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History
Gesneriaceae were one of the last major families to take
on its present taxonomic form. Originally two separate families were
recognized: the New World “Gesneriaceae”, and the Old World
“Didymocarpaceae” = “Cyrtandraceae” (described almost at the same time). In
1839, the famous British botanist Robert Brown united the two families into
a single one and this was universally accepted. Bentham (1876) and Fritsch
(1893/94) were the first to publish overall accounts of Gesneriaceae in its
wide sense. C.B. Clarke (1883) wrote an important revision of the Old World
“Cyrtandreae”.
In 1963, B.L. Burtt, from the Royal Botanic Garden
Edinburgh, presented a new classification of Old World Gesneriaceae. He
recognized that this group (subfamily Cyrtandroideae) shares a conspicuous
and uncommon feature: the anisocotylous seedling. He distinguished five
tribes: Cyrtandreae, Trichosporeae, Didymocarpeae, Klugieae, and Loxonieae.
Additional tribes (Saintpaulieae, Rhynchotecheae, Ramondeae, Titanotricheae)
were then suggested by Russian and Chinese authors, but were not generally
accepted.
Not included in subfamily Cyrtandroideae were the
Australian and SW Pacific genera Fieldia, Lenbrassia, Negria,
Coronanthera
and Depanthus. These were referred to tribe Coronanthereae and, together
with the temperate South American tribe Mitrarieae (Mitraria, Sarmienta,
Asteranthera) added to the New World Gesnerioideae (Burtt 1963).
In 1983, The German-American botanist Hans Wiehler united
the tribes Coronanthereae and Mitrarieae and raised them to a subfamily of
its own (Coronantheroideae). The morphological character linking its genera
is that the nectary is adnate to the ovary. Thus, the last formal
subdivision of Gesneriaceae (Burtt & Wiehler 1995) recognized three
subfamilies:
-
Coronantheroideae,
comprising the single tribe Coronanthereae,
-
Gesnerioideae,
comprising the tribes Gloxinieae, Episcieae, Beslerieae, Napenatheae, and
Gesnerieae, and
-
Cyrtandroideae
(now to be called Didymocarpoideae for priority reasons), comprising the
tribes Klugieae (now Epithemateae),
Cyrtandreae, Trichosporeae and Didymocarpeae.
The advent of cladistics in Gesneriaceae research
(starting with Boggan 1991) and molecular systematics (starting with Smith &
Carroll 1997) provided a novel set of data and showed that the former
morphology-based classifications were in need of considerable modification.
Present
classification of New World Gesneriaceae
Regarding
the New World Gesneriaceae, molecular systematics has roughly confirmed the
tribes formerly distinguished, and has recognized the new tribes Sinningieae
and, most recently, Sphaerorhizeae. The list of tribes thus reads:
-
Coronanthereae
-
Beslerieae
-
Napeantheae
-
Gesnerieae
-
Gloxinieae
-
Sinningieae
-
Sphaerorhizeae
-
Episcieae
It is
important to note that the Coronanthereae indeed are closely associated with
the New World Gesneriaceae. They apparently do not represent a link to Old
World Gesneriaceae, though they contain several genera distributed in the
Old World (New Caledonia, New Zealand, NE & SE Australia). Moreover, it
became clear that Beslerieae and Napeantheae are the most primitive tribes
and closely related to each other. They form a group of its own opposed to
all other tribes.
Present
classification of Old World Gesneriaceae
Molecular
systematics has proceeded less in Old World Gesneriaceae and this group is
in fact more difficult to classify. It became clear that the previous
subdivision into tribes cannot be maintained. Two groups can be clearly
separated, the former small tribe Epithemateae (here informally referred to
as Epithematoid Gesneriaceae) and the large remainder, comprising the genera
of Didymocarpeae in which the members of Cyrtandreae and Trichosporeae are
nested at various places (here informally referred to as Didymocarpoid
Gesneriaceae). In the Didymocarpoid Gesneriaceae, a succession of the
following morphological-geographical groups can be observed (Weber 2004). At
the present state of knowledge an application of formal names would not make
much sense:
-
Epithematoid Gesneriaceae (= the former tribe Epithemateae)
-
Didymocarpoid Gesneriaceae
-
Basal Asiatic + European genera
-
African genera
-
Advanced Asiatic genera
-
Genera with twisted
capsular fruits
-
Genera with straight
capsular or indeshiscent fruits
The basal
Asiatic-European genera include the Asiatic rosette plants Jerdonia and
Corallodiscus as well as the three European genera Haberlea,
Ramonda and
Jancaea, also of rosette habit. The group comprises, however, also tall,
somewhat woody plants such as Tetraphyllum, Boeica and Rhynchotechum (with
indehiscent fruits). Rhynchotechum was previously asscociated with Cyrtandra
and placed in tribe Cyrtandreae. Another peculiar member of that group the
tiny unifoliate plant Platystemma.
The African
genera form a well-defined, compact group. Streptocarpus (with twisted
fruits) seems to be the most primitive genus, from or in which the other
genera seem to have originated.
The Advanced
Asiatic genera are split into two major groups: one with twisted fruits (a
parallelism to the African genera?) and the large remainder with straight
fruits. This is the group which comprises the species-rich and well-known
genera Henckelia, Chirita, Didymocarpus, Aeschynanthus etc.
The
molecular data provide clear evidence that a number of genera are
artificial. In particular, this confirms the view that the genus Chirita is
an assemblage of largely non-related species, held together only by the
technical character of a stigma with upper lobe reduced and the lower one
being bilobed. The molecular data show unambiguously that this character is
indicative of relationships only to a limited extent. |