Botanical Journal of !he Linnean Sock@ ( 1 982) 84: 355-377. With 15 figures
The delimitation of the tribe Anthospermeae
and its affinities to the Paederieae (Rubiaceae)
CHRISTIAN PUFF
Institute of Botany, University of Vienna,
Rennweg 14, A-1030 Vienna, Austria
Received September 1981, accepted for publication February 1982
A critical comparison of many characters suggests that the Rubiaceae tribe Anthospermeae is closely
allied to the tribe Paederieae. The delimitation of the tribe Anthospermeae from other tribes is
redefined as to include only wind-pollinated genera, and characters of fruit structure, pollination
biology and distribution patterns support the subdivision of the tribe into the three subtribes:
Anthosperminne, Operculariinae and Coprosminae.
All insect-pollinated genera previously placed in the Anthospermeae are transferred to the
Paederieae. It is shown that the genus Neogaillonia Linchevskii (=Gaillonia A. Rich. ex. DC.),
previously included in the Spermacoceae, also belongs to this tribe; the genera Pteragaillonia
Linchevskii, Pseudogaillonia Linchevskii, Jaubertia Guill. and Choulettia Pomel are considered synonyms
of .Neogaillonia.
The Anthospermeae are believed to be closely, and the Paederieae more remotely, allied to the tribe
Theligoneae.
KEY WORDS :-anemophily~Anthospermeaeclassificationentomophily ~Paederie;ie~Theligoneae
CONTENTS
Introduction . . . . . . . . . . . . . . .
Historical survey . . . . . . . . . . . . . .
Circumscription of the Anthospermeae . . . . . . . . .
Wind-pollinated flowers of the Anthospermeae . . . . . . .
Insect-pollinated flowers of the Anthospermeae, Paederieae and .Neoguillonia
Further characters of the Anthospermeae, Paederieae and .Neogaillonia. .
Conclusion
. . . . . . . . . . . . . . .
Taxonomic conspectus . . . . . . . . . . . . .
Relationships to other tribes
. . . . . . . . . . .
Acknowledgements. . . . . . . . . . . . . .
Appendix. Synonyms of.Neogaillonia; new combinations . . . . .
References. . . . . . . . . . . . . . . .
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355
356
357
357
359
364
369
370
373
373
374
375
INTRODUCTION
During several years of intensive investigation on African and Madagascan
Rubiaceae Anthospermeae, evidence has accumulated to suggest that the tribe
Anthospermeae as circumscribed in the widely accepted classifications of the
+
0024-4074/82/040.355 23 S03.00/0
355
0 1982 The Linnean
Society of London
356
C. PUFF
Rubiaceae (Hooker, 1873; Schumann, 1891) is a highly unnatural, inflated group
with only vague relationship to other tribes. An attempt is made to redefine the
tribe on the basis of all available data.
HISTORICAL SURVEY
Shortly after Chamisso & von Schlechtendal (1828) described the tribe
Anthospermeae, Richard (1830) also studied the group and included the genus
Coprosma. His remarks concerning the Anthospermeae were noteworthy but
unfortunately their significance was either not appreciated at the time or
overlooked by later authors. While Chamisso & von Schlechtendal distinguished
the members of the tribe by their two-seeded fruits usually dehiscing into two oneseeded mericarps, Richard (1830: 56) pointed out that “ . . . le vrai caractere qui
la distingue . . . consiste uniquement dans le forme et la structure des
stigmates. . . . on trouve deux trks-longs stigmates filiformes, subults, poilus”, the
first time that an indirect reference was made to the anemophilous flowers of this
group (see p. 357).
Richard ( 1830) also described the tribe Opercularieae (Opercularia, Pornax) and
stated that the genus Opercularia, upon which the tribe is based, should not be
excluded from the Rubiaceae and placed in a family of its own (cf. de Jussieu,
1804).He also noted its affinity to the Anthospermeae “par ses deux longs stigmates
subults et poilus . . . ”. De Candolle (1830) accepted Richard’s classification and
also listed (de Candolle, 1830: 343) the elongated hairy stigmas as being a
diagnostic character of the Anthospermeae.
During the 43 years which elapsed between the publication of the accounts by
Richard (1830) and de Candolle (1830) and Hooker’s (1873) system, many new
genera were described. Hooker added seven new genera to the Anthospermeae semu
Richard and de Candolle and also included the Opercularieae and another five
genera previously placed in two different tribes by de Candolle (cf. Table 1) .
The subfamilial classificationof the Rubiaceae by Baillon ( 1880),although never
widely accepted, deserves special consideration. He drastically reduced Hooker’s
25 tribes of Rubiaceae to 15 and included the tribe Paederieae in the
Anthospermeae. In Die natiirlichen Pjanzenfamilien, Schumann ( 1891) accepted
Hooker’s classification and added only the genus Cremocarpon. But the latter as well
as other genera described after Schumann’s publication were sooner or later
excluded from the Anthospermeae (Table 1). It is noteworthy that, in his remarks
concerning the Paederieae, Schumann pointed out the very close affinity to the
Anthospermeae and stated that “the berry-like or indehiscent fruits, in at least
most cases, allow a distinction” between the two tribes. Thus both Hooker and
Schumann placed the Paederieae next to the Anthospermeae. For reasons
unexplained, both Verdcourt (1958) and Bremekamp (1966) did not follow suit
and placed the two tribes far apart from each other without mentioning or
discussing possible affinities between them. Bremekamp merely stated that “in its
present delimitation this tribe (Paederieae) makes a rather unnatural impression”,
For a satisfactory discussion of tribal delimitation, yet another genus,
.Neogaillonia Linchevskii, has to be considered. Linchevskii ( 1973), correctly
substituting the name Reogaillonia for the illegitimate name Gaillonia A. Rich ex
DC., also created the segregate genera Pterogaillonia and Pseudogaillonia and upheld
the genera Jaubertia and Choulettia which were, in the past, considered synonyms of
RUBIACEAE
35 7
Gaillonia. In my opinion, these four genera are not sufficiently different from
.Neogaillonia to warrant their separation ; they should be considered synonyms (see
Appendix for the necessary new combinations). The genus had been placed in the
tribe Spermacoceae by all of the authors mentioned above. Both Hooker (1873)
and Schumann (1891) categorically stated that Neogaillonia has an “ovule attached
to the middle of the septum”, although Richard (1830: plate 5, Fig. 3F) showed
clearly, and correctly, that the solitary, anatropous ovules are basally attached as
in the Anthospermeae. The basal attachment of the ovules can also be seen in a
figure of .Neogaillonia putorioides (A. R. Smith) Linchevskii (as Jaubertia putorioides in
Smith, 1971). Qarar. (1973) also suggested an affinity of Neogaillonia to the
Anthospermeae but stated that its position within the Rubiaceae is uncertain.
Hemsley (in Aitchison, 1882: 162) pointed out affinities between Aitchisonia
(Paederieae) and Neogaillonia.
A re-investigation of the genera traditionally placed in the Paederieae and a
carefiil study of the available literature indicates a clear affinity with the
Anthospermeae. Genera placed in both the Anthospermeae and Paederieae sensu
Hooker (1873) and Schumann (1891) are discussed below.
CIRCUMSCRIPTION OF T H E ANTHOSPERMEAE
From the tribes with one, basally-attached, anatropous ovule per locule, Hooker
( 1873) distinguished the Anthospermeae by the following characters: “flowers
more often unisexual, stamens inserted at the base of the corolla, rarely at the
throat, ovary with one to four locules, style fused or branches more often filiform,
fruits baccate or indehiscent”. Schumann (1891) used “ovary two- or seldom
more-locular, stamens usually attached to the base of the corolla tube, mostly
dioecious plants”. These circumscriptions are, in part, incorrect and unsatisfactory.
The description of filament attachment is particularly misleading. A reinvestigation has shown that only certain genera (genera 1-12 in Table 1) have the
filaments attached to the base or seldom just below the middle of the corolla tube
(Carpacoce species with unusually long corolla tubes). All the others have the
filaments attached to the throat or always above the middle of the corolla tube.
Both Hooker (1873) and Schumann (1891) used these characters in their keys to
divide the tribe into two groups and as a result contradicted themselves as regards
to filament attachment characters in the tribal key of the Rubiaceae. They
included the genus Serissa in the 12 genera (Table 1) with filaments attached at or
near the base of the tube, although its filaments are joined to the tube near the
throat. They were misled by the fact that the stamen vascular system remains more
or less distinct and prominent below the actual point of insertion of the filaments,
and can therefore be followed to the base of the corolla tube.
WIND-POLLINATED FLOWERS OF THE ANTHOSPERMEAE
The same group of 12 genera (Table l ) , discussed in the previous section, is
characterized by having anemophilous flowers. The stigmas are long, filiform and
the receptive surface is increased by unicellular hairs of varying length (Figs 2 4 ,
below) ; hairs are distributed over the whole surface of the stigmas and not confined
to their inner portion. The style is united, either (sub-)obsolete or, if welldeveloped, never longer and commonly much shorter than the stigmatic areas.
Richard
-
Guettardeae: “gen. obsc.”
CofFeeae
Baillon
I
I
“Operculariaceae”, cf. de Jussieu (1804)
Operculariaceae, to Dipsacales? cE Bremekamp (1966)
Anthospermeae of Chamisso &
von Schlechtendal (1828)
Comments
Anthospermeae
-
-
Guettardeae subtribe Cuettardinae Anthospermeae
Ant hospermeae
Guettardeae subtribe Cuettardinae Anthospermeae
Paederieae
to Dipsacales?, cf. Bremekamp (1966)
The correct name of the genus is
Spermadictyon (cf. Backer & Backhuizen van den
Brink, 1965: 347)
now generally considered synonyms
of Paederia (cf. Verdcourt, 1976:175)
Spermacoceae subtribe Putoriinae Anthospermeae
Spermacoceae subtribe Putoriinae Anthospermeae
Anthospermeae
Guettardeae subtribe Guettardinae Chiococceae
Ant hospermeae
Spermacoceae subtribe Putoriinae Anthospermeae
Anthospermeae cf. Robbrecht & Puff (1981); to Hedyotideae (Pu&
1982)
Anthospermeae to Psathureae (= Psychotrieae), Bremekamp (1958)
to Knoxieae, Verdcoun (1958)
Anthospermeae
Anthospermeae
Anthospermeae
Anthospermeae
Anthospermeae
Anthospermeae
Anthospermeae
Guettardeae subtribe Guettardinae Anthospermeae
Anthospermeae
Anthospermeae
Opercularieae
Anthospermeae
Opercularieae
Anthospermeae
Anthospermeae
An thospermeae
An thospermeae
Anthospermeae
Anthospermeae
de Candolle
* = I n Schumannonly; ** =generadescribedata laterdate butassociatedwith the Anthospermeae. Authorsofgeneraaccording to Index NorninurnGenericorum (Plantarum) (Farretal.. 1979).
24. I.cptodermis Wall.
25. Pseudopyxis Miq.
26. Aifchisonia Hemsl. ex Aitch.*
Paederieae
22. Paed& L.
(syn. Lygodisodea Ruiz & Pav.
honfea A. Rich. ex DC.)
23. Spermadupon Roxb.
(syn. Hamiltonia Roxb.)
Anthospermeae
2. Ncnax Gaenn. (syn. Ambraria Cruse)Anthospermeae
Anthospermeae
3. Galopina Thunb.
4. Plyllis .I
An thospermeae
5. Carpacme Sond.
6 . CoprosmaJ. R. & J. G. A. Font.
Anthospermeae
7. Nertera Banks & Soland. ex Gaertn.“Guettardaceae dubiae”
8. Coynula Hook. fil.
9. Nomandia Hook. fil.
10. Opprcularia Gaenn.
Opercularieae
Opercularieae
1 1. Pmax Soland.
12. Ehfheranthus F. Muell.
(“Eleuthranthes”)
13. Plocmno Ait.
Spermacoceae subtribe Putoriiae
14. Putoria Pea.
Spermacoceae subtribe Putoriiae
15. Crqllis E. Mey. ex Hook. fil.
16. Mikhella L.
“Guettardaceae dubiae”
17. Wlogia Torr. ex Bentham
Spermacoceae subtribe Putoriinae
18. SnisSo Comm. ex Juss.
19. Otwphora Zucc.
(syn. Mericocabx Bamps**)
20. Crnnocarpon Baill.*
21. Calanda K. %hum.**
(syn. Ofocephalur Chiov.**)
