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37
BLUMEA
tuberous epiphytes
The
A
335-340
(1993)
taxonomic
history
C.R.
Department
of Plant
Sciences,
Oxford
of the Rubiaceae 6:
of
the Hydnophytinae
Huxley
University,
OX1
Road, Oxford,
South Parks
3RB,
U.K.
Summary
first described
Rumphius
recognize
the
briefly placed
further
the two
elsewhere and
genera of the
major
and Jack described
work,
confused,
and
until
admirable
Beccari’s
added. Valeton’s revision
genera were
Hydnophytinae in 1750, but
Hydnophytum
was
in
Myrmecodia
Linnaeus
1823. These
did not
genera were
monograph (1884-86)
when
three
sadly incomplete.
Introduction
The taxonomic
history
is
a
presented
as
this
volume), Myrmephytum
(Jebb,
to
the
first
The first person
Rumphius
to
& Jebb, in
Becc.
& Jebb,
Jack
and
1991b),
1991a)
background
&
(Huxley
to
Jebb,
Becc.
Squamellaria
valid publication
who worked for the Dutch East India
His classic work 'Herbarium Amboinense'
until 1750.
published
of the
work,
Joseph
did
on
(ms)
(Beaglehole,
The third
by
till
names
prepared during
for the
of fire and
ant-plants,
described the
long
plant
as
published,
were not
the latter halfof
blindness,
and
Nidus formicarum
pre-Linnean. Linnaeus, although
not
niger
aware
it into his system.
after J.D.
Epidendroides
and the
ant-plants
tetrandra,
of this
discovery
Hooker had described it
encounter
with
ant-plants
descriptions
of
(1823)
Sumatra,
described
Hydnophytum formicarum
niger.
No
was
as
in
Queens-
but his
manu-
remained
plant
Myrmecodia
described
and identified it with
specimens
have existed,
or to
of
exist
also tragic: William Jack's
Hydnophytum
Island off
to
was
ravages
Georg
was
Ambon in the Moluc-
at
beccarii
1962).
fire and his
micarum
joint
therefore
are
after his premature death. Jack
known
the
Company
the first voyage of the 'Endeavour' found
and illustration
unrecognized
ruber,
incorporate
not
Banks
land. Solander
delayed by
Rumphius'
and Nidus formicarum
script
the
concerns
prep.), Myrmecodia
(Huxley
& Jebb,
Hydnophytinae (Huxley
series since it
describe the ant-inhabited tuberous Rubiaceae
the seventeenth century,
lost
in this
1991).
Pre-Linnean
cas.
(Huxley
of the
genera
separate paper
Jack
Hydnophytum
of the
and
Myrmecodia
Myrmecodia
Rumphius'
Nidus
specimens
were
were
published
tuberosa from Nias
formicarum
ruber. He
from Sumatra and identified it with Nidus for-
ant-plants
today.
associated with these three authors
are
BLUMEA
336
Two years after Jack's
They
neither Rumphius
described it
as
while H.
oblong leaves,
Jack described H.
H.
was
species
a new
H.
-
feature mention-
a
in
western
he said had
Java and
short-petioled
leaves. Since, however,
ovate
the dis-
having short-petioled elliptic-ovate leaves,
as
Later J.D. Hooker
tenuous.
Java, which he placed
western
spiny tuber,
species
had subsessile
formicarum
Logani-
sank H.
(1881)
into
montanum
formicarum.
Gaudichaud
(1830)
described collections made in the Moluccas
age of the 'Uranie'. He
codia he described
also sank the genus
mis which
without reference
wrote
Mirmecodia [sic]
as
M. tuberosa Jack the
name is
A. P. de Candolle
H. formicarum Jack and his
to
indicate
a
M.
species
according
to
H.
nizing
For
a
M.
armata
formicarum
while there
DC. is
Jack and H.
was
fruit of what is in fact
were
Becc.
later
(=
as
montanum
used the
name
which he identiHe created
Rumphius.
a
of
species
followed
(1833)
in
in
(M.
and H.
a
tuberosa Jack,
thinking
name
recog-
but
most
authors
apparently
meant
he had found numerous seeds
Hydnophytum with
loranthifolium (Benth.)
but
Jack,
hispida,
two
pyrenes. He revived
in it
species
two
Hydnophytum by Beccari (1885); they
oblonga)
He
Blume.
mentioned M.
an error
with
M. iner-
De Candolle followed Blume in
genus in the Rubiaceae and described
a
placed
L.
correct.
