I
Psychotria heterosticta 0
Psychotria kimuenzae I o
Psychotria
Psychotriapunctata
asiatica V.
Psychotria auxopoda
Psychotria capensis
Psychotria eminiana 0
Psychotria mapourioides '
Psychotria remota
Psychotria djumaensis -
Psychotria orophila
Psychotria succulenta
Psychotria daphnoides
Psychotria loniceroides 0
Psychotria sarmentosa 0
Psychotria tephrosantha
Psychotria rupicola -
Carapichea guianensis -o
Chassalia subochreata
Geophila repens 0
Hymenocoleus hirsutus (
Margaritopsis acuifolia
Palicourea guianensis
Notopleura guadaloupensis o
Rudgea hostmanniana
hvaothetical
Figure 5. Stipule persistence optimised on the tree resulting from the jackknife analysis of rpsl6 intron data.
pruned to comprise only those terminal taxa for which detailed carpological data are available. The hypothet
group represents a summary of all groups presumed to be possibly the sister group of Psychotrieae. Branche
terminals for which information is missing are marked as equivocal.
182

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
marginal PGSs Hydnophytum formicarium

present ? Myrmecodia tuberosa
C absent Squamellaria imberbis
equivocal/polymorphic Calycosia lageniformis
Psychotria turbinata -o
Amarcarpus kochii o
Dolianthus vaccinioides 9.
Psychotria chrysantha (
Psychotria gyrulosa
Psychotria mariana
Psychotria brownei
Psychotria ernestii
Psychotria viridis -
Psychotria
Psychotria horizontalis
flava -
Psychotria trichotoma 3
Psychotria nervosa 1
Psychotria calva 0
Psychotria heterosticta ?
Psychotria kimuenzae
Psychotria punctata
Psychotria asiatica
Psychotria auxopoda
Psychotria capensis
Psychotria eminiana '
Psychotria remota
I Psychotria orophila
Psychotria loniceroides
Psychotria sarmentosa
Psychotria rupicola
Carapichea guianensis -o
Geophila repens
Hymenocoleus hirsutus
Margaritopsis acuifolia 8
Palicourea guianensis
Notopleura guadaloupensis
Rudgea hostmanniana x
hypothetical
Figure 6. Presence of marginal, preformed germination slits (PGSs) on pyrenes, optimised on the same tree as that of
fig. 5. Branches leading to polymorphic terminals have been marked as equivocal.
183

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Optimisation (fig. 7) shows that the pattern of abaxial ridging is largely equivocal over the tree,
indicating that this character is of limited use for diagnosing major groups. The use of the charac-
ter is also burdened by coding problems, because there are quite a number of polymorphic taxa and
because ridges vary quite gradually from high and conspicuous over inconspicuous and incomplete
to none. Nevertheless it may be concluded that presence of many, distinct and complete abaxial
ridges is strong evidence that a taxon belongs in Psychotria s. str., whereas absence of these fea-
tures is uninformative.
Optimisation (fig. 8) fails to decide whether presence of an SCP is a plesiomorphic or apomorphic

trait in Psychotrieae. However, the state is unequivocally absent in the Palicourea complex ances-
tor and present in the Psychotria complex ancestor. Presence of an SCP is thus a diagnostic of the
Psychotria complex rather than of Psychotria s. str., as presumed by Petit (1964). The fact that this
character worked so well for the African Psychotrieae studied by Petit is simply that no Pacific
clade members are present in Africa.
A ruminate endosperm was emphasised by Bremekamp (1934, 1961, 1963) as one of the main
defining characters of Mapouria (i.e., Psychotria s. str.). He did not include in his concept of rumi-
nate those seeds that only have one median adaxial testa invagination and, in 1963, he segregated
such species as the genus Apomuria. Petit (1964) did not give the same weight to rumination, not-
ing that every conceivable intermediate exists between strongly ruminate over adaxially invaginat-
ed to completely entire endosperm. Piesschaert (2001) has shown that the variability found among
Africari species is also found among Neotropical ones.
Continuous variation creates difficulties for coding of rumination characters. In order to be able
to test Bremekamp's (1963) implicit hypothesis that presence of only a single median and adaxial
testa invagination is a characteristic independent of "true" rumination, this was coded as a separate
state (tab. 2). Among the species of the present sample, there were no difficulties making a distinc-
tion between this and presence of "true" rumination, i.e., testa folded in more than one place and inde-
pendent of pyrene ridges. However, among species with true rumination no consistent further subdi-
vision could be made according to amount or details of location. In a couple of cases Piesschaert says
that it is dubious if any rumination is present at all. These were coded as missing data.
As optimised (fig. 9), it appears that rumination is primitively absent in Psychotrieae. It is fur-
thermore primitively absent in the Pacific clade and the Palicourea complex. The ancestral state of
Psychotria s. str. is "truly" ruminate. Optimisation further confirms Petit's notion that presence of
rumination is a quite unreliable character for recognition of major groups in Psychotrieae and
rejects Bremekamp's idea that presence of an single adaxial invagination characterises a natural
group. Although resolution of the tree is too poor to reveal much detail of character transformation,
it rather appears that a single ventral invagination has been repeatedly derived from "true" rumina-
tion. From this point of view it seems justified to merge these two states into one, "rumination pre-
sent". This is not to deny that detailed patterns of rumination may still be characteristic of smaller
natural groups.
Johansson (1992) examined pollen of six species of the Pacific clade (as found in the present analy-
sis and the ITS tree of Nepokroeff et al. 1999) and 23 species of Psychotria s. str. All the species
of the Pacific clade were found to have double apertures (colporate), although in P. grandiflora (the
only species of his type XIII) they were more often simple (3-5-colpate). These species were
referred to five different pollen types, which means that there is considerable variation with respect
184

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
ridges on abaxial pyrene surface Hydnophytum formicarium

[" 1 (0) none Myrmecodia tuberosa
(1) 1-3 1Squamellaria imberbis
S (2) x4 Calycosia lageniformis
equivocal/polymorphic Psychotria brackenridgei
(possible states on branches) Psychotria brevicalyx
Psychotria turbinata -
Psychotria unicarinata 0
Amarcarpus kochii o
Dolianthus vaccinioides ".
I Psychotria chrysantha O
Psychotria gyrulosa
Psychotria mariana
Psychotria brownei
Psychotria
emestii "
Psychotria viridis "-
Psychotria horizontalis
Psychotria flava =r W
o0 grand
Psychotria
Psychotria peduncularis x
Psychotria nervosa -o
1 Psychotria calva -o C.
Psychotria heterosticta
Psychotria kimuenzae =r
Psychotriaasiatica
Psychotria punctata= 0"
Psychotria auxopoda
Psychotria capensis -
Psychotria eminiana
Psychotria remota
Psychotria djumaensis =
Psychotria orophila
Psychotria daphnoides '
Psychotria sarmentosa o
Psychotriarupicola
Psychotria tephrosantha -
Carapichea guianensis 0-T
Chassalia subochreata -
Geophila repens o
Hymenocoleus hirsutus
MI argaritopsis acuifolia 0
Palicourea guianensis 3
Rudgea hostmanniana I
hypothetical
Figure 7. Ridge pattern on abaxial side of pyrenes, optimised on the same tree as that of fig. 5. Branches leading to
polymorphic terminals have been marked as equivocal. Figures inside boxes on equivocal branches represent possible
character states.
185

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Syst. Geogr. P1. 72 (2002)
soluble seed coat pigment Hydnophytum formicarium
'I absent Myrmecodia tuberosa

pSquamellaria imberbis
present Calycosia lageniformis
equivocal/data missing Psychotria brackenridgei
Psychotria
Psychotria turbinata
unicarinata -o
Psychotria sp. "2489" 0
Amarcarpus chrysanthakochii
Psychotria D .
Psychotria gyrulosa
Psychotria mariana
Psychotria brownei
Psychotria carthagenensis o
Psychotria emestii
Psychotria viridis - 0
Psychotria horizontalis 0
Psychotria flava o
Psychotria nervosa
Psychotria calva 0
Psychotria heterosticta 0I
Psychotria kimuenzae o
Psychotria punctata
Psychotria asiatica =
Psychotria auxopoda
Psychotria capensis
Psychotria eminiana 3
Psychotria mapourioides O
Psychotria remota
Psychotria orophila
Psychotria loniceroides 0
Psychotria sarmentosa 0
Psychotria rupicola
IChassalia
Carapichea guianensis -
subochreata
Geophila repens 8
I Hymenocoleus hirsutus
I Margaritopsis acuifolia )
I Palicourea guianensis 3
Notopleura
Rudgea guadaloupensis -
hostmannianax
.hypcothetical
Figure 8. Presence of ethanol-soluble

terminals with unknown or un
186

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
endosperm Hydnophytum formicarium

[ (0) non-ruminate Myrmecodia tuberosa
(1) with adaxial, T-shaped intrusion Squamellaria imberbis
(2) ? distinctly, typically ruminate Calycosia lageniformis
equivocal/polymorphic (possible Psychotria brackenridgei
states on branches), or data missing Psychotria brevicalyx
Psychotria turbinata "a
Psychotria unicarinata 0
Psychotria sp. "2489" o
I-Amarcarpus kochii o
Dolianthus vaccinioides CL
Psychotria
Psychotria chrysantha
confertiloba "
Psychotria gyrulosa
Psychotria mariana
Streblosa myrocarpa
Psychotria brownei
Psychotria carthagenensis "o

Psychotria emestii ,
Psychotria
Psychotria viridis
horizontalis ' -U "
Psychotria flava '
Psychotria peduncularis a
Psychotria vogeliana x
Psychotria nervosa
Psychotria calva , -
Psychotria heterosticta 2 0
Psychotria kimuenzae P
Psychotria punctata
Psychotria asiatica. =
Psychotria auxopoda
Psychotria capensis
Psychotria eminiana .
Psychotria mapourioides o
Psychotria
remota= .
Psychotria orophila
Psychotria daphnoides .
Psychotria sarmentosa a
Psychotria rupicola
I Carapichea guianensis
Geophila repens
I Hymenocoleus hirsutus
I Margaritopsis acuifolia
Palicourea guianensis 8
Rudgea hostmanniana ,
-hypothetical
Figure 9. Presence and type of endo

terminals for which the state is un
branches represent possib
187

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
to exine sculpture. All the species of Psychotria s. str. except one were found to have simple aper-
tures [3-4 (5)-colpate or 2-porate]. The exception was P. limonensis, which according to Johansson
has inaperturate pollen. The identification is dubious, however, because Johansson's specimen
came from an area where R limonensis is otherwise not reliably recorded (C.M. Taylor, pers.
comm.). 17 of the 23 species of Psychotria s. str. were referred to the same pollen type (XVI),
which suggests fairly homogeneous pollen in this clade. Although sampling is still sparse with
respect to detailed pollen data, Johanson's observations suggest, when compared to recent
sequence-based phylogenies, that the Pacific clade and Psychotria s. str. differ with respect to aper-
ture structure.
4.3 Psychotria sensu stricto

Nepokroeff et al. (1999) suggested that "the genus Psychotria should be redefined to include m
bers of subgen. Psychotria, subgen. Tetramerae, and members of subtribe Hydnophytinae", i
they suggested exclusion from Psychotria of such species found to belong in the Palicou
complex, but inclusion of all species of the Psychotria complex, including the myrmecophile
They also say that "a new description of the genus should include lack of preformed germinati
slits on the pyrenes". This is consistent with the way PGS characters are optimised on their tr
but, as discussed above, their optimisation is faulty. They offered no detailed arguments in fav
of their relatively broad circumscription of Psychotria, their focus being rather on segregating
ments of the Palicourea complex. However, it may be assumed that their main arguments were
the one hand their finding (with which the present analysis agrees) that the Psychotria complex
monophyletic and on the other hand the assumption (proven wrong) that this complex is charac
isable morphologically by absence of PGSs. Although the Psychotria complex is indeed morph
logically characterisable (by caducous stipules and presence of an SCP), I find this circumscripti
inconvenient. The main reason for this is that it entails synonymisation of the myrmecophiles und
Psychotria. Besides the the fact that this group is strongly aberrant morphologically, a very lo
tradition speaks against such a drastic rearrangement. Nor does it appear necessary and it may
even be helpful in creating morphologically characterisable taxa. As will be further discussed
below, there are considerable problems within the Pacific clade, but to merge the Pacific clade w
Psychotria would solve such problems only in a nomenclatural sense. I would suggest th
Psychotria is restricted to correspond to the clade labelled "Psychotria s. str." in fig. 1, i.e., excl
ing members of the Pacific clade. Monophyly of this group is still quite strongly supported, w
89% jackknife support in the rpsl6 analysis, 96% jackknife support in the combined analysis, an
81% in the ITS analysis of Nepokroeff et al.
Based on the observation that marginal PGSs are present in Psychotria asiatica, A. Davis an
D. Bridson (pers. comm.) have suggested a much narrower circumscription for Psychotria, nam
ly to include only species with marginal PGSs. This does not seem feasible in view of the prese
results. First, this character seems to be variable even within R asiatica. Second, such a sample
species would clearly be paraphyletic, including most members of the Pacific clade (except so
myrmecophiles) and some, though far from all, species of the Psychotria s. str. clade. Quite cle
ly, whether narrowly or broadly circumscribed, the genus Psychotria cannot be at the same ti
monophyletic and diagnosed by any single feature of pyrene morphology.
With the circumscription proposed here, Psychotria s. str. is characterised morphologicall
mainly by the combination of absence of PGSs, presence of multiple abaxial ridges on the pyren
and pollen with simple apertures. Thus circumscribed, it corresponds conceptually to Psychotri
Petit's (1964) sense, Psychotria subgen. Psychotria in Steyermark's (1972) sense, and Mapouria
the sense of Mtiller (1881) and Bremekamp (1934, 1961). Besides the type species of Psychotri
188

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
(P. asiatica), the present sample includes the type species of both Mapouria (P. mapourioides) and
Psychotrophum (P brownei). Although the type species was not sequenced, two species referrable
to Cremocarpon are also included in the present sample. This may be taken to support Piesschaert's
(2001) suggestion that Cremocarpon should be synonymised under Psychotria. Based on morpho-
logical evidence, Piesschaert argues further that Pyragra should go with Cremocarpon. No species
referrable to Pyragra was included in the present sample, however.
Only one of the species included by Bremekamp (1963) in Apomuria is included in the present
sample, viz. P. punctata. This is nested in subclade II of the Psychotria s. str. clade. Apomuria was
segregated from Mapouria mainly because the endosperm is not "truly" ruminate but has only one
adaxial testa invagination. Several more species with a single testa invagination are included in the
present sample, although they were not treated by Bremekamp: P. auxopoda, P. brevipuberula, P
comperei, P. gabonica, and P. lucens. Although some of these may be related, there is strong sup-
port that all of them are not and that all of them have close relatives with a "truly" ruminate
endosperm. Thus, this character is is homoplasious and the genus Apomuria appears polyphyletic
as concieved by Bremekamp. Since no other characters were given for the genus, I feel confident
to conclude that it should be synonymised under Psychotria.
Results of the present analysis do not suggest a simple way in which to subdivide Psychotria.
Although a fair number of strongly supported species groups are identified, the majority of species
are found at two large polytomies.
The only detailed infrageneric classification of Psychotria in the present sense is that of Petit
(1964, 1966) and it takes only Africcan representatives into account. Petit's segregation of subgenus
Tetramerae is essentially supported by present results in as much as circumscription is concerned.
With few exceptions species referred to Tetramerae by Petit are found in subclade II of Psychotria
s. str. (fig. 3) in which is found only one species (P. butayei, listed by Petit among species of uncer-
tain position) that do not have bacterial nodules. Subclade II seems to be characterized morpho-
logically also by having non-caducous stipules (fig. 5). However, although Tetramerae is thus
monophyletic in essentially the circumscription given by Petit, to recognize it as a subgenus leaves
subgen. Psychotria paraphyletic.
Three of the species referred by Petit to sect. Flaviflorae appear in the moderately supported
monophyletic group in clade III (fig. 3, "k") and the other two species (P. albicaulis, P. gabonica)
of this section are attached to the same polytomy. Two species referred to sect. Confertiflorae form
a moderately supported group (fig. 3, "1") but the third species (P. comperei) of the same section is
grouped with ambiguous support with P. succulenta, which Petit referred to sect. Holostipulatae.
The other three species (P. djumaensis, P. orophila, P. zombamontana) referred to sect.
Holostipulatae all appear at the polytomy of subclade III (fig. 3, "m"). In conclusion, although
Petit's sections of subgen. Psychotria are only partly supported as monophyletic, their monophyly
is not strongly refuted by the present analysis.
Well-supported smaller groups found within Psychotria are essentially consistent with geo-
graphical distribution. Thus, all species of the groups a, c, d, and j (fig. 3) are Neotropical. All
species of the groups b, k, 1, m, n and clade II are African, those of group i are East Asian or
Indonesian, those of group o are Indomalaysian and those of group p are Fijian. Furthermore,
species of subclade IV, which includes groups o and p are all from Asia and Australasia.
These results suggest that a subdivision of Psychotria into smaller monophyletic groups, possi-
bly recognizable as sections, may well be possible although this analysis does not suggest precise-
ly how this should be done. Such a subdivision will obviously need to depend on morphological
characters and will have to take into account a much broader sample of species than the present one.
189

84.115.220f:ffff:ffff:ffff on Thu, 01 Jan 1976 12:34:56 UTC
Syst. Geogr. PL. 72 (2002)
Considering that both the present data set and the ITS data of Nepokroeff et al. (1999) fail to pro-
vide trees with detailed and well supported resolution, it does not seem likely that sequence data
will achieve such a goal, unless quite long sequences from several different loci are employed,
preferably in combination with morphological data.
Psychotria s. str., as represented in the present sample, includes numerous species from the
Neotropics and Africa, several from Madagascar, a few from SE Asia, two from Sumatra, one from
Borneo, one from New Caledonia, two from Fiji, and four from Australia. To these may be added
R manillensis DC. from the Philippines and R decorifolia Moore from New Guinea, which
Nepokroeff et al. (1999) found to be nested in their corresponding clade. However, it is a striking
feature of the present analysis as well as of that of Nepokroeff et al., that although a large number
of "Psychotria" species are recognized in tropical parts of Asia, Australasia and the Pacific, most
of those sampled for molecular studies have turned out to belong in the Pacific clade. The impres-
sion that true Psychotria are poorly represented in the Pacific becomes even stronger when adding
the circumstance that quite a number of "Psychotria" species from this area have been shown
(Andersson 2001) to belong in the Palicourea complex. It is also fully consistent with Piesschaert's
(2001) pyrene data: most species he investigated from Indonesia, New Guinea, the Philippines and
Fiji have pyrenes with distinct marginal PGSs and an abaxial surface with few and/or incomplete
ridges, which suggests that they belong in the Pacific clade. In conclusion, Psychotria s. str. appears
to be much richer in species in Africa and the Neotropics than it is in Asia and the Pacific. The fair
number of species in Australia may be due to secondary radiation.
4.4 The Pacific clade

Having circumscribed Psychotria as done above, the question remains how to treat taxonomic
the Pacific clade. Since it is poorly resolved, this analysis leaves open several options. My p
ry concern is that larger, strongly supported clades should be formally recognized, if possibl
concerns in the first place the myrmecophiles and the Calycosia clade, but recognising these
nomically leads to consequences for the rest of the groups as well.
Hydnophytum. Monophyly of the myrmecophiles is strongly supported in this analysis as w

in those of Andersson & Rova (1999) and Nepokroeff et al. (1999). The group was subdivided
five genera by Huxley & Jebb (1991). From the present analysis it is clear that Hydnophyt
paraphyletic as presently circumscribed. From the analysis of Nepokroeff et al. (1999), based
largely different taxon sample, it is clear that also Myrmecodia is paraphyletic. In my view, th
solution to this problem is to merge all myrmecophiles in one genus. The other option, to sub
the group into monophyletic genera appears to entail further splitting of both Hydnophytu
Myrmecodia, which, I feel, is not in the interest of nomenclatural stability. The fairly small
ber of species in this clade is a strong argument against proliferation of generic names. Si
Hydnophytum and Myrmecodia have equal priority, having been published in the same publica
a choice must be made between the two names. I herewith propose that Hydnophytum is use
the emended genus, since this is the choice that leads to the smallest number of new combin
and is thus the one that best serves the goal of nomenclatural stability.
Calycosia. Calycosia has been recorded from Fiji, the New Hebrides, Samoa, and the Sol
Islands (Darwin 1979). Its delimitation has varied considerably among authors and, conseque
records of its distribution vary as well.
190

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
The present sample includes two species, C. lageniformis and C. macrocyatha which were
referred to Calycosia by Fosberg (1942), Darwin (1979), and Smith & Darwin (1988). As circum-
scribed by these authors, Calycosia was characterised mainly by its strongly congested, involucrate
inflorescences. However, in the present analysis C. lageniformis and C. macrocyatha are included
in a strongly supported group, which also includes species that do not have involucrate heads and
are rather referrable to Psychotria in the sense of Smith and Darwin (1988). This Calycosia group
is internally totally unresolved because of very slight sequence divergence (fig. 2). The strong sup-
port suggests that the Calycosia group does merit recognition, but it is also evident that it is more
inclusive than presumed by Fosberg (1942), Darwin (1979), or Smith and Darwin (1988), and that
it must be differently diagnosed.
When originally described (Gray 1858), Calycosia included two species, C. petiolata and
C. sessilis. In Gray's original conception, Calycosia was distinguished mainly by an enlarged,
infundibular and caducous calyx. Somewhat later Gray (1860) added two more species to the
genus, but these do not possess the congested inflorescences. Gillespie (1930) transferred the two
species (C. lageniformis and C. petiolata) with congested inflorescences to "Uragoga L.", retain-
ing in Calycosia species with enlarged calyx but without congested inflorescences. He did not treat
the second of Gray's original species. As pointed out by Smith (1936), Gillespie changed the cir-
cumscription of Calycosia such that both original species were excluded. Smith seems to have
agreed with Gillespie's generic concepts, however. He lectotypified Calycosia by C. petiolata and
referred species with an enlarged calyx but lax, non-involucrate inflorescences to a new genus,
Calycodendron A.C.Sm. Fosberg (1942) included Calycodendron in Psychotria, as did Darwin
(1979) and Smith & Darwin (1988).
The species associated with Calycosia in the present sample are not consistently characterised
by an involucrate inflorescence, nor by a conspicuously enlarged calyx. However, Darwin (1979)
found that pollen grains of all species of Calycosia have a thin exine and are not "very resistent to
acetolysis". Johansson (1992) found the same thing for the three Fijian species Psychotria
gibbsieae S.Moore, P gillespieana A.C.Sm., and P. pubiflora (A.Gray) A.C.Sm., which he refer-
red to pollen type VII. Four more Fijian species were referred to the same pollen type, but
those seem to have thicker exine. Johansson distinguished this pollen type mainly on comparatively
short colpi and small to very small lumina. None but these Fijian species were referred to this pollen
type.
All species of the Calycosia clade, except the unidentified "2489" have none-ruminate
endosperm. In the exceptional species rumination consists of few and very shallow invaginations
confined to the adaxial side. Piesschaert (2001) examined pyrenes of P. gibbsieae, R gillespieana,
P. pubiflora and P. storckii Seem., all of which were found by Johansson (1992) to have pollen of
type VII. All of these species were found to have non-ruminate endosperm. It may be, therefore,
that non-ruminate or weakly ruminate endosperm is a character that would serve to distinguish
Calycosia from Straussia.
According to Piesschaert (2001), pyrenes with long marginal PGSs and non-ruminate
endosperm occur in R pickeringii A.Gray and R platycocca A.Gray, two of the four species referred
by Gray (1858) to Psychotria subgen. Piptilema. None of these species has been sequenced, nor
were they examined by Johansson (1992). In spite of this it appears justified, based on pyrene mor-
phology, to let this name go into synonymy under Calycosia.
Most of the presently sampled species of the Calycosia clade have no or very scarce SCP (fig.
8). However, Piesschaert (2001) found SCP in P brackenridgei as well as several of the non-
sequenced species discussed above. It may well be that the impression given by fig. 8, that
Calycosia is diagnosed by absence of SCP, is due to a sampling artefact.
191

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
In conclusion, this analysis suggests that the type of Calycosia belongs in a group that is strong-
ly supported as monophyletic. This speaks for continued recognition of the genus. It is clear, how-
ever, that neither a congested involucrate inflorescence, nor a conspicuously enlarged calyx serve to
diagnose it. It is possible, but far from clear, that it may be characterised by non-ruminate endosperm
and the peculiar pollen type VII of Johansson. The question clearly needs a more detailed study, par-
ticularly of pollen morphology in a broader sample of Fijian "Psychotria" species.
Amaracarpus and Dolianthus. Amaracarpus is supposed to be distributed from Malesia eastward

to Micronesia and Fiji (Darwin 1979) and Dolianthus is endemic to the Highlands of New Guinea
(Davis & Bridson 2001).
The present analysis does not give any strong indications concerning the status of Amaracarpus.
The two sampled species form a moderately supported group together with P. insularum, but its
relationships to Calycosia and Straussia are not revealed. Nepokroeff et al. (1999) included P. insu-
larum in their ITS analysis, based on the same accession used here. They found it nested among
Straussia species, but relationships in this part of their tree were unsupported, except for one weak-
ly supported branch. Nepokroeff et al. also included one unnamed ["Amaracarpus sp. = Psychotria
hombroniana (Baillon) Fosberg"] species of Amaracarpus, which they found to form a strongly
supported group together with an unnamed ("Tinian Isl.") species of Psychotria. This group formed
the sister group of an unsupported cluster comprising mainly Straussia species. Davis & Bridson
(2001) have argued that Amaracarpus is distinct based on its dorsiventrally organized branches
with leaves in two rows, pseudolateral inflorescences, and tetramerous flowers. These characters
are ceratainly not unique and Piesschaert (2001) found no unique features in their pyrene mor-
phology. If it should be synonymised, it is under Straussia rather than Psychotria.
Dolianthus was synonymised under Amaracarpus by van Royen (1973), but recognized by
Davis & Bridson (2001). Davis & Bridson argued that Dolianthus is distinct from both
Amaracarpus and Pacific "Psychotria" by its large bi- or multicoloured flowers with stamens
inserted above the ring of hairs. The present analysis is inconclusive in that the only included
species of Dolianthus does not show any supported relationships to other members of the Pacific
clade. This may be taken as a weak argument not to include Dolianthus in Amaracarpus.
Straussia. The genus Straussia was established by Gray (1858), when raising de Candolle's (1830)
Coffea sect. Straussia to generic rank. Gray, like de Candolle, included only Hawaiian species.
Although Farr et al. (1979) state that a generic type has not been designated, the genus should in
fact be considered lectotypified by Schumann's (1891: 112) statement that "S. kaduana (Cham.) A.
Gr. ist die typische Art". Straussia was accepted as a distinct genus by Hooker (1873) and
Schumann (1891), who both included only Hawaiian species.
Fosberg (1964), who treated all Hawaiian Psychotrieae under Psychotria, reduced Straussia to
a section of that genus. He recognized ten Hawaiian species in section Straussia, four of them
included in the present sample: P. hathewayi, P. kaduana, P. mariniana and P. psychotrioides
(Heller) Fosb. He also recognized two other sections among Hawaiian Psychotria, P sect.
Psychotria and R sect. Pelagomapouria. To sect. Psychotria he referred only P. insularum and to
sect. Pelagomapouria two species, P. grandiflora and P. hexandra. Straussia was synonymized also
by Sohmer (1977) and Darwin (1979). Darwin says that "the differences between [Straussia and
Psychotria] seem trivial when compared to the variation which must be tolerated in Psychotria in
the strictest sense".
Mainly as a matter of convenience, I prefer to provisionally maintain Straussia as separate

genus. When doing so, I prefer to include all "Psychotria" species of the Pacific clade that are not
192

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
strongly supported as members of the Calycosia clade. Admittedly, there is nothing in the present
analysis suggesting that this is a monophyletic group. The ITS analysis of Nepokroeff et al. (1999)
shows Straussia as monophyletic (if including sect. Pelagomapouria), but without jackknife support.
The present analysis seems to suggest that it is rather a complex of smaller groups, which together
form a paraphyletic assemblage in which Calycosia and Hydnophytum (in the wide sense advocat-
ed above) are nested. If this proves to be true, then Straussia should be further subdivided into nat-
ural groups. At present, however, neither the taxon sample, nor the amount of sequence data are suf-
ficient to decide how this should be done. Pending more and better data, I prefer to take a moder-
ately conservative point. To recognize Straussia makes the problem of its delimitation and natural-
ness evident, whereas sinking it into Psychotria will only sweep the problems under the carpet.
As far as data are presently available, Straussia appears to be diagnosable morphologically by
pyrenes that are smooth to knobbly abaxially, or provided with a single median crest, which have
usually long marginal PGSs, and by an endosperm which is ? ruminate, at least insofar as having
a ? deep adaxial intrusion. Presence of long and distinct marginal PGSs would serve to distinguish
Straussia from Psychotria. A ruminate endosperm would serve to distinguish Straussia from
Amaracarpus and Dolianthus, which both have entire endosperm (Davis & Bridson 2001, pers.
comm.). As discussed above, Calycosia may be morphologically distinct from Straussia in pollen
type as well as absence of endosperm rumination.
The name Straussia has never been applied to species from outside Hawaii. It seems clear, how-
ever, that with the circumscription proposed here, this complex has a wider distribution in the
western Pacific and Indian Ocean. Psychotria confertiloba from Fiji would belong here based on
how it comes out in the present analysis and on its possession of long marginal PGSs in combina-
tion with ventrally ruminate endosperm (Piesschaert 2001). Psychotria chrysantha from New
Guinea, R gyrulosa from Borneo and P. mariana from the Mariana Islands belong in Straussia
based on their position in the present analysis as well as their carpological features. Piesschaert
reports the same combination of carpological characters in one species from Malaysia, two from
Sri Lanka, some five from Java and Borneo, about a dozen from New Guinea, at least four from the
Solomon Islands, several (more) from Fiji, some five from the Philippines, and at least one from
Australia. Among the species which, based on Piesschaert's data, are likely to belong here are
R eumorphanthus Fosb., the type species of Eumorphanthus A.C.Sm., and P. nigra (Gaertn.)
Alston, the type species of Grumilea Gaertn. Grumilea, which has priority over Straussia, is fur-
ther discussed below.
Streblosa. Only a single species of Streblosa was included in the present study, which consequent-
ly cannot evaluate whether the genus is monophyletic. It appears in this analysis as the sister to all
other members of the Pacific clade, but this relationship is quite weakly supported. Its membership
in the Pacific clade is strongly supported, however, and this is enough to reject it's synonymisation
under Psychotria (Backer & Backhuizen van den Brink 1965). As argued by Piesschaert (2001)
there are several morphological characters, which seem to imply that the genus is monophyletic and
diagnosable: herbaceous habit, reticulate leaf venation, pseudoaxillary, boragioid inflorescences,
small fruits, absence of PGSs and presence of a ventral cavity on the adaxial pyrene surface. In
absence of strong molecular evidence to the contrary, I feel that Streblosa should be recognized.
Bremer & Manen (2000) say that "it is not obvious from morphology that [Margaritopsis,
Readea and Streblosa] belong [in Psychotrieae]". They do not specify the reasons of their doubts,
however. In fact, morphology is fully consistent with the placement of Streblosa in Psychotrieae:
presence of raphides, ovules solitary in locules and basally attached, and drupaceous fruit
(Bremekamp 1947).
193

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
4.5 Names of unclear or uncertain application

Cephaelis. The name Cephaelis Sw. has been applied to Neotropical species as well as to African
and Asiatic ones. This broad generic concept was based on congested, involucrate inflorescences.
Steyermark (1972) included all Neotropical "Cephaelis" species treated by him in Psychotria
subgen. Heteropsychoria. The type species, Cephaelis muscosa (Jacq.) Sw. [= Psychotria muscosa
(Jacq.) Steyerm.] has been shown by sequence data (Nepokroeff et al. 1999, Andersson 2001) to
belong in the Palicourea complex and, from a nomenclatural point of view, Cephaelis is a synonym
of Palicourea Aubl. With very few exceptions all Neotropical species referred to Cephaelis belong
in Palicourea s. lat., but three of them belong in Carapichea (Andersson 2002).
Schnell (1957, 1960) argued that African species referred to Cephaelis were more closely relat-
ed to African Psychotria than to Neotropical "Cephaelis". Petit (1964) agreed with this and referred
them to Psychotria. Two of them were included in the present sample, R psychotrioides (DC.)
Roberty (which includes Cephaelis congensis Hiern; Petit 1964) and R vogeliana (which includes
C. cornuta Hiern and C. talbotii Wernh.; Petit 1964). Present results are consistent with the con-
clusions of Petit and Schnell.
The Asiatic species Cephaelis stipulacea Bl. was transferred by Piesschaert et al. (2001) to
Chassalia. This is supported by sequence data (Andersson unpubl.) in the sense that C. stipulacea
clearly belongs in the Palicourea complex and is related to Chassalia, Geophila and
Hymenocoleus. However, sequence data do not presently support monophyly of Chassalia
(Andersson 2001).
Chaetostachydium. Chaetostachydium Airy Shaw is presumed to be a genus of three species, all

endemic to New Guinea (Ridsdale 1975). Its salient features are a peculiar habit with large leaves
and densely spicate, pendent inflorescences (Piesschaert 2001). Ridsdale (1975) suggested that it
might not be sufficiently distinct from Psychotria to warrant generic distinction, a view with which
Piesschaert (2001) disagreed. None of the three species has been sequenced but Piesschaert (2001)
described the pyrenes of the type species, C. versteegii (Valeton) Airy Shaw. They have a slightly
rugulose, ecostate abaxial surface and long and distinct marginal PGSs, the testa lacks an SCP and
the endosperm is non-ruminate. The stipules are caducous, judging from Valeton's illustration
reproduced by Piesschaert (2001). By these characters it would fit in the Pacific clade rather than
in Psychotria s. str. and the non-ruminate endosperm suggests affinity to Calycosia. Its combina-
tion of morphological features is unique, however, and its precise relationships remain uncertain.
Coelopyrena. Only a single Malesian species, C. salicifolia Valeton, has been referred to
Coelopyrena. It has not been sequenced, nor was it examined by Piesschaert (2001). However, the
ilustration of Valeton's original description (reproduced by Piesschaert 2001) is fairly detailed and
Piesschaert examined two "Psychotria" species which appear closely similar to C. salicifolia.
According to Valeton, the stipules of C. salicifolia are caducous and judging from his illustration
the seeds have a non-ruminate endosperm. The pyrenes examined by Piesschaert had two short but
distinct marginal PGSs, an SCP was present and the endosperm is ruminate, at least in the upper
part of the seeds. These features suggest that Coelopyrena belongs in the Psychotria complex. At
least the two species examined by Piesschaert appear related to Straussia. The fact that the
endosperm appears non-ruminate on Valeton's illustration may be because the section was made at
a level where rumination does not show.
Gillespiea. Gillespiea A.C.Sm. was originally described (Smith 1936) based on a single species
from Fiji and has remained monotypic (Darwin 1979, Smith & Darwin 1988). It was considered
194

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
close to Calycosia because of its large (but not widened) calyx. Darwin (1979) and Smith & Darwin
(1988), on the other hand, suggested an affinity to Readea, which Andersson (2001) included in
Margaritopsis of the Palicourea complex. The only species has not been sequenced, nor were its
fruits available to Piesschaert (2001). The stipules were described by Smith (1936) as soon
deciduous, which speaks for Smith's original hypothesis in as much as it implies a placement in the
Psychotria complex. Without access to mature fruits, or to material suitable for sequencing, the
case remains open, however.
Grumilea. The name Grumilea was coined originally (Gaertner 1789) for a single species
[Grumilea nigra Gaertn. = Psychotria nigra (Gaertn.) Alston] from Sri Lanka, but it has been wide-
ly applied to Psychotrieae species of Africa and Asia. The genus was recognized as distinct by
Schumann (1891), but synonymised under Mapouria by Bremekamp (1961) and under Psychotria
by Petit (1964). Even if using different names, Bremekamp and Petit reached essentially the same
conclusion, since Mapouria sensu Bremekamp and Psychotria sensu Petit are conceptually more
or less the same. Schumann distinguished Grumilea from Mapouria mainly on the basis of
Mapouria having "seeds" with flat adaxial surface whereas Grumilea was said to have "seeds" with
a narrow furrow. Bremekamp argued, correctly, that Schumann's distinction was based on faulty
observations and that, because the two genera agree in having caducous stipules and ruminate
endosperm, they are essentially the same. However, as shown by the present results (fig. 5, 9),
caducous stipules and ruminate endosperm occur both in Psychotria s. str. and the Pacific clade.
The present analysis gives support for Bremekamp's and Petit's conclusions as far as African
species are concerned. Seven African species of the sample have been referred to Grumilea:
P. capensis, R djumaensis, R kirkii, P lauracea, R psychotrioides (DC.) Roberty, R riparia, and
P succulenta. All of these are included in Psychotria s. str. However, the type of Grumilea was not
sequenced. According to Piesschaert (2001), this species has long and distinct marginal PGSs in
addition to a shorter mediodorsal one and the adaxial pyrene surface has a single median ridge. The
endosperm is deeply ruminate on both adaxial and abaxial side. This combination of characters
does not occur in any sequenced species of Psychotria s. str., but it does occur among species of
the Pacific clade. Based on pyrene morphology it appears that Grumilea should possibly be applied
to the group of species discussed above as Straussia.
Hedstromia. The genus Hedstromia was established by Smith (1936) to accommodate one species
thought to differ from other Fijian "Psychotria" in having free petals. Darwin (1979) and Smith &
Darwin (1988) also noted that the stipules differ from those of other Fijian "Psychotria" in being
free nearly to the base. As suggested by Piesschaert (2001), the "free" petals are likely to be a mis-
interpretation of a very short corolla tube. Furthermore, among other Psychotrieae, stipules may
vary from nearly free to completely fused and calyptrate even among closely related species. This
is hardly more than a specific character. The type and only species of Hedstromia has not been
sequenced. Piesschaert (2001) described the pyrenes as having marginal PGSs and found an SCP
present. He could not observe the endosperm, however, since the seeds were not fully mature.
Caducous stipules, marginal PGSs and SCP clearly suggest that Hedstromia belongs in the Pacific
clade of the Psychotria complex, but in the absence of information on endosperm rumination it can-
not be decided if it belongs in Calycosia or Straussia.
Lecariocalyx. Lecariocalyx Bremek. was distinguished mainly because of its rotate calyx and
accrescent fruit (Bremekamp 1940). It seems to be known only from the type collection (from
Borneo) of its only species and no material was available to Piesschaert (2001), nor has it been
195

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
sequenced. Bremekamp stated that the endosperm is ruminate and suggested a relationship to
Grumilea or Peripeplus. He did not mention whether or not PGSs are present. The presence of a
ruminate endosperm clearly suggests that this species is a member of the Psychotria complex and
its geographical origin may be taken as an indication that it belongs in the Pacific clade. Clearly,
no further conclusions can be drawn in the absence of sequence data and detailed information on
pyrene morphology.
Mapouria. A wide usage of the name Mapouria seems to date back to MUiller (1886) and Schumann
(1891). For reasons that were not explained, they applied the name Psychotria to species with an
adaxially furrowed seed, leaving the ones with an adaxially flat seed in Mapouria. With Miller,
who only treated Brazilian species, Psychotria came to correspond quite closely to Palicourea s.
lat., whereas Mapouria corresponded to Psychotria s. str. Schumann extended these concepts to a
worldwide scale. The name Mapouria was used in approximately this sense also by Bremekamp in
1934 and 1961, but in 1963 he restricted Mapouria to comprise only species with "truly" ruminate
seeds, referring the ones with only a single median testa invagination to Apomuria.
There is some confusion about application of the name Mapouria guianensis Aubl. It was
synonymised under Psychotria carthagenensis by Zappi & Nunes (2000). According to Dr. C.M.
Taylor (pers. comm.), Aublet's type belongs to the species that I have identified as P ernestii.
Whichever is correct, the synonymisation of Mapouria under Psychotria is unaffected.
Peripeplus. Petit (1964) accepted the African genus Peripeplus as distinct from Psychotria, main-
ly, it appears, because he found the pyrenes to "probably" possess four dorsal PGSs. Petit also
reported that the endosperm is ruminate. From his key it appears that the testa lacks an SCP. No
pyrenes were available to Piesschaert and none of the two species has been sequenced. The rela-
tionships of this genus remain obscure. The lateral inflorescences, presence of dorsal PGSs and
absence of SCP speak against its inclusion in Psychotria. On the other hand, absence of marginal
PGSs and ruminate endosperm as well as its geographical range speak for such inclusion. The
unusual features may well be autapomorphies and thus uninformative. A final judgement would,
however, need confirmation from sequence data and, besides, Petit's observations need to be con-
firmed.
Psathura. Psathura is considered (Bremekamp 1963) as a genus of ca. 8 species from Madagascar
and the Mascarenes. No species has been sequenced and Piesschaert (2001) did not see any mate-
rial. Bremekamp (1963) described the stipules as soon marcescent (rather than deciduous) and the
endosperm as non-ruminate. This combination of features is reminiscent of the Palicourea complex
rather than the Psychotria complex.
Uragoga. Petit (1964) designated the Neotropical Callicocca ipecacuanha Brot. as lectotype of
Uragoga. Andersson (2002) showed that this species belongs in the Palicourea complex and is
closely related to Carapichea guianensis Aubl. [= Psychotria ligularis (Rudge) Steyerm.]. Thus,
when Carapichea Aubl. is recognized as a distinct genus of the Palicourea complex, as Andersson
suggested it should, Uragoga becomes a synonym of Carapichea. The name Uragoga has been
widely and vaguely used, however. Sometimes, as with Kuntze (1891), a wide concept was based
in a refusal to accept the nomenclatural starting point. Then, Uragoga ("Ouragoga") was ascribed
to Linnaeus (1737) and given priority over Psychotrophum (Browne 1756) and Psychotria
(Linnaeus 1759). On the other hand, Schumann (1891) and De Wildeman (1936), who ascribed the
196

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
name to Linnaeus, used it as the correct name for a genus circumscribed approximately as
Cephaelis. Schumann listed, among others, Cephaelis and Carapichea as synonyms. This concept
is apparently behind Gillespie's (1930) transfer of two Calycosia species to Uragoga.
De Wildeman (1936) states, without explanation, that Cephaelis peduncularis Salisb. is the type
of Uragoga, but since this name was published only in 1806 it cannot be the type of a name
ascribed to Linnaeus. Nor can it be the type of Uragoga L. ex Baill., since it is not mentioned by
Baillon (1879), who, furthermore, made it quite clear that his concept is based on Linnaeus', i.e.
including Callicocca ipecacuanha Brot.
5 Conclusion: a synopsis of genera of the Psychotria complex
Amaracarpus Bl., Bijdr. 945 (1826-1827). Type: Amaracarpus pubescens Bl.