1. Anfhospmum L.
Anthospermeae
Hooker, Schumann
Table 1. Genera belonging to tribe Anthospermeae and tribe Paederieaeaccording to Hooker ( 1873)and Schumann ( 1891), and the positionof
these genera in the classification systems of Richard ( 1830), de Candolle (1830) and Baillon (1880)
RUBIACEAE
359
There are usually two stigmas, even in species in which one of the two carpels or
ovules is reduced (except in Carpacoce). In a few species of Coprosma, the number of
stigmas and carpels is increased from typically two; in C. lucida, for example, there
are usually two stigmas, but flowers with up to five stigmas also occur (Wild &
Zotov, 1906). The long slender filaments are basally attached and bear relatively
large, dangling, dorsifixed anthers (Fig. 1). The flowers are odourless and have no
trace of a nectar-producing disk ; the corollas are inconspicuously coloured,
commonly greenish to greenish yellow, and, in African genera in particular, the
corolla lobes of hermaphrodite and male flowers are frequently conspicuously
rolled back (Fig. 1). In most species there is a marked dimorphism in corolla size
and, to a certain extent, in corolla shape between the hermaphrodite and male
flowers and purely female flowers. The corollas of the female flowers are much
smaller and, sometimes, even absent as compared with the corollas of the
distinctive hermaphrodite and male flowers (Figs 1, 2). Hermaphrodite flowers are
either protandrous (Anthospermurn, Carpacoce, Galopina, Phyllis) or protogynous
(Coprosma, .Nertera, .Nomzandia, Corynula, Pomax). Nenax is the only genus in which all
species are strictly dioecious. Often, other genera also have some dioecious species,
but include hermaphrodite species, some gynodioecious species and some species
with much more complicated sex distributions. Corynula and Eleutheranthus seem to
have hermaphrodite flowers only. It must be stressed that, in contrast to
Verdcourt’s (1958) and Bir Bahadur’s (1968) views, heterostyly is absent in these
genera. Dioecism in the species of the wind-pollinated group is, furthermore, not to
be considered “the ultimate consequence” of heterostyly.
Wind-pollination in this tribe was first pointed out by Thomson (1881) for
Coprosma and Nertera; Cheeseman (1887) cofirmed this for Coprosma. Delpino
( 1889) independently mentioned these two genera as being anemophilous, in
addition to Phyllis, Opercularia, Pomax and Anthospermurn. Schumann (1891) must
have overlooked these publications as he did not refer to wind pollination in his
discussion of general characters of the Rubiaceae or in connection with the
Anthospermeae. Although Loew (1905), and later Werth (1923) and Wettstein
(1935), did refer to the anemophilous flowers of at least some of these genera, this
was not mentioned at all by Verdcourt (1958) or Bremekamp (1966).
INSECT-POLLINATED FLOWERS OF T H E ANTHOSPERMEAE, PAEDERIEAE AND
.NEOGAILLO.NIA
In the majority of taxa (Table 1, genera 13-18 and 22-26), the flowers are
hermaphrodite and frequently heterostylous (consult Table 2). Crocyllis is an
exception in that it is gynodioecious, but there is no corolla size or shape difference
between hermaphrodite and female flowers (Puff & Mantell, 1982). Paederia,
according to Schumann ( 1891) , has hermaphrodite and polygamous-dioecious
flowers. All species of Neogaillonia are, according to Qarar ( 1973), hermaphrodite
and homostylous.
Flowers have corollas with funnel-shaped to more or less cylindrical, often very
long corolla tubes (Figs 5, 6). The tubes are frequently hairy inside, ranging from
sparsely hairy to densely pilose. Hairs may be restricted in distribution, occurring
only at or below the throat, or the tube may be hairy to the base. The stamens are
inserted at the throat or above the middle of the tube and have relatively short
filaments so that the anthers are only slightly exserted or are enclosed in the
Anthospmnum
Anthosperminae
Anthospermeae
PAQ/J/O
4(5)
4
5
4-7
-
F
+M
(creeping) perennial herbs
5
-
-
small trees, shrubs
(creeping) perennial herbs
+M+
F
FF
5
4-5
shrubs
dwarf shrubs (perennial herbs)
-
+v
+M
+M
dwarf shrubs
perennial herbs
small trees, large shrubs+
perennial herbs
small trees-rdwarfshrubs
D
D
D
D +M;-
+M
FF
2
2
2
1(2)
D
- 4-5(10)
-
-
4-5
-
+
+
-
-
+
-
+
+ -
+
+
+
South Africa, Namibia
southeastern Afiica
Canary Islands and Madeira
southAfiica:swtmvestancapeplwina
66
22
22
22
22
44
40
220
c. 154
New Caledonia
as Coprosma, but further north and south
and occurring in South America
western South America
22
(south temperate) amphi-transpacific,
?32 but not reaching South America
44
-68)
Africa, Madagascar; Yemen Arab
Republic
44
22
Table 2. Summary of the most important characters of the genera of the tribe Anthospermeae Cham. & Schlecht. ex DC., subtribe
Anthospermineae, subtribe Copmsminae Puff, and subtribe Operculariinae Bentham, and the tribe Paederieae DC
+
+
+
+
+
+
-?
Aitchisonia
Serissa
Paederia
+
+
4
4-5
‘
H
I
5
4-5
5
4-5
4-5
3-6
5
5-7
+,-
+
+
+
+
+
+
+
+
+
+
2-3
2-3
2
5
FF
D
D
D
D
D
4-5
5
4
F
FF
D
D
FF
D
2
2-3
2
2
2
D
D
D
+V
+M+
+V
shrubs
climbing shrubs (or perennial
herbs)
dwarf shrubs
(dwaqshrubs
perennial herbs
shrubs
+V
+V
-
+M
-
*M
dwarf shrubs
subsrubs, perennial herbs
(sub)shrubs, short-lived perennial
herbs
annual(?)
dwarf shrubs
shrubs
shrubs
perennial herbs
(creeping) perennial herbs
+M
+M;-
+V(?)
+O
+O
+
+
+
?-
?-
+
+
+
+
+
+
?-
+
+ +
+
+
+
+
+
+
+
USSR, Afghanistan,
Pakistan-Arabia-northwestern Africa ;
Socotra
Mediterranean; E. to Iraq
Canary Islands
southern Namib desert
southwestern China; western U.S.A.
Korea and adjacent parts of USSR,
Japan; eastern North America to Mexico
Japan
eastern India, Malaysian Peninsula to
China
southwestern Australia
(south)eastern Australia
Australia, Tasmania
+
22
24
26
22 Himalayas, East Bengal to northeastern
44 China, Japan
>60
Afghanistan and adjacent parts of Iran
and Pakistan
22 China?; widely cultivated in East Asia
44 southeast Asia to Japan; to Mauritius,
Madagascar. trop. and subtrop. eastern
Africa; South America
22
22
44
22
22
+ +
FLOWERS: Insertion of stamens above the middle of the corolla tube = +, below the middle (usually at base) = - ; style much longer than the stigma lobes = ,stigmatic area much
enlarged, style very small or *O= -. FRUITS: Very fleshy=FF, ffleshy=F, dry=D; dehiscence: fruits separating into indehiscent m e n c a p = + M ; Mericarps themselves dehiscing
= M ;exocarp splitting into valves = +V. Inhctescence opening by means ofan operculum = +O; fruits indehiscent = -. PHYTOCHEMICAL DATA: hperubsidic glycosides and
foetid odour: definite absence = - (blanks: no data available).
s
+
+
+
Pseudopyxis
Spermdutyon
Mitchella
-
-?
?
+
+
+
Kellogia
cr~vllis
Puloria
Plocama
+
+
+
+
+
+
.Nmgaillonia
Paederieae
(415
1
less
4-5
Eleutheranthus
-
1
Q
1
5
4-5 or
Pmax
Opercularia
Operculariinae
362
C. PUFF
Figure 1-4. SEM. Figs 1, 2. Male and female flower ofNenax divaricatn (South Africa, western Cape
Prov., Puff 790712-2/1, WU); note corolla size and shape differences and insertion of filaments and
rudimentary ovule (ro) in male flower, t. x 26. Fig. 3. Portion of inflorescence of female plant (st:
stigmas) of Anthospermurn herbaceum (South Africa, Transvaal, Puff 791201-1/2, WU), c. x 2. Fig. 4.
Stigma hairs of Anfhospemum randii (Malawi, Chongoni For. Res., Puff 780215-2/1, WU),c. x 165.
RUBIACEAE
Figures 5, 6. Fig. 5. Flower pair of Mikhellu repens (U.S.A., Mississippi, photo PuJJ 710724-2/4)
subtended by a pair of leaf-like bracts (br);short styled flower with exserted stamens (Fa: filaments);
the inside of corolla lobes and tube is densely pilose, c. x 6. Fig. 6. Part of inflorescence of Paelria sp.
(Madagascar; photo Professor S.Vogel), c. x 0.5.
363
364
C. PUFF
uppermost part of the tube. It is only in short-styled flowers that the anthers may
be more or less conspicuously exserted (Fig. 5). In Aitchisoniu the stamens are
inserted at different levels, a feature which led Bremekamp (1966) to suggest a
placement of the genus in the Dipsacales rather than the Rubiaceae. Loew (1905)
and Qarar (1973) recorded the same pattern in various species of Puederiu and
Neogailloniu respectively.
While there are usually only one or two ovules per flower in the wind-pollinated
Anthospermeae, flowers of the insect-pollinated taxa more often have five or four
ovules and carpels. There is a long, more-or-less filiform, united style which is
terminated by two to five papillose, but never distinctly hairy, short stigma lobes,
depending on the number of carpels. The stigmatic surfaces are always much
smaller than the stylar portions of the gynoecium. Certain Puederiu species are
aberrant in that the styles are sometimes split nearly all the way to the base, but
the papillose stigmatic area remains considerably shorter than the free, nonreceptive stylar segment. Disks are present in all genera and are sometimes massive
(cf. Puederiu pospischilii K. Schum., in Verdcourt, 1976: fig. 19).
In conclusion, it can be said that all the insect-pollinated genera of the
Anthospermeae and Paederieae, and Neogailloniu are so similar in floral
morphology that it becomes impossible to draw a clear demarcation line between
them.
FURTHER CHARACTERS OF THE ANTHOSPERMEAE, PAEDERIEAE AND NEOCAII.LO.NIA
From the previous two sections it is clear that, on the basis of floral biology and
morphology, genera 1-12 of the Anthospermeae and the remaining genera form
two clear-cut groups. It is now necessary to determine (1) whether there are any
other characters which could test the close affinity between the two groups and (2)
whether the insect-pollinated genera presently placed in different tribes share
features other than floral characters which would group these genera together.
Fruits: At first sight, the different genera seem to exhibit a bewildering range of
different fruit types. However, one character common to all genera is evident : the
seed itself is never released .but remains enclosed in an indehiscent endocarp
(Figs 7, 13-14), a mericarp (Fig. 15) or an entire fruit. The fruit wall is either
Figure 7. Cross section ofmencarp of Nomandia neocaledonica (New Caledonia, White 2007, K ) ; the exocarp (ex)
with five vascular bundles (vb) is about to separate from the thick, sclerenchymateous endocarp (en) which
encloses the seed (te: testa, a:endosperm, co: cotyledons); camera lucida drawing of microtome section, c. x 55.
RUBIACEAE
365
Figures 8 12. SEM. Figs 8 1 I . fomax urnbellah (SW Australia, Zerny s.n., W). Fig. 8. Side view of
infructescence with calyx-like bracts, c. x 22. Fig. 9. Removed operculum, c. x 22. Figs 10, I 1. Dorsal and
ventral view of endocarp containing the seed, c. x 22. Fig. 12. Ventral view of mericarp of Nomandia
neocakdonzca (New Caledonia, White 2007, K ) ; exocarp (ex) separated from ‘stone’ (endocarp plus
seed), c. x 22.
differentiated into two layers with an exo- and endocarp, or three layers with an
exo-, meso- and endocarp.
The fruits are distinctly fleshy, more-or-less fleshy or dry. Distinctly fleshy fruits
are indehiscent and are drupes containing two to five, depending on the number of
carpels present, ‘stones’or pyrenes (e.g. Plocama, Coproma).The more-or-less fleshy
fruits, more-or-less transitional to dry fruits, split into two indehiscent mericarps.
In Putoria, for example, the fleshy exocarp dries up and remains on the endocarp as
a skin and in Normandia the exocarp separates from the endocarp (Figs 7, 12).
366
C. PUFF
Figures13-15.SEM.Figs 13,14,Pomaxumbcllufa. (S.W.Australia,<mys.s.n.,W).Sectionofendocarp (en)
plus seed (te: testa, es: endosperm, c. x40. Fig. 14. Endocarp (en) removed to expose testa (te) ofseed,
c. x 40.Fig. 15. Cross section offruit ofAnfhospenumsp. near A. acfhwpicum (South Africa, eastern Cape
Prov., Puff 79011+2/1); the mericarps have a distinct endocarp (en) made up of sclerenchyma cells,
however, the exocarp (with a massive, tannin containing epidermis; vb: vascular bundle) does not
separate from it-the mericarps themselvesremain indehiscent; the thick arrow points to the abscission
zone between the two mericarps; te: testa; c. x 110.
Dry fruits are most common. They are indehiscent or dehiscent in various ways.
Most frequently, fruits separate into two indehiscent mericarps (e.g. Phyllis,
Anthospermurn, Neogaillonia) . The mericarps show a distinct differentiation into an
endocarp and a thin exocarp (Fig. 15). In Anthospermurn and some Nenax species,
the mericarps sit on variously-shaped carpophores. Similar structures occur in
some Neogaillonia species (see for example, Jaubert & Spach, 1843: fig. 77), but are
RUBIACEAE
367
absent in all other genera. Transitions to indehiscent fruits occur in Nenax and
Neogaillonia; also the fruits of Crocyllis do not always dehisce readily.
In some genera, such as Carpacoce and Pseudopyxis, the exocarp splits into several
valves, the number equalling the number of carpels or calyx lobes. The released
diaspores are the endocarps containing seeds. The fruits of Paederia belong to this
group, but the endocarps are often conspicuously winged. In Leptodennis, and
perhaps Spermadictyon, the hard endocarp is surrounded by a net-like mesocarp.
Among the genera with dry fruits, Pomax and Opercularia are unusual in that they
produce ‘infructescences’ which develop from fused ovaries of flowers of a partial
inflorescence. These infructescences open by means of an operculum (Figs 8, 9).
The diaspores are endocarps enclosing the seeds (Figs 10, 11, 13, 14), and not
the seeds themselves as suggested by Schumann (1891). Similarly, in Mitchella, the
four-carpellate ovaries of fused flower pairs give rise to ‘double drupes’ with two
groups of four pyrenes.
Inzorescences: The basic and most common “central type” of rubiaceous
inflorescence in the sense of Weberling (1977) is a typical thyrse or more accurately
a ‘thyrso-paniculate’ inflorescence (Troll, 1964: 95). It is found in a number of
genera across the present tribal boundaries, in Phyllis, Plocama and Spennadictyon for
example. In other genera, the inflorescences are variously modified into three main
groups: ( 1) a congested pseudoverticillate arrangement of flowers (e.g.
Anthospermurn), (2) more-or-less head-like inflorescences (e.g. Aitchisonia), and (3)
much reduced inflorescences with one-flowered partial inflorescences (e.g. Nutera,
and Nenax species). These modifications are considered to be derived from typical
thyrsus inflorescences. In the larger genera (e.g. Anthospermurn and Coprosma) there
is a wide range of inflorescence organization incorporating all the variations
described above.
The inflorescences of Pomax and Opercularia deserve a special mention. In these
genera, the ovaries of partial inflorescences are fused, thus forming single units of
three or more flowers. In Pomax, these units are arranged into a n umbel-like
inflorescence, which according to Weberling ( 1977), represents a modified thyrse.
Growth forms: There is a very wide range in both growth form and habit. For
example in the large genus Anthospennum, there are small trees, large shrubs, erect
or cushion forming dwarf shrubs and perennial herbs, some of which can be rather
short-lived. Coprosma exhibits a similar range of growth forms, although there are
no perennial herbs (Oliver, 1935).
In general, woody plants predominate, although there is a conspicuous absence
of larger trees. Amongst these, Paederia is the only genus with climbing vine-like
species. Some genera are purely herbaceous, for example Mitchella, Pseudopyxis,
.Nertera, and all of them are perennial or sometimes biennial. There may not be any
true annuals although material of the little-known genus Eleutheranthus still awaits
re-examination. It is listed as an annual by both Bentham (1866) and Schumann
(1891).
Leaves and stipules : Broad and more-or-less large leaves are characteristic for most
herbaceous genera. Woody genera tend to have smaller, often narrower, lanceolate
to linear, and tougher leaves. It is particularly noteworthy that various kinds of
xeromorphic leaves such as convolute or revolute rolled leaves and equifacial
368
C. PUFF
round leaves ( = “kon- und revolute Rollblatter, aquifaziale Rundblatter” of
Napp-Zinn, 1973) occur in all of the African genera, except Galopina, and
Neogaillonia (Qarar, 1973). In species with leaves round in cross section, stomata
are dispersed over the whole surface, and in certain species of Carpacoce, a species of
Coprosma (Foweraker, 1916) and in Neogaillonia (Qarar, 1973) the more-or-less flat
leaves may be amphistomatic. Numerous species of Coprosma are known to have
domatia (Oliver, 1935), but there are no records of their occurrence in any other
genus of this group.
Aitchisonia is unusual in having peculiar, multicellular, stalked glands, perhaps
homologues of colleters, occurring on leaves, bracts and stems. They are quite
conspicuous on new growth, but disappear on the older parts.
Stipules of herbaceous genera tend to be much-dissected and relatively large
except in Nertera with its more reduced, more-or-less triangular stipules. Woody
genera tend to have stipules with two or one more-or-less triangular lobes, or show
a reduction of stipules to stipular sheaths. In the larger genera (Anthospennum in
particular) there is much variation in stipule structure. Stipules are deciduous in
species of Paederia, but persistent in all other genera.
Phytochemical data: Available data are few. According to Kooiman (1969) and the
literature cited therein, all investigated genera of the Anthospermeae and
Paederieae sensu Hooker and Schumann possess asperulosidic glycosides, although
these glycosides do not occur in a number of Coprosma and Nertera species.
Several taxa are known to produce an unpleasant foetid odour when the leaves
are crushed. Paederoside, a compound structurally related to asperulin (Hegnauer,
1973), appears to be responsible for the odour. It was first isolated from plants of
Paederia scandens (Lour.) Merrill by Inouye et al. ( 1968).Foetid odour or paederoside
has been also recorded in species of Nertera (Allan, 1961), Spermadictyon and Serissa
(Backer & Backhuizen van den Brink, 1965), Leptodermis (Ohwi, 1965), Coprosma
and Opercularia (Hegnauer, 1973), Plocama and Putoria (Mendoza-Heuer, 1977),
Aitchisonia (Hemsley, in Aitchison, 1882), Kellogia (Dempster, personal
communication), Carpacoce and Crocyllis (Puff & Mantell, 1982). There is little
doubt that remarks as to the “rather unpleasant odour” of the plants on herbarium
sheets of Neogaillonia (for example Lamond I 3 9 in Edinburgh herbarium) are also a
clear reference to the presence of paederoside.
Outside the Anthospermeae and Paederieae, this distinct foetid odour or
paederoside is known only to occur in Theligonum (Theligoneae ;Wunderlich, 1971),
Oldenlandia hirsuta L.f. (Hedyotideae; Makino, 1909) and Pentanisia foetida Verdc.
(Knoxieae; Verdcourt, 1958).
Chromosome numbers: Published counts (compiled from Fedorov, 1969) and my
own unpublished karyological investigations confirm a base number of x = 1 1 for
most genera. Hamiltonia (Spenadictyon) suaveolens Roxb., with numbers of 2n = 22,
24, 26, and Nertera setulosa Hook.fil., with 2n =40, seem the only exceptions. The
chromosome number of 2n= 32 for Coprosma montana Hillebr. appears doubtful.
So far only diploid species have been detected in Nenax, Galopina, Carpacoce,
Crocyllis (Puff & Mantell, 1982), Phyllis, Putoria, Mitchella, Spermadictyon and Serissa
(Vij, Shekhar & Kuthiala, 1981; Fagerlind, 1937, with 2n = 22 for S.foetida Lam.).
Tetraploids alone have been found in Nertera and Plocama. Diploids and higher
polyploids have been found in Anthospermum, Coprosma and Leptodennis.
RUBIACEAE
369
Pollen : Pollen grains are generally 3-colpate or 3-colporate (Robbrecht, 1982).
Mature grains are trinucleate in Anthospermum, Nenax, Galopina, Carpacoce, Phyllis,
Paederia and Serissa, and binucleate in Leptodermis (see Wunderlich, 1971, and
references included therein).
Embryology : Little information is available. The archesporium is unicellular in
Phyllis (Fagerlind, 1936a) and Anthospermum, but multicellular in Putoria
(Fagerlind, 1936b) and Crocyllis (Puff & Mantell, 1982). Nuclear endosperm has
been recorded for Phyllis, Putoria (Fagerlind, 1936a, b) and Paederia (Hashmi &
Siddiqui, 1974). The suspensor is non-haustorial and uniseriate in Leptodermis and
Putoria (Fagerlind 1937, 1936b), biseriate in Paederia (Hashmi & Siddiqui,
1974), and multiseriate in Phyllis (Fagerlind, 1936a). The obturator of Phyllis is a
small, finger-like projection (Fagerlind, 1936a); the obturators of Putoria,
Leptodemzis (Fagerlind 1936b, 1937) and Crocyllis (Puff & Mantell, 1982) are very
similar to each other in having special elongated hair-like cells.
Distribution : The genera of the Anthospermeae and Paederieae are essentially
tropical or subtropical, with extensions to both the north and south. They are
relatively poorly represented in South America, but there is a marked
concentration of taxa in both southeast Asia and Africa.
Widely distributed genera are Paederia, occurring from southeast Asia to
Mauritius, Madagascar and tropical and subtropical eastern Africa, and to South
America, and the transpacific genus Coprosma (Oliver, 1935: map, fig. 3). Nertera is
even more widely distributed than the former and extends further away from the
tropics (van Steenis, 1962: map, fig. 14; in van Steenis, 1965: map, fig. 6,
Madagascar is incorrectly included in the distribution map). Anthospmum is
widely distributed in Africa and Madagascar and also occurs in the Yemen Arab
Republic. Neogaillonia occurs from southwest Asia to northwest Africa (Qarar,
1973: maps 1-4). Opercularia is found over most of Australia (Bentham, 1866).
Spermadictyon, Leptodermis and Serissa occur from southeast Asia to Japan. Putoria
occurs in the Mediterranean region, and east to Iraq (Ehrendorfer & SchonbeckTemesy, 1980).
Kellogia and Mitchella, consisting of two species each, show interesting southeast
Asian-North American disjunctions. K . chinensis Franch. occurs in the mountains
of the Yunnan Province in China (Smith, 1921; Handel-Mazzetti, 1936;
Dempster, 1975), K . galioides Torr. occurs in the western U.S.A. (Hitchcock &
Cronquist, 1973); M . undulata Sieb. & Zucc. occurs in Japan, Korea and adjacent
parts of the U.S.S.R. (Ohwi, 1965), M . repens L. in the eastern U.S.A. (Fernald,
1950) and south to Mexico.
The other genera have more limited distributions. In the northern hemisphere,
Phyllis and Plocama occur in the Canary Islands and Madeira, Aitchisonia occurs in
Afghanistan and adjacent parts of Iran and Pakistan and Pseudopyxis occurs in
Japan. In the southern hemisphere, .Nenax, Carpacoce, Galopina and Crocyllis are
found in southern and southwestern Africa, Pomax and Eleutheranthus in
southeastern and western Australia, Normandia in New Caledonia and Corynula
occurs in western South America.
CONCLUSION
From this comparison of the characters it seems that the genera of the
Anthospermeae and the Paederieae form a related, interconnected group. Most of
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C. PUFF
the characters overlap between genera and show much variation, but the floral
structures and pollination mechanisms allow a clear distinction, and a separation
into two groups. Rather than including all genera in a single tribe (i.e. the
Anthospermeae; viz. Baillon, 1880), it would appear most reasonable to place all
wind-pollinated genera in the tribe Anthospermeae s e w strict0 and all insectpollinated genera in the Paederieae. In this context, it is important to emphasize
that there is a clear demarcation line between wind- and insect-pollinated taxa;
there are no taxa showing gradual transitions from insect- to wind-pollinated
flowers or vice versa.
Darwin (1976) should be consulted for detailed information on correct names
and authors of the tribes and subtribes dealt with in the following section. Table 2
gives a summary of the most important characters of the genera.
TAXONOMIC CONSPECTUS
Tribe Anthospermeae Cham. & Schlecht. ex DC.
The tribe occurs in tropical, subtropical and temperate regions of the southern
hemisphere, with a few extensions to the north. Details of fruit structure,
pollination biology, particularly differences in protandry and protogyny, together
with a consideration of distribution patterns can be used to make subdivisions
within the Anthospermeae.
Subtribe Anthosperminae
Genera Anthospermurn, Galopina, Nenax, Carpacoce, Phyllis
This subtribe is restricted to Africa, Arabia in the Yemen Arab Republic,
Madagascar, the Canary Islands and Madeira and is characterized by dry,
dehiscent fruits with the exception of dry, indehiscent fruits in a few Nenax species,
and unisexual and/or protandrous hermaphrodite flowers.
There are many indications that both Nenax and Galopina (endemic to southern
Africa) may be closely allied to the large, widely distributed genus Anlhospermum, in
contrast to the Cape endemic genus Carpacoce which is more likely to have ancient
links with non African genera. The same is, perhaps, true for Phyllis, although it
might have closer links to Anthospermurn than Carpacoce.
Subtribe Coprosminae Puff, Subtribus nova
Plan tae cum Anthosperminis et Operculariinis floribus anemophilis congruentes,
sed fructibus carnosis vel carnulosis differt.
The subtribe agrees with the Anthosperminae and the Operculariinae in having
wind-pollinated flowers, but differs in having fleshy or more-or-less fleshy fruits.
Genera Coprosma, ,Normandia, Nertera, Corynula
This subtribe has a more-or-less south temperate amphi-transpacific distribution
(cf. van Steenis, 1962). It is characterized by the very fleshy to more-or-less fleshy
indehiscent to dehiscent fruits and the unisexual and/or protogynous
hermaphrodite flowers.
RUBIACEAE
37 I
The woody, monotypic, New Caledonian, genus Normandia is closely
allied to the large, woody genus Coprosma. The herbaceous genus .Nertera is
probably also closely allied and related to Coprosma. It has a wider distribution
than the latter. The South American endemic genus Corynula is most likely a
segregate and very close ally of Nertera.
Subtribe Operculariinae Bentham
Genera Opercularia, Pomax, Eleutheranthus
The subtribe is restricted to Australia and Tasmania and is characterized by the
umbel-like or compound head-like inflorescences. The partial inflorescences,
whose flowers have fused ovaries, give rise to infructescences opening by means of
an operculum. The flowers are unisexual and/or protogynous hermaphrodite.
I do not support the placement of Opercularia and Pomax into a separate tribe
Opercularieae as was done by Richard (1830) and de Candolle (1830) on the basis
of their infructescences. Details of floral morphology and protogyny rather suggest
a relatively close affinity to the Coprosminae.
The little known, monotypic genus Eleutheranthus probably belongs here. It is
supposed to have head-like inflorescences, in which the ovaries of partial
inflorescence are not fused. A thorough re-investigation is needed which may well
provide evidence for its inclusion in Opercularia.
There is no support for the assertion (Bremekamp, 1966) that these three genera
should be removed from the Rubiaceae and placed in the Dipsacales as a separate
family, the Operculariaceae (cf. de Jussieu, 1804).
Tribe Paederieae DC.
The basic floral structure of all genera in this group is quite uniform, although
size and shape of corolla tubes and number of carpels present, ranging from two to
five in number, are variable. The fruits, in parallel to the Anthospermeae, are
distinctly fleshy, more-or-less fleshy or dry.
The tribe occurs in (sub)tropical southeast Asia, with extensions to the south, the
Mediterranean and North America. Its genera are concentrated in the northern
hemisphere with the exception of Crocyllis, an endemic genus in the southern
Namib desert.
A subdivision of the Paederieae into discrete groups does not seem as simple and
clear-cut as in the Anthospermeae since less data are available for the tribe.
Therefore, it is preferred to only delimit and discuss a number of “species groups”
without giving them formal taxonomic recognition.
Group 1 : ..Veogaillonia, Crocyllis, Putoria, Plocama, Kellogia
.Neogaillonia, occurring in the Saharo-Sindian and Irano-Turanian regions
(Qarar, 1973), shows undeniable and, in some characters, very close
morphological similarities to the southern Namib genus Crocyllis (Puff & Mantell,
1982). This suggests a close affinity between the two genera and probably an
ancient xeromorphic palaeotropic common ancestor. The genus pair provides
another example of the close floristic links between the arid regions ofsouthwestern
Africa and the north (cf. Burtt, 1971; de Winter, 1971).
The Mediterranean genus Putoria is morphologically quite similar to
572
C. PUFF
Neogaillonia, but many xeromorphic features found in Neogaillonia are absent.
Flowers are similar, fruits are slightly fleshy but, as in Neogaillonia, separate into
two indehiscent mericarps. Distribution patterns of other (non-rubiaceous) taxa
(cf. Davies & Hedge, 1971 : 3. Irano-Turanian-Mediterranean connections) seem
to support, from a phytogeographical point of view, a connection between the
genera via a southwest Asiatic ancestral group.
The tetraploid, monotypic genus Plocama, although diffcying in its fleshy fruits,
appears to be affiliated to Neogaillonia on the basis of vegetative characters,
including xeromorphic features, and to Crocyllis on the basis of its short, more-orless funnel-shaped corolla tubes.
The flowers, particularly the short, more-or-less funnel-shaped corolla tubes, of
the disjunct herbaceous genus Kellogia are similar to those of both Plocama and
Crocyllis; Neogaillonia has more-or-less long-tubed flowers. Fruits of Kellogia, Crocyllis
and Neogaillonia are almost identical except for the presence of carpophores in some
species of Neogaillonia. These morphological similarities seem to point to a
relationship with the other genera of this group, although Kellogia is, on account of
its distribution (southwest China-western U.S.A.), more-or-less isolated.
Group 2 : Mitchella, Pseudopyxis
A purely herbaceous group, in which Mitchella shows an east Asian-North
American disjunction similar to that of Kellogia, while Pseudopyxis is endemic to
Japan. The two genera are similar both vegetatively and in their floral structure.
Mitchella has flowers in pairs (partial inflorescences) with fused ovaries. In
Pseudopyxis, the partial inflorescences are similarly few-flowered, but the ovaries of
adjacent flowers remain unfused. Mitchella has fleshy, indehiscent ‘double fruits’,
Pseudopyxis dry, dehiscent fruits.
The position of these two genera within the Paederieae and their links to other
genera is not entirely understood at present. Perhaps there is an ancient connection
to the genera of group three.
Group 3 : Spermadictyon, Leptodermis, Aitchisonia, Serissa
This group comprises woody genera occurring in central and southeast Asia
with the exception of Aitchisonia, which is endemic to Afghanistan and
neighbouring areas to the west and south. The group is characterized by flowers
with more-or-less long, funnel-shaped to cylindrical corolla tubes, more-or-less
large and broad-leaved foliage frequently with more-or-less triangular, single
stipular lobes. The fruits are variable. Spermadictyon and Leptodermis have fivecarpellate fruits dehiscing into valves; Aitchisonia has bicarpellate fruits splitting
into two dehiscent mericarps. Serissa has two- to three-carpellate fleshy indehiscent
fruits.
Spermadictyon and Leptodermis appear to be closely related on the basis of their
fruit characters, similar flower structure and general appearance. However, in
Leptodermis, the flowers and fruits are subtended by an often very conspicuous cuplike structure consisting of a pair of fused bracts which are not to be confused with
the carpophores of some Neogaillonia species. This cup-like structure, although not
as conspicuous, is also present in Aitchisonia; it may point to a closer affinity
between the two genera. Aitchisonia, in spite of its unique stalked glands, also seems
to have ‘cross links’ to Neogaillonia with its bicarpellate flowers and anther filaments
inserted at different levels in the corolla tube.
RUBIACEAE
373
The diploid genus Serissa seems to be isolated because of its fleshy, two- to threecarpellate fruits. However, the flower structure, vegetative characters and general
appearance are similar to Spermadictyon and Leptodemis. Spermadictyon is diploid, but
also shows aneuploidy ( n = 1 1, 12, 13) ; Leptodemis has di-, tetra- and hexaploid
species.
Group 4 : Paederia
Paederiu appears to be the most derived genus within the tribe because of its
climbing habit, deciduous stipules, anther filaments inserted at different levels in
the corolla tube, its winged endocarps and tetraploid chromosome number. It also
is the most widely distributed genus in the tribe, although the majority of species
occurs in southeast Asia.
RELATIONSHIP TO OTHER TRIBES
There are many indications to suggest that the monotypic tribe Theligoneae is
closely allied to the Anthospermeae (as originally suggested by Wunderlich, 1971,
on the basis of a careful comparison of a wide range of characters) and, more
remotely, to the Paederieae. I am certain that the genus Theligonum belongs to the
Rubiaceae in spite ofdoubts recently expressed by Praglowski (1973) and Nowicke
& Skvarla ( 1979). Their conclusions are based primarily on palynological
characters; the strong arguments in favour of its placement in the Rubiaceae
presented by Wunderlich (1971) are largely ignored.