(1834)
M. echinata. Then Bentham made
a
name
unfortunately
Myrmecodia
ruber of
formicarum
peace; Henschel
followed De Candolle. Richard
Lasiostoma
of
spineless species
plant
illegitimate.
the
published
voy-
Myrme-
Blume) and also Gaudichaud's M. echinata. Since Gaudichaud's
superfluous,
was
and hence
include Blume's collection from Java
armata to
Freycinet's
on
Blume. The
collection from the Moluccas.
new
the genera, but
(1830) re-separated
or
but since he identified the
into Mirmecodiaand
fied with M. tuberosa Jack and Nidus
new
echinata,
Sprengel
to
nomenclaturally superfluous
Hydnophytum
comprised
M. inermis Gaudich.
in
a
Hydnophytum
This
montanum.
from
species
in the
position by Blume (1826/27).
from
Jack. He also found
formicarum
somewhat
transferred both
(= Strychnos L.)
proper
but noted that it had
Jack,
nor
their
to
of Myrmecodia
specimen
a
in M. tuberosa
tentatively
tinction
1993
2,
Lasiostoma Schreber
to
however, returned
were,
Blume also collected
by
No.
publication Sprengel (1825)
their genera in the Rubiaceae
ed
37,
1824-1872
Confusion
aceae.
Vol.
Becc.
H.
are
(=
L.
(1843).
These
oblongum (Benth.)
loranthifolia)
from
New Ireland and New Guinea respectively.
Miquel (1857) recognized
the
pre-Linnean
num
name
Blume's
Blume; he also observed that these
Myrmecodia
name
niger
for these
two
In 1858 Asa
American
M. echinata Gaudich.
He
described
Myrmecodia
no
small
two
was
an
of Arts and
new
species,
imberbis which
mellaria. Gray
is
Gray presented
account
one
about the
by
one or
the
to
be
to
this
by
following
In
Jack
de Candolle's name M.
was
not
published
wrote
Fiji
to
the
until 1860.
and the other
to a new
and
ar-
followed Blume.
longiflorum
Beccari
species
monta-
include Blume's collection
existing literature
old
H.
distinguished.
M. inermis,
(1861) Miquel
Hydnophytum
two
but he transferred
of Seemann's collections from
later transferred
rather confused
obscurity
was
to
ignoring
Sciences, though
was
scarcely
name
collections. In his flora of Sumatra
Academy
Hydnophytum
are
superfluous
from Java and Gaudichaud's from the Moluccas,
mata
of
from H. formicarum Jack
species
two
he removed De Candolle's
instead. He retained the
species
two
Nidusformicarum
genus
sadly
of this genus and
Squa-
"There
respecting
C.R.
the distinction between it and
Seemann, apparently
up."
vitiensis, but
M.
name
More confusion
on
of
still
was
collections
arise.
to
be
to
lected
an error
by Zippelius
removed it from the
tion of
Jack and H.
Baron
next
von
From the
for H.
Another red
montanum.
and referred
ant-plant
montanum
by referring
by Zippelius
report
Mueller
was
of
(1871)
collected
ant-plants
referred them to
‘Genera Plantarum’
the distinction between
well
floral
as
M. imberbis A.
(1873)
ones.
They
In 1874 Beccari entered the
(1884)
to
herring
and
also
a
by
with
arena
to a new
Myrmecodia again
myrmecophytic
he
the
saw
genus
and that the
zippelianum.
and M.
Australia.
armata.
succeeded in
(1873)
stem
clarifying
and tuber char-
that M. inermis A.
Gray (meaning
describe it.
not
of Myrmecodia selebica which
Myrmephytum
Becc. Von Mueller in
them to
Papua, assigning
on
1875
H. formicarum
the Andaman Is-
Jack.
(1880) suggested
should be
two
no
flowers. No-one
ideas. In 1880 Britten mentioned M.
mistake for M. inermis. In the
(1886)
that
regarded
Hydnophytum
as
be sunk
section of the
a
non-
L. He also mentionedLasiostoma Bentham of which
Uragoga
but found
specimens
were
(1884-86)
description
a
genus
Beccari
Cape York,
Myrmecodia, noting
recognized
H. formicarum
as
ovatum
Blume which ap-
called H.
(1885)
Nares from
distinct genus, but did
Baillon in his 'Histoire des Plantes'
into
that he
his H. lanceolatum col-
was
echinata. In 1877 Kurz found Hydnophytum
lands and identified it
H.
species
ellipticum
H. formicarum
listed D'Albertis' collections from southern
Myrmecodia
a new
Cephaelis laevigatum;
Beccari
to
Hydnophytum
Gray) belongs
he later transferred
and
plant
same
clear
1861 the
also mentionedanother incomplete collec-
Miquel
genera.
to
H.
Bentham and Hooker in their 'Genera Plantarum'
acters as
in
Blume and concluded they
to
from ?New Guinea which Beccari
Zippelius'
The
the
was
named
Miquel (1869)
distinct. He confused later writers
pears
out, this
to
ought
published
and De Vriese from Temate. He discussed the sim-
by Teijsmann
ilarity between H. formicarum
not
Gray's description,
Gray (1862) pointed
as
337
Hydnophytinae
which the Dutch botanists
Hydnophytum
unaware
the
history of
longiflorum.
had called H.
based
Taxonomic
Huxley :
same
glabra
seems to
have followed Baillon's
from Borneo, but this
year Beccari cited
two
species
was
of
probably
Myrmecodia
a
in
Count d'Alberts' book 'New Guinea': M. albertisii and M. muelleri.
J.D. Hooker in
carum
lected in
that
1881 took the step of
Jack. Home in the
one
to
species,
omissions,
morphology
of the
of these
Odoardo Beccari
were not
H. wilsonii,
Squamellaria
sinking
year listed
Fiji, though descriptions
of these
transferred it
merous
same
was
two
H.
montanum
species
published
not a
of the
Blume into
Hydnophytum
till 1883
by
group,
in
gave
a
a
paper
H. formi-
he had col-
Baker who realized
typical Hydnophytum
in 1886. Franz Antoine
taxonomy
of
;
history,
indeed Beccari
albeit with
mainly devoted
nu-
to
the
as
the
plants (1882).
published
his monumental
study of
ant-associated
plants
second part of his three-volume work 'Malesia' between 1884 and 1886. This volume
is subtitled 'Piante
Ospitatrici; ossia piante
descritte ed illustrate da O. Beccari';
plants,
but the
Hydnophytum
was
greatest part
was
based
familiar with the
concerns
on
type
formicarie della Malesia
it deals
extensively
e
della
Papuasia
with other ant-associated
the Rubiaceae. His revision of Myrmecodia and
considerable
material in
knowledge
Europe.
of the
He added
plants
16
in the field and he
species
to
the
existing
BLUMEA
338
of Myrmecodia and 29
two
he removed
form the
to
the six of
to
new
Vol.
genus
phytum
accommodate his
to
not
believe
six-merous corolla and
ters,
the
of
species
ants
From Beccari
Valeton
to
After Beccari there
species
but
with
a
species
species
of
in
finds.
of
Archipelago.
from the British
of
specimen
ted
new
and
of
piecemeal
work with
in
began
colony
of
(1927).
But he
of
J.D.
species.
new
plant
distinctive
a
which had been
and moved the
an error
a
careless
and
colony
two
of
as
of
Drake
H.
and
Myrmecodia
species
led
exploration
new
Hydnophytum
species
a new
moment
new
species
activity
1915)
of
one
from
Hydnophytum
described
(1895)
costatum;
this
was
a
spot-
to
of
Hydnophytum
at
an
elderly
Kew,
and
and Elmer
(1911,
of
man;
he
and he left undescribed
ant-plants
saw
cies in New Guinea followed
of three and
Hydnophytum fared a
has taken up
of
two are
little better. His
two
of
by descriptions
photographs
of these
from New Guinea
in his
of
key
were not
Myrmecodia
von
(1911,
Papuasien'
seven
and
consists of
new
a
and H.
key
to
of which
species,
16
but
spe-
frag-
species
new
have survived and P.
(H. vaccinifolium
specimens.
on
only fragmentary,
all that survived. His
manuscripts
names
from the
the material neither at Florence
names
many of the collections he described
account
1913, 1934)
Myrmecodia
be sunk.
species
ailing
in 1905 and
ceased but American authors
and four of
Hydnophytum
named several
later lost in Berlin. Thus his
(1983)
the
was
species
named
(1907, 1908, 1913,
eight species
already
was
nor
Unfortunately,
ments
of
when he undertook the Rubiaceae for Lauterbach's 'Flora
(Beccari's)
were
German
added
(1892)
In
Papua.
a
(1898)
Hooker
Almost all of these will have
Valeton had
1912a, b)
Merrill
total of
Philippines.
charac-
Beccari also recorded
description
named three
1894)
in 1913. Then German
two more
a
had
Myrmedoma,
vegetative
1924.
earnest:
described
genus
Myrmecodia beccarii,
but found that he had made
Schumann and Lauterbach described a
Rechinger
genus,
Myrmecodia.
Scyphiphora hydrophyllacea (Rubiaceae)
Pittard in
by
monotypic
in Australia. This
while Schumann
Myr-mecodia,
as
Myrme-
1898.
Warburg (1891,
the Bismarck
new
of the tubers.
Northeastern New Guinea had become
to new
a
Banks and Solander. Schumann in 1888 honoured Beccari
Hydnophytum,
Psychotria
to
phase
a
vegetation
published by
not
?wil-
(1927)
was
from mangrove
seen
origin
honoured Beccari's
(1886)
Hooker
and H.
be ant-associated
to
recognized
from each genus
Gray
present in the ant-plants, discussed theirrelationship with
and considered the
plants,
Squamellaria
but these
differed in
genera
large bracts,
resembling respectively Hydnophytum
the different
species
imberbis A.
previously described Myrmecodia selebica. This satellite
with his other
genus Myrmephytum, along
a
One
Hydnophytum.
observation of tubers. He also
Gray's
1993
2,
Squamellaria (M.
sonii Home ex J.G. Baker). He did
he missed
No.
37,
van
of
Royen
crassicaule).
After Valeton
In the
species
same
These three
viously
In
year that Valeton's work
collected by Brass
species
described
by
of
on one
Hydnophytum
Hooker and
1940 Fosberg described
a
appeared,
of the many and
were,
Beccari,
new
Moore
(1927)
important
described three
Archbold
however, all very close
which Moore did
species
of
not
Hydnophytum
to
Expeditions.
species
apparently
from the
pre-
examine.
Solomon
Huxley :
C.R.
the
history of
Taxonomic
339
Hydnophytinae
Islands, and in 1942 Bremekamp published H. inerme (Gaudich.) Bremek.
gaudichaudii
combinationfor H.
type for H.
lections
came
of these
species
Finally,
of
from
areas
species
see
missed.
were
a
existing types,
num-
of the col-
Many
and
variability
range
therefore often under-estimated.
was
in 1967 A.C.
the flora of
Smith, studying
and called it S.
Squamellaria
the
and 15 of Hyd-
Myrmecodia
not
before and little since, the
explored
not
did
they
as
of
species
seven
and identities with earlier
relationships
a new
was
Jack.
Perry published
from the Brass collections. But
nophytum
specimen
as
which
the type for M. inermis Gaudich. since this
not
was
formicarum
In 1945 Merrill and
ber of
Becc.
gaudichaudii
included H.
species
Becc. But the Gaudichaud
found
Fiji,
distinct
a
species
major.
CONCLUSION
The taxonomic
was
the
This led him to
the available
its
types.
His
plants
as
the
styly
of only the
commonest
existing specimens
synthesis
or
of these
first-hand over much of theirrange and
at
is the
only
which
one
Beccari
understanding';
unique symbiosis
interest in the
ochlospecies H. formicarum,
edge
to
an
covers
subgeneric classification, attempts
and the association with
led
by
treatment
discusses
interpret
we
the
study
distribution,
what
of the group shows little
history
author stimulated
only
to
to
the group
plants.
examine all
throughout
define the variation of
and which considers hetero-
Valeton, Merrill, and Perry had first-hand knowl-
ants.
with their limited acquaintance
species; this, together
their work
little more than
being
listing
a
of
new
with
collections.
ACKNOWLEDGEMENT
I
would like
to thank
F. White.
supervisor
my
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