Stipules (always?) persistent. Pyrenes planoconvex, adaxial side smooth, often with short median
ridge at base, abaxial side ecostate or with 3-4 ? distinct ridges, with distinct marginal PGSs. Testa
dark but pigment not soluble in ethanol (only 1 unidentified species examined). Endosperm entire.
According to Piesschaert (2001) Amaracarpus is morphologically distinguishable by its dor-
siventral branches, pseudoaxillary inflorescences, and tetramerous corolla. According to Darwin
(1979) there are ca. 60 species in an area from Malesia to Fiji.
Calycosia A.Gray, Proc. Amer. Acad. Arts. Sci. 4: 47 (1858). Type: Calycosia petiolata A.Gray
(lectotype designated by Smith 1936: 153).
Psychotria sect. Piptilema A.Gray, Proc. Amer. Acad. Arts Sci. 4: 46 (1858). Type: Psychotria
pickeringii A.Gray (lectotype designated here).
Calycodendron A.C.Sm., Bernice P. Bishop Mus. Bull. 141: 155 (1936). Type: Calycosia pubiflora
A.Gray = Calycodendron pubiflorum (A.Gray) A.C.Sm.
Stipules caducous. Pyrenes planoconvex to triangular, adaxial side ? smooth, often with short medi-
an ridge at base, sometimes with shallow depressions on either side of crest, abaxial side usually
with 1-4 ? pronounced ridges in upper part, more rarely ecostate and bumpy, or ridged throughout,
with long and distinct, marginal PGSs. Testa with or without ethanol-soluble pigment. Endosperm
usually not ruminate, rarely with sparse and shallow invaginations on adaxial side.
Calycosia may comprise some 60 Fijian species, i.e. those referred by Smith & Darwin (1988)
to Calycosia and the informal Psychotria groups Brackenridgeae, Filipedes, Macrocalyces,
Pickeringieae and Turbinatae. In addition, Smith (1979) referred a small number of species from
Samoa and the Solomon Islands to this genus. Considering that Smith and Darwin (1988) circum-
scribed Calycosia more narrowly than is done here, the number of species may be larger in these
areas, but sequence data and detailed data on endosperm structure are to scarce to substantiate this.
Chaetostachydium Airy Shaw, Kew Bull. 18: 271 (1965). Chaetostachys Valeton, Nova Guinea 8:
495 (1911), non Benth. Type: Chaetostachys versteegii Valeton - Chaetostachydium verstegii
(Valeton) Airy Shaw.
Stipules caducous. Pyrenes concave-convex, adaxial surface concave, with ? distinct crest in centre
of cavity, abaxial surface ecostate, slightly rugulose, marginal PGSs present. Testa without soluble
pigment. Endosperm entire.
Ridsdale (1975) referred three species to Chaetostachydium, all from New Guinea.
197

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Coelopyrena Valeton, Icon. Bogor. tab. 279 (1909). Type: Coelopyrena salicifolia Valeton.
This may be a synonym of Straussia, but position is uncertain.
Dolianthus C.H.Wright, Kew Bull. 1899: 106 (1899). Type: Dolianthus vaccinioides C.H.Wright.
Stipules persistent or caducous. Pyrenes planoconvex, adaxial side smooth, often with an incon-
spicuous medial ridge, abaxial side ecostate or with 1-4 ? distinct ridges, with distinct marginal
PGSs. Presence of soluble testa pigment not studied. Endosperm entire.
According to Davis & Bridson (2001) Dolianthus is distinguishable by its large flowers with
stamens inserted on the corolla above (instead of within) the ring of hairs. It is endemic to the high-
lands of New Guinea and may comprise ca. 13 species (Davis & Bridson 2001).
Gillespiea A.C.Sm., Bernice P. Bishop Mus. Bull. 141: 158 (1936). Type: Gillespiea speciosa
A.C.Sm.
This may be a synonym of Calycosia, but in absence of sequence data and detailed data on polle
and pyrene morphology, its position remains uncertain.
Grumilea Gaertn., Fruct. 1: 138 (1788). Type: Grumilea nigra Gaertn.

Grumilea has commonly been synonymized under Psychotria, but may be the oldest name for
complex here treated as Straussia. Its position should be considered as doubtful.
Hedstromia A.C.Sm., Bernice P. Bishop Mus. Bull. 141: 146 (1936). Type: Hedstromia latifol
A.C.Sm.
Hedstromia is likely to be a synonym of Calycosia or Straussia, but in the absence of sequence da

and detailed information on pollen and pyrene morphology its position remains uncertain.
Hydnophytum Jack, Trans. Linn. Soc. London 14: 124 (1823). Type: Hydnophytum formicarium
Jack.
Myrmecodia Jack, Trans. Linn. Soc. London 14: 122 (1823). Type: Myrmecodia tuberosa Jack.
Myrmephytum Becc., Malesia 2: 92 (1884). Type: Myrmephytum selebicum Becc.
Myrmedoma Becc., Malesia 2: 94 (1884). Type: Myrmedoma arfakiana Becc.
Squamellaria Becc., Malesia 2: 228 (1886). Type: Myrmecodia imberbis A.Gray - Squamellaria
imberbis (A.Gray) Becc. (Jebb 1991).
Anthorrhiza Huxley & Jebb, Bull. Jard. Bot. Nat. Belg. 60: 420 (1990). Type: Anthorrhiza echinel-
la Huxley & Jebb.
Lecariocalyx Bremek., Kew Bull. 1940: 192 (1940). Type: Lecariocalyx borneensis Bremek.
This may be a synonym of Straussia, but in the absence of sequence data and detailed information
on pyrene morphology its position remains uncertain.
Peripeplus Pierre, Bull. Mens. Soc. Linn. Paris, s6r. 2 1: 66 (1898). Type: Peripeplus kleineanus
Pierre.
In the absence of sequence data and detailed information on pollen and pyrene morphology the
position of this genus remains uncertain.
Psathura Comm. ex Juss., Gen. 206 (1789). Type: Psathura borbonica J.F.Gmel.
The position of Psathura remains uncertain in the absence of sequence data and detailed morpho-
logical information. It may belong in the Palicourea complex rather than in the Psychotria com-
plex.
198

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Psychotria L., Syst. ed. 10 929 (1759), nom. cons. Type: Psychotria asiatica L., lectotype (Petit
1964: 11-12).
Myrstiphyllum P.Browne, Nat. Hist. Jamaica 152 (1756), nom. rej. Type: Psychotria myrstiphyllum
Sw.
Psychotrophum P. Browne, Nat. Hist. Jamaica 160 (1756), nom. rej. Type: Psychotria brownei
Spreng., lectotype (Petit 1964: 11-12).
Mapouria Aubl., Hist. Pl. Guiane 1: 175 (1775). Type: Mapouria guianensis Aubl. = Psychotria
mapourioides DC.
Camptopus Hook.f., Bot. Mag. tab. 5755 (1869). Type: Camptopus mannii Hook.f. (non Psychotria
mannii Hiern).
Cremocarpon Baill., Bull. Mens. Soc. Linn. Paris 1: 192 (1879). Type: Cremocarpon boivinianum
Baill.
Megalopus K.Schum., Engl. Bot. Jahrb. 28: 490 (1900). Type: Megalopus goetzei K.Schum.
[= Psychotria goetzei (K.Schum.) E.M.A.Petit].
Pyragra Bremek., Candollea 16: 174 (1958). Type: Pyragra obtusifolia Bremek.
Apomuria Bremek., Verh. Konink. Nederl. Akad. Wet., Natuurk., Tweede Reeks 54(5): 88 (1963).
Type: Apomuria mollis Bremek.
Stipules mostly caducous, rarely persistent. Pyrenes planoconvex, adaxial surface plane or with two
longitudinal depressions, abaxial side usually with multiple longitudinal ridges extending through-
out the length of pyrene, rarely ? smooth, PGSs usually absent, marginal ones rarely present. Testa
with reddish, ethanol-soluble pigment. Endosperm usually ruminate.
Straussia (DC.) A.Gray, Proc. Amer. Acad. Arts Sci. 4: 42 (1858). Coffea sect. Straussia DC.,
Prodr. 4: 502 (1830). Uragoga sect. Straussia (A.Gray) Baill., Adansonia 12: 327 (1879).
Psychotria sect. Straussia (DC.) Fosb., Brittonia 16: 264 (1964). Type: Coffea kaduana Cham.
& Schltdl. = Straussia kaduana (Cham. & Schltdl.) A.Gray = Psychotria kaduana (Cham. &
Schltdl.) Fosb. (lectotype designated by Schumann (1891: 112).
Eumorphanthus A.C.Sm., Bernice P. Bishop Mus. Bull. 141: 157 (1936). Type: Eumorphanthus
fragrans A.C. Sm. (= Psychotria eumorphanthus Fosb.).
Stipules caducous. Pyrenes planoconvex (to triangular in cross section), adaxial surface smooth,
often with a short median crest at base, sometimes with shallow depressions on either side of crest
and/or a marginal grove, abaxial surface smooth to rugulose, or with 1-3 ? distinct longitudinal
ridges, which are usually confined to distal two-thirds, marginal PGSs distinct and usually long.
Testa usually with ethanol-soluble pigment. Endosperm with an adaxial, median invagination, or
with several to many invaginations that are independent of ridges.
Straussia is widely distributed from Sri Lanka to Fiji and Hawaii having, possibly, its centre of
diversity in Hawaii.
Streblosa Korth., Ned. Kruidk. Arch. 2(2): 245 (1851). Type: Streblosa tortilis (Bl.) Korth. (lecto-
type designated by Bremekamp 1947: 151).
Stipule persistence unknown (probably non-caducous). Pyrenes concave-convex, adaxial side with
or without median crest, abaxial side with 3-4 distinct crests extending throughout length of
pyrene, PGSs absent. Testa with ethanol-soluble pigment. Endosperm entire.
199

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
The major diagnostic features of Streblosa are the monochasial inflorescences, the small fruits,
and ventrally excavated pyrenes. Bremekamp (1947) recognised 25 species, 23 from the Malay
Peninsula and Indonesia and two from the Philippines.
Acknowledgements. - This study was supported by grants from the Swedish Natural Science
Research Council. In addition to the herbaria listed in tab. 1, material for sequencing was put at my
disposal by the the Lyon Arboretum, the Jodrell Laboratory of the Royal Botanic Gardens, Kew, the
Royal Botanic Gardens, Edinburgh, the University of Oxford Botanic Garden, the University of
Vienna Botanic Garden, the National Tropical Pacific Garden, Kauai, Hawaii, the National Botanic
Garden of Belgium, Meise, Dr. Asl6g Dahl, Mr. Claes Gustafsson, Mr. Ivan Jaimes, Dr. Claes
Persson, Dr. Johan Rova, and Dr. Bertil Stdihl. Ms. Vivian Alddn gave invaluable technical assis-
tance. Valuable comments on the manuscript were received from Dr. Victor Albert, Dr. Charlotte
Taylor, and an anonymous reviewer. To all these persons and institutions I am most grateful.
References
Andersson L. (2001) Margaritopsis (Rubiaceae, Psychotrieae) is a pantropical genus. Pl. Syst. Geogr 71: 73-85.
Andersson L. (2002) Re-establishment of Carapichea (Rubiaceae). Kew Bull. 57: 363-374.
Andersson L. & Andersson S. (2000) A molecular phylogeny of Tropaeolaceae and its systematic implications. Taxon 49:
721-736.
Andersson L. & Chase M. (2001) Phylogeny and classification of Marantaceae. Bot. J. Linn. Soc. 135: 275-287.
Andersson L. & Rova J.H.E. (1999) The rpsl6 intron and the phylogeny of the Rubioideae (Rubiaceae). PIl. Syst. Evol.
214: 161-186.
Backer C.A & Bakhuizen van den Brink R.C. Jr. (1965) Rubiaceae. In: Flora of Java (Spermatophytes only) 2: 111-160.
Groningen, N.V.P. Noordhoff.
Baillon H. (1879) M6moire sur les Uragoga. Adansonia 12: 323-335.
Bremekamp C.E.B. (1934) Notes on the Rubiaceae of Surinam. Rec. Tray. Bot. Njerl. 31: 248-308.
Bremekamp C.E.B. (1940) Additions to the flora of Borneo and other Malay islands: XXVII. Bull. Misc. Inform. Kew
1940: 192-193.
Bremekamp C.E.B. (1947) A monograph of the genus Streblosa Korthals (Rubiaceae). J. Arnold Arbor 28: 145-185.
Bremekamp C.E.B. (1961) On the identity of the genera Mapouria Aubl. and Grumilea Gaertn. (Rubiaceae, Psychotrieae.
Acta Bot. Neerl. 10: 307-319.
Bremekamp C.E.B. (1963) Sur quelques genres de Psychotri es (Rubiac6es) et sur leurs repr6sentants malgaches et
comoriens. Verh. Kon. Ned. Akad. Wet., Afd. Nat., ser 2 54(5): 1-181.
Bremekamp C.E.B. (1966) Remarks on the position, the delimitation and the subdivision of the Rubiaceae. Acta Bot. Neerl.
15: 1-33.
Bremer B. & Manen J.-F. (2000) Phylogeny and classification of of the subfamily Rubioideae (Rubiaceae). PIl. Syst. Evol.
225: 43-72.
Browne P. (1756) The civil and natural history of Jamaica. London.

Candolle A.P. de (1830) Prodromus systematis naturalis regni vegetabilis 4. Paris, Truettel & Wuirtz.
Darwin S.P. (1979) A synopsis of the indigenous genera of Pacific Rubiaceae. Allertonia 2: 1-44.
Davis A.P. & Bridson D.M. (2001) A taxonomic revision of the genus Dolianthus (Rubiaceae). Blumea 46: 421-446.
Davis A., Bridson D., Jarvis C. & Govaerts R. (2001) The typification and characterization of the genus Psychotria L.
(Rubiaceae). Bot. J. Linn. Soc. 135: 35-42.
De Wildeman E. (1936) Remarques a propos de formes du genre Uragoga L. (Rubiac6es). MWm. Inst. Roy. Colon. Belge,
Sect. Sci. Nat. Mid., coll. in-8", 4: 187 pp.
Farr E.R., Leussink J.A & Stafleu F.A. (1979) Index nominum genericorum (Plantarum) III. Regnum Veg. 102.
Farris J.S., Albert, V.A., Kiillersji, M., Lipscomb, D. & Kluge, A.G. (1996) Parsimony jackknifing outperforms
neighbor-joining. Cladistics 12: 99-124.
200

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Fosberg F.R. (1942) Rubiaceae. In: A.C. Smith et al., Fijian plant studies 2. Sargentia 1: 118-140.
Fosberg F.R. (1964) Studies in Pacific Rubiaceae: V. Brittonia 16: 255-271.
Gaertner J. (1788) De fructibus et seminibus plantarum, vol. 1. Stuttgart.
Gillespie J.W. (1930) New plants from Fiji - I. Bernice R Bishop Mus. Bull. 74: 1-99.
Gray A. (1858) Notes upon some Rubiaceae, collected in the United States South Sea exploring expedition under Captain
Wilkes, with characters of new species, &c. Proc. Amer Acad. Arts Sci. 4: 33-50.
Gray A. (1860). Notes upon some Rubiaceae, collected in the South Sea exploring expedition under captain Wilkes. Proc.
Amer Acad. Art Sci. 4: 306-318.
Hamilton C.W. (1989) A revision of Mesoamerican Psychotria subgen. Psychotria (Rubiaceae). Part III: species 48-61
Ann. Missouri Bot. Gard. 76: 886-916.
Hooker J.D. (1873) Rubiaceae. In: Bentham G. & Hooker J.D. (eds.) Genera Plantarum 2 (pp. 7-151). London, Reeve &
Co.
Huxley C.R. & Jebb M.H.P. (1991) The tuberous epiphytes of the Rubiaceae 1: a new subtribe - the Hydnophytineae.
Blumea 36: 1-20.
Jebb M.H.P. 1991. The tuberous epiphytes of the Rubiaceae 4: a revision of Squamellaria. Blumea 36: 53-61.
Johansson J.T. (1992). Pollen morphology in Psychotria (Rubiaceae, Rubioideae, Psychotrieae) and its taxonomic signifi-
cance. A preliminary survey. Opera Bot. 115: 1-71.
Kuntze 0. (1891) Revisio generum plantarum 2. Leipzig.
Linnaeus C. (1737) Hortus Cliffortianus. Amsterdam.
Linnaeus C. (1759) Systema naturae ed. 10. Stockholm.
Melchior H. (1964) A. Engler's Syllabus der Pflanzenfamilien 2, ed. 12. Berlin, GebrUder Borntraeger.
Miiller J. 1881. Rubiaceae. In: Martius, C. von (ed), Flora Brasiliensis 6(5): 1-485.
Nepokroeff M., Bremer B. & Sytsma K.J. 1999. Reorganization of the genus Psychotria and tribe Psychotrieae
(Rubiaceae) inferred from ITS and rbcL sequence data. Syst. Bot. 24: 5-27.
Oxelman B., Liden M. & Berglund D. (1997) Chloroplast rpsl6 intron phylogeny of the tribe Sileneae (Caryophyllaceae).
Pl. Syst. Evol. 206: 393-410.
Persson C. (2000) Phylogeny of Gardenieae (Rubiaceae) based on chloroplast DNA sequences from the rpsl6 intron and
trnL(UAA)-F(GAA) interegenic spacer. Nord. J. Bot. 20: 257-269.
Petit E. (1964) Les especes africaines du genre Psychotria L. (Rubiaceae). I. Bull. Jard. Bot. Etat, Bruxelles 34: 1-229.
Petit E. (1966) Les especes africaines du genre Psychotria L. (Rubiaceae). II. Bull. Jard. Bot. Etat, Bruxelles 36: 65-190.
Piesschaert E 2001. Carpology and pollen morphology of the Psychotrieae (Rubiaceae-Rubioideae). Towards a new tribal
and generic delimitation. Dissertation, Katholieke Universiteit Leuven.
Piesschaert E, Andersson L., Jansen S., Dessein S., Robbrecht E. & Smets E. (2000a) Searching for the taxonomic posi-
tion of the African genus Colletoecema (Rubiaceae): morphology and anatomy compared with an rpsl6-intron analysis of
the Rubioideae. Can. J. Bot. 78: 288-304.
Piesschaert E, Huysmans S., Jaimes I., Robbrecht E. & Smets E. (2000b) Morphological evidence for an extended tribe
Coccocypseleae (Rubiaceae-Rubioideae). Pl. Biol. 2: 536-546.
Piesschaert E, Robbrecht E. & Smets E. (2001) Patterns in pyrenes: the systematic significance of pyrene morphology in
Chassalia (Rubiaceae-Psychotrieae) and related genera. Flora 196: 121-131.
Plunkett G.M. & Downie S.R. (1999) Major lineages within Apiaceae subfamily Apioideae: a comparison of chloroplast
restriction site and DNA sequence data. Amer J. Bot. 86: 1014-1026.
Richard A. (1834) M6moire sur la famille des Rubiac6es, contenant la description g6n6rale de cette famille et les caracteres
des genres qui la composent. Mim. Soc. Hist. Nat. Paris 5: 81-304.
Ridsdale C.E. (1975) Notes on New Guinea Rubiaceae. Chaetostachydium. Blumea 22: 267-269.
Robbrecht E. (1988) Tropical Woody Rubiaceae. Opera Bot. Belg. 1: 1-271.
Robbrecht E. (1994) Supplement to the 1988 outline of the classification of the Rubiaceae. Index to genera. Opera Bot.
Belg. 6: 173-196.
Royen P. van (1973) Sertulum Papuanum 18. Two superfluous genera in the New Guinean flora. Bot. Notiser 126: 417-425.
Schnell R. (1957) Notes sur les Psychotriees (Rubiac6es) B l'ouest africain. Mim. EI.FA.N. 50: 50-93.
Schnell R. (1960) Note sur le genre Cephaclis, et le probleme de l'evolution parallele chez les Rubiac6es. Bull. Jard. Bot.
ttat Brux. 30: 357-373.
Schumann K. (1891) Rubiaceae. In: Engler A. & Prantl K. (eds.) Die natiirlichen Pflanzenfamilien IV.4. (pp. 1-156).
Leipzig, Wilhelm Engelmann.
Smith A.C. (1936) Fijian plant studies. Bernice P. Bishop Mus. Bull. 141: 1-166.
201

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC
Smith A.C. & Darwin S.P. (1988) Rubiaceae. In: Smith A.C. (ed.) Flora Vitiensis Nova. A New Flora of Fiji
(Spermatophytes Only) 4 (pp. 143-377). Lawai, Pacific Tropical Botanical Garden.
Sohmer S.H. (1977) Psychotria L. in the Hawaiian Islands. Lyonia 1: 103-186.
Sohmer S.H. (1978) Morphological variation and its taxonomic and evolutionary significance in the Hawaiian Psychotria
(Rubiaceae). Brittonia 30: 256-264.
Steyermark J.A. (1972) Rubiaceae. In: B. Maguire et al., The botany of the Guayana highland - part IX. Mem. New York
Bot. Gard. 23: 277-832.
Swofford D.L. (1999) PAUP*. Phylogenetic Analysis Using Parsimony (*and other methods). Sunderland, Massachusetts,
Sinauer Associates.
Taylor C.M. (1996) Overview of the Psychotrieae (Rubiaceae) in the Neotropics. Opera Bot. Belg. 7: 261-270.
Verdcourt B. (1975) Studies in the Rubiaceae-Rubioideae for the 'Flora of Tropical East Africa': I. Kew Bull. 30: 247-3
White T.J., Burns T.S., Lee S. & Taylor J. (1990) Amplification and direct sequencing of fungal ribosomal RNA genes for
phylogenetics. In: PCR protocols: a guide to methods and applications. San Diego, Academic Press.
Zappi D. & Nunes T.S. (2000) Notes on the Rubiaceae of northeastern Brazil. I. Erithalis, Psychotria and Rudgea. Ke
Bull. 55: 655-668.
Manuscript received September 2001; accepted in revised version April 2002.
202

84.115.220.56 on Sun, 07 Nov 2021 10:41:46 UTC

2002 - Relationships and Generic Circumscriptions in The Psychotria Complex (Rubiaceae

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

2002 - Relationships and Generic Circumscriptions in The Psychotria Complex (Rubiaceae

Uploaded by

Copyright:

Available Formats

Relationships and Generic Circumscriptions in the Psychotria Complex (Rubiaceae,

Stable URL: https://www.jstor.org/stable/3668767

This content downloaded from

Botanical Institute, Goteborg University,

Key words: Amaracarpus, Calycosia, cpDNA, Cremocarpon, Gentianales, Grumilea, Hydnophy-

This content downloaded from

This content downloaded from

This content downloaded from

2 Material and methods

2.1 Taxon sample

This content downloaded from

2.2 Laboratory procedures

Table 1. List of taxa used in cladistic analyses.

Taxon Origin Voucher NCBI acc no.

Amaracarpus grandifolius Valeton New Guinea Takeuchi 8926 (L) AF410678* -

C. macrocyatha Fosberg Fiji Rova & Gustavsson 2486 (GB) AF410671* -

C. subochreata (De Wild) Tanzania Frimodt-Moller 2 (C) AF369841

"M." macrochlamys Bremek. Madagascar Davis 1005 (K) AF410689* -

This content downloaded from

Taxon Origin Voucher NCBI acc no.

Myrmephytum selebicum (Becc.) Becc. unknown3 without voucher AF410691* AF034916

This content downloaded from

Taxon Origin Voucher NCBI acc no.

PF leucopoda E.M.A.Petit Kenya Polhill & Robertson 4831 (BR) AF410730*

P? aff. pichisensis Standl. Bolivia Persson &

FP rubra (Lour.) Poir. Japan Aiba 4 (GB) AF410749* -

PF verschuerenii De Wild."1 unknown'" without voucher AF410762" -

This content downloaded from

2.3 Alignment and indel coding

2.4 Morphological data

Table 2. Morphological characters used for optimisation (fig. 5-9).

Taxon Source Characters

This content downloaded from

Taxon Source Characters

2.5 Cladistic procedures

This content downloaded from

This content downloaded from

4.1 Major features

This content downloaded from

99.C Hydnophytum grandifiorum

Figure 2. Results of jackknife analysis of rpsl6 intron data (part of jackknife

This content downloaded from

Psychotria auxopoda Q Psychotria coptosperma o

Figure 3. Results of jackknife analysis of rpsl6 intron data (part o

This content downloaded from

s99/501s 9Myrmecodia tuberosa

100/100100 Myrmephytum selebicum

63/<50ro4 Psychotria insularum

100/(r Psychotria kaduana D

97,s3i<o ~Psychotria ernesti

65/-/91 Psychotria loniceroides

71/-/88 - Notopleura guadalupensis

This content downloaded from

4.2 Character distribution

This content downloaded from

stipules caducous Hydnophytum formicarium

This content downloaded from

marginal PGSs Hydnophytum formicarium

This content downloaded from

Optimisation (fig. 8) fails to decide whether presence of an SCP is a plesiomorphic or apomorphic

This content downloaded from

ridges on abaxial pyrene surface Hydnophytum formicarium

This content downloaded from