Theligonum, like all Anthospermeae, has wind-pollinated flowers with distinct
sexual dimorphism. There is only one ovule per flower and the drupaceous fruit
has a hard endocarp enclosing the seed. The species are annual, or possibly shortlived perennials. Their leaves show a strong anisophylly in the upper portions of
the stems. A clear trend to anisophylly is also found in some species ofAnthospemum
and Curpucoce. The plants produce the foetid odour characteristic of many
Anthospermeae and Paederieae and the compounds asperuloside and
galiumglucoside are also present (Kooiman, 1971 ). The geographical distribution is
noteworthy in that it overlaps with that of several genera of the Paederieae;
Theligonum occurs in the Canary Islands, the Mediterranean area and east to
Japan (Ulbrich, 1934).
The similarities pointed out above seem to suggest that the three tribes
Paederieae, Anthospermeae and Theligoneae form a related group. Amongst
these, the Anthospermeae and Theligoneae appear to exhibit more ‘derived’
characters than the Paederieae. The Anthospermeae and Theligoneae are the only
two groups within the Rubiaceae with anemophilous flowers and this certainly
must be considered a derived feature in a large family with predominantly
zoophilous flowers.
This group of three tribes may, in turn, be linked with the large, more or less
pantropical, primarily woody tribe Psathureae ( = Psychotrieae) . Similarities in
details of floral structure are undeniable, and the drupaceous fruits with two to
many pyrenes suggest some relationship. However, available data are insufficient
to draw definite conclusions.
ACKNOWLEDGEMENTS
I am indebted to Dr E. Robbrecht for reviewing the manuscript and making
helpful suggestions for its improvment. I thank Professor S. Vogel for providing me
C. PUFF
314
with photographs of Paederia; Miss L. T . Dempster for field observations on
Kellogia; Miss D. Mantell for critically checking the manuscript, and Miss J. Dawe
for numerous chromosome counts. The CSIR/Pretoria (CSR/P75; 1979), the
Senate Research Committee of the University of the Witwatersrand/Johannesburg
(SRG/P990; 1978-79) and the Hochschuljubilaumsstiftung der Stadt Wien ( 1982)
provided research funds. SEM investigations were supported by the Fonds zur
Forderung der wissenschaftlichen Forschung in Osterreich (Projekt 3681) .
APPENDIX
Synonyms of Neogaillonia ; new combinations
Neogaillonia Linchevskii, Novosti Sistematiki Q-sshich Rastenij, 10: 226 ( 1973),
Gaillonia A. Rich. ex DC., Prodromus 4: 574 (1830), nom. illeg. (non
Gaillona Bonnemaison, 1828).
Jaubertia Guill., Annales des Sciences Naturelles, Paris, skrie 2, 16: 60 (1841).
TYPE: 3. aucheri Guill.
Choulettia Pomel Nouveaux Mathiauxpour la Flore Atlantique: 81 (1874) TYPE : C.
reboudiana (Coss. & Dur.) Pomel.
Pterogaillonia Linchevskii, Novosti Sistematiki Q-sshich Rastenij, 10: 233 ( 1973).
TYPE : P. calycoptera (Jaub. & Spach)Linchevskii.
Pseudogaillonia Linchevskii, Novosti Sistmatiki 5-sshich Rastenij, 10: 235 ( 1973).
TYPE : P. hymenostephana (Jaub. & Spach)Linchevskii.
SYNONYMS:
LECOTYPE:
N , olivieri (A. Rich. ex DC.)Linchevskii.
Neogaillonia calycoptera (Decne.) Puff, comb. nova
SYNONYMS: Spennacoce caly(co)ptera Decne., Annales des Sciences Naturelles, Paris, skrie 2,
2: 267 (1834). Gaillonia calycoptera (Decne.) Jaub. & Spach, Annales des Sciences
Naturelles, Paris, drie 2, 20: 86 ( 1843). Pterogaillonia calyoptera
(Decne.)Linchevskii,Novosti Sistematiki Q-sshich Rastenij, 10: 233 ( 1973).
TYPE:
Eygpt, Mt Sinai, Bout! 208, (P!, W!).
Neogaillonia stscherbinovskii (Linchevskii) Puff, comb. nova
Pterogaillonia stscherbinovskii Linchevskii, Novosti Sistematiki Vjkshich
Rastenij, 10: 234 (1973).
SYNONYM :
TYPE :
Iran, Ab-Gjarm, 40 km north of Bender-Abbas, Stscherbinovski (LE).
Neogaillonia crucianelloides (Jaub. & Spach)Puff, comb. nova
Gaillonia crucianelloides Jaub. et Spach, Annales des Sciences Naturelles,
Paris, skrie 2, 20: 86 ( 1843). Pterogaillonia crucianelloides (Jaub. & Spach)
Linchevskii, Novosti Sistmatiki Q-sshich Rastenij, 10: 235 ( 1973).
SYNONYMS:
TYPE:
Iran, Mt. Ghino, Aucher-Eloy 4681 (P!, W ! ) .
RUBIACEAE
375
Neogaillonia hpenostephuna Uaub. & Spach)Puff, comb. nova
Gaillonia hymenostephana Jaub. & Spach, Annales des Sciences Naturelles,
Paris, sirie 2, 20: 85 (1843). Pseudogaillonia hymenostephana (Jaub. &
Spach)Linchevskii, Novosti Sistematiki Vjkshich Rastenij, 10: 236 ( 1973). Gaillonia
humijiisa Jaub. & Spach, Annales des Sciences Naturelles, Paris, skie 2,20: 85 ( 1843).
SYNONYMS :
TYPE:(Oman) Maskat, Aucher-Eloy 4677 (K, P!, W!).
Neogaillonia aucheri (Guill.)Puff, comb. nova
SYNONYMS : Jaubertia
aucheri Guill., Annales des Sciences Naturelles, Paris, skrie 2, 16: 60
( 1841). Gaillonia aucheri (Guill.)Jaub. & Spach, Annales des Sciences Naturelles,
Paris, skrie 2, 20: 87 (1843).
TYPE:(Oman) Maskat and Iran, province Laristan, Aucher-Eloy 4680 (P!, W ! )
Neogaillonia reboudianu (Coss. & Dur.)Puff, comb. nova
SYNONYMS : Gaillonia
reboudiana Coss. et Dur., Bulletin de la Sociktt! Botanique de France,
2: 250 (1855). Choulettia reboudiana (Coss. & Dur.)Pomel, Nouveaux Mdteriauxpour
la Flore Atlantique: 8 1 ( 1874).
TYPE:Algeria, Oued-en-Nsa, Reboud 22 (P!).
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