Systematic Anatomy of Euphorbiaceae Subfamily Oldfieldioideae I. Overview

23 Pages·1994·11.6 MB·

by  W John Hayden

Number
2 Jensen
Volume 81 et 179
al.
,
1994 Leguminlike Proteins
3-46
Pp. in S. L. Jury, T. Reynolds, D. F. Cutler Securinega durissima F. Gmel., Bot. Gard. Stutt-
J.
& Evans The Euphorbiales: Chemistry, gart-Hohenheim
F. (editors),
J.
Taxonomy and Economic Botany. Academic Press, Strophioblachia fimbricalyx Boerl., seed stock Lehr-
London. Jensen
stuhl
1994. Synopsis of the genera and suprage- Vernicia montana Lour., seed stock Lehrstuhl Jensen
neric taxa of Euphorbiaceae. Ann. Missouri Bot. Gard.
33-144. Anacardiaceae
81:
Handbuch
WETTSTEIN, R. 1935. der Systematischen
Anacardium occidentale L., Bot. Gard. Basel
Botanik. Vierte umgearbeitete Auflage. Deuticke,
Spondias mombin Bot. Gard. Wien 1984
L.,
Leipzig.
Toxicodendron vernicijiua (Stokes) Barkl., Bot. Gard.
Wunderlich, R. 1968. Some remarks on the taxo-
Gottingen
nomic significance of the seed coat. Phytomorphology
310-311.
17:
Cneoraceae
Appendix. List of species used, with their sources. Cneorum tricoccon Bot. Gard. GieBen
L.,
Euphorbiaceae
Flacourtiaceae
Adelia riclnella L., seed stock Lehrstuhl Jensen Casearia gladiiformis Mast., seed stock Lehrstuhl Jen-
Aleurites montana (Lour.) Wils., Bot. Gard. Sri Lanka sen
1985 1978
Flacourtia inermis Roxb., Bot. Gard. Basel
Andrachne Wien 1988
colchica Fisch. et Mey. ex. Boiss., Bot. Idesia polycarpa Maxim., Bot. Gard.
Gard. Wien 1984
Antidesma
bunius (L.) Sprene., Bot. Gard. Salisburg Linaceae
1977
Hugonia orientalis Engl., seed stock Lehrstuhl Jensen
Baccaurea
tetrandra Muell. Arg., Bot. Gard. Basel
Linum usitatissimum Reformhaus Sattran, Bay-
L.,
Breyniafruticosa (Muell. Arg.) Hook., seed stock Lehr-
reuth
stuhl Jensen
Bridelia tomentosa Wien
Bot. Gard.
Bl.,
Moraceae
Chrozophora
tinctoria (L.) A. Juss., Bot. Gard. G6t-
tingen 1986 seed stock Lehrstuhl Jensen
Antiaris africana Engl.,
Croton tiglium L., Bot. Gard. GieBen 1984 Cudrania tricuspidata (Carr.) Bur. ex Lavall., Bot.
Dalechampia
scandens Wien 1988
L., Bot. Gard. Basel Gard.
Dry & Wien 1988
petes australasica Pax Hoffm., seed stock Morus rubra L., Bot. Gard.
Lehrstuhl Jensen
Eremocarpus
setigerus (Hook.) Benth., Bot. Gard. Ho- Passifloraceae
nolulu
Harms, seed stock Lehrstuhl
Adenia pechuelii (Engl.)
Euphorbia
lathyris L., Bot. Gard. Rostock
Jensen
txcoecaria
bicolor Hassk., Bot. Gard. Basel
Rosdorf
caerulea Terra,
Passiflora L.,
Wochtdion
glaucum
Muell. Arg., Bot. Gard. Basel
Hevea
brasiliensis (Willd. ex A. Juss.) Muell. Arg.,
Rutaceae
Bot. Gard. Sri Lanka 985
1
Homalanthus Wien Dictamnus albus seed stock Lehrstuhl Jensen
populifolius Grah., Bot. Gard. L.,
1984 japonicum Maxim., Bot. Gard. Basel
Phellodendron
ffura crepitans L., Bot. Gard. Sri Lanka 1985 Ruta graveolens L., seed stock Lehrstuhl Jensen
Jntropha curcas 1984 Zanthoxylum simulans Hance, seed stock Lehrstuhl
L., Bot. Gard. Bogor
Joanne.ua
princeps Lyon Arboretum, Ho- Jensen
Veil., Cutt.
nolulu
Mallotus
philippinensis (Lam.) Muell. Arg., Bot. Gard. Thymelaeaceae
Bogor
1984
Lehrstuhl Jensen
Daphne gnidium seed stock
L.,
Manihot
esculenta Crantz, Bot. Gard. Dusseldorf DDaavphhne mczcrcum seed M<>< k Lehrstuhl Jm••irSiCsen
L.,
^rgaritaria
discoidea Webster, Bot. Gard. Jensen
(Baill.) seed stock Lehrstuhl
Pimelea ligustrina Labill.,
Basel
Vercurudu
perennis
L., Bot. Gard. Basel
Tropaeolaceae
"rrantheum
ericoides Desf., seed stock Lehrstuhl
1989
Jensen Garden
Tropacolum majus L., priv.
°mphalea
trichotoma
Muell. Are., seed stock Lehr-
«uhl
Jensen Ulmaceae
Wlostigma
Jensen
quadriloculare F. Muell., seed stock occidental seed stock Lehrstuhl
L.,
Celtis
Lehrstuhl
1984
Jensen Gard. Siena
Bot.
Imus parvifolia Jacq.,
yUanthiu /
ralycinus Labill., Bot. Gard. Basel Zelkova carpinifolia (Pall.) K. Koch, seed stock Lehr-
"onjua
roxburghii
Wall., Bot. Gard. Basel
Jensen
Walts stuhl
trisperma
(Blanco) A. Shaw, Bot. Gard. Bo-
gor 1984
Urticaceae J™
P,
a arrnaria A Gra Webster Texas
1986°''' coll.
Y>
- moro„lrs Wedd., Bot. Gar,l
/,„,„,„,« <-«"><W»
;W.en 1988
Gard.
l ' ln "s communis L., Bot. Gard. Sri Lanka 1985 Parietaria effidnalu I... B«,t.
s 1984
P*um rwn Bot. (iard. ^rlit-Dald.-.n
sebiferum 1983 dioiec I..
' (L.) Roxb., Bot. Gard. Siena I
ANATOMY
SYSTEMATIC
OF Hay
John den
IV.
EUPHORBIACEAE
SUBFAMILY
OLDFIELDIOIDEAE.
OVERVIEW
1
I.
Abstract
The
biovulate subfamily Oldfieldioideae of Euphorbiaceae, characterized by spiny pollen, an otherwise apparently
is
diverse assemblage of mostly Southern Hemisphere trees and shrubs that traditionally have been allied with genera
of Phyllanthoideae and Porantheroideae sensu Pax and Hoffmann. Although fairly diverse anatomically, the following
structures characterize the subfamily with only a few exceptions: pinnate brochidodromous venation with generally
randomly
organized and
tertiary higher order venation; foliar and petiolar glands absent; unicellular or unbranched
uniseriate trichomes; latex absent; mucilaginous epidermis or hypodermis; brachyparacytic stomata; vessel elements
with simple perforation plates and alternate, often very small, intervascular pits; thick-walled nonseptate imperforate
tracheary elements; numerous narrow heterocellular rays; and abundant axial xylem parenchyma in diffuse to somewhat
banded patterns and often bearing prismatic crystals. Anatomically, the shrubby Australian ericoid genera forma
well-defined group with obvious affinities to the more arborescent Australasian genera, which show clear relationships
to each other; the African and neotropical genera bearing compound leaves form another distinct group; the remaining
genera are somewhat more isolated and seem to represent, in various cases, elements that are primitive within the
subfamily or elements derived from the group bearing compound leaves. Presence of theoid teeth and palmately
compound
leaves in Oldfieldioideae are features consistent with Dilleniid origin for Euphorbiaceae.
As a taxonomic member
entity the euphorbiaceous sub- dendron was shown to be a of Oldfieldioi-
&
family Oldfieldioideae Kohler Webster Hayden 1984,
dates deae (Hayden, 1977; see also et al.,
conceptually from the palynological studies of Punt and Hakki, 1985), an anatomical survey of the
(1962) and Kohler (1965) who noted the spiny entire subfamily was initiated (Hayden, 1980) to
pollen that characterizes the group; the assemblage assess relationships from data independent ot re
recognized by Based in
pollen structure was subsequently productive (including pollen) morphology.
formalized nomenclaturally new o
as a subfamily part on these anatomical studies, classification
(Webster, Webster -
1967). In essence, from
pollen characters Oldfieldioideae has been modified
circumscribed a group of genera that previously (1975) original proposal; two broadly similar clas-
had been preceding
assigned
to the biovulate subfamilies Phyl- one by G. Webster (the
sifications, L.
lanthoideae and Porantheroideae in the system of issue) and another by G. A. Levin and M-
&
Pax Hoffmann
(1931). Webster's (1975) Simpson are proposed elsewhere.
first (this issue),
classification of oldfieldioid genera contained
sev-
eral novel taxonomic
associations at variance with
History
&
earlier classifications, notably those of Pax Hoff-
mann
ft*
(1931) and Hutchinson from the
(1969). Euphorbiaceae
Classifications of
V*
Interest in the anatomy of Oldfieldioideae half of the 19th century Jussieu, 1824;
(e.g.,
stemmed
from
the need for comparative data 1836-1840) insight into rela-
to licher, offer little
aid placement of the problematic genus Picroden- among genera. At this
tionships oldfieldioid
dron Planchon Euphorbiacea^
(Hayden,
1977). Despite a number mative time in the definition of
kno<^
of
publications dealing with the anatomy genera
of old- (Webster, 1987) the few oldfieldioid
fieldioid genera (Appendix family*
2), detailed information were often scattered widely in the
for most of the subfamily is lacking. Once Picro- over, some were referred to other families.
thank •nneg*-
™»wu»i«,g anu
guiuance; n. iv. Airy Shaw, L. riickey, O. A. Levin, a. i«-
r •m J.
anda G. iL. Webster were
\ particularly helpful in providing specimens and/or comments; numerous others
specimens.
2 Department
of Biology, University of Richmond, Richmond, Virginia 23173, U.S.A.
Ann.
Missouri Bot. Gard. 180-202.
81: 1994.
Number Hayden 181
2
Volume
81
,
Euphorbiaceae Subfamily Oldfieldioideae
1994
Pseudanthinae (or Caletieae in the traditional/ included, using current nomenclature, Dissiliaria,
of
Longetia, Austrobuxus, Hyaenanche, Mischo-
narrow sense) were the first to be classified to-
undoubtedly because of their shared ericoid don, Oldfieldia, Piranhea Baillon, and only one
gether,
and common provenance nonoldfieldioid genus, Bischofia Blume, within gen-
xeromorphic habit in
grouped Micrantheum era 46-54. Although these plants constituted three
Baillon (1858)
Australia.
Bentham
Desf. with Pseudanthus Sieber ex Sprengel, to discrete groups in his conspectus, (1878)
and
which Agardh (1858) added Stachystemon Plan- expressed some doubt about their relatedness
Neoroepera and Petalostig-
chon, establishing the composition of Caletieae that he chose to associate
was followed in all subsequent studies until the ma with other phyllanthoid genera. Pax (1890)
much had Bentham;
addition, of Neoroepera Muell. Arg. by Koh- treated oldfieldioid genera as
first,
Tetracoccus Engelm. ex C. Parry, however, was
(1965) and, now, the inclusion of Austral-
ler all
&
proposed by Levin & Simpson included, and Pax Hoffmann (1931) added Peta-
asian oldfieldioids as
Androstachys Prain, and Aristogeiton-
(1994, this issue) and Webster (1994). During the lostigma,
intervening years, however, the use of cotyledon ia Prain to various subtribes consisting largely of
width as the primary criterion for subdivision of oldfieldioid genera. Perhaps the first clear indica-
& genera can
the family (Mueller, 1866; Pax, 1890; Pax tion of the relatedness of oldfieldioid
on Euphorbiaceae,
Hoffmann, 1931) relegated the genera of Pseu- be found in Pax's (1925) essay
reproduced
danthinae (as Caletieae) to the Stenolobeae, a small which includes the phylogenetic tree
much
Although of the detail in this
group of Australian xerophytes with narrow cot- in Figure 1.
may
be challenged seriously in
yledons. Their somewhat Sten- phylogenetic tree
isolated position in
show
knowledge, does old-
olobeae minimized association of Pseudanthinae with the light of present it
minus Pseudanthinae)
other genera (as usual,
oldfieldioid genera. fieldioid
from other biovulate
Other comprising a clade distinct
oldfieldioid genera similarly suffered early
Euphorbiaceae.
taxonomic
assignments that delayed consideration
Hurusawa's (1954)
of genera,
of In the disposition
relationships with the rest of the subfamily. Bail-
Euphorbiaceae essentially follows
of
lon (1858) submerged Podocalyx Klotzsch within classification
&
Pax Hoffmann (1931) and thus contains
the phyllanthoid genus Richeria Vahl, a disposition that of
gen-
followed well into this century. Oldfieldia Benth. no new insights on classification of oldfieldioid
phylogeny and because
& Pax's 925)
Hook. was temporarily considered sapinda- era. In light of ( 1
f.
penological data available
ceous (Mueller, 1866; Baillon, 1878). Worse, the of its disregard for the
Hutch-
1962; K5hler, 1965),
time (Punt,
relationships of Picrodendron were long obscured at the
was a retrograde de-
b a (1969) classification
y series of misassignments both the generic inson's
at
genera were widely distrib-
and
velopment; oldfieldioid
familial (Hayden
levels 1984).
et al.,
with three tribes
2 biovulate tribes,
Aside from genera uted in five of 1
Caletieae sensu the
stricto,
elements.
and phyllanthoid
mixing oldfieldioid
°1 Oldfieldioideae were scattered widely in Muel-
was however, to associate
ler's ( 1 866) treatment of the family for the Prod- Hutchinson the first,
gen-
Kuhlm. with other oldfieldioid
romus; Paradrypetes
Mischodon
Thwaites was included with
jiniovulate
genera, an error of era.
following earlier
doubts
Shaw (1965) entertained serious
Kaillon Airy
(1858), was banished
Oldfieldia to
still
genera
in
of several oldfieldioid
^apindaceae, about inclusion
and
the remaining five oldfieldioid
he proposed Andro-
Accordingly,
genera Euphorbiaceae.
in his treatment were each assigned to dif-
accom-
uncertain relationships, to
ferent stachydaceae, of
subtribes of Phyllantheae. Greater cohe-
Androstachys; he recognized Pi< roden-
eness modate
^ apparent
is in a later publication of Baillon
Euphorbiaceae;
87 from but allied to
For draceae as distinct
8). example: was
Choriceras Baillon in-
Calaenodendron
Aristogeitonia,
Jluded within and he viewed
' Caletieae; Austrobuxus Miq. (as Bur-
and Piranhea
"'""« Mischodon, Oldfieldia,
Standley,
Baill.), Longetia Baillon ex Muell. Arg.,
between these
^lostigma an intermediate position
occupying
F. Muell., and Hyaenanche Lambert as
Airy
1972, 1973).
Shaw, 1966,
°rrned
a (Airy
sequence; and Dissiliaria F. Muell. ex families
?R * accept an isolated tax-
°wed Shaw (1983) continued to
fo,, Richeria (which included the old-
Pseud-
e Stenolobeae (including
Wioid onomic position for
Podocalyx). com
(See Webster 987) for
1
(
group the re-
jnentary on but he did, consistently,
anthinae),
Baillon's peculiar "serial" system to
m
genera of Oldfieldioideae in
,cate maining Australasian
relationships.) elements were
Additional
**"By informal or tentative
several of his
accreted adjacent tribe, in
nucleus
to this loosely defined
Shaw, 197b,
° Airy
systems
f (e.g.,
^fieldioid classification
genera. Bentham
(1880), in his treat-
men
of 1980b, 1983).
< Euphorbiaceae the Genera Plantarum,
in
^
182
Annals
of the
Missouri Botanical Garden
WIELANDIINAE
ANDRACHNINAE
PSEUDOLACHNOSTYLIDINAE
PETALOSTIGMATINAE
ANTIDESMINAE
Petalostigma
AMANOINAE
DRYPETINAE
Drypetes
Heywoodia
DISCOCARPINAE
Lingelsheimia
Neoroepera
Tetracoccus
UAPACINAE
BISCHOFFIINAE
PAIVAEUSINAE
SAUROPODINAE
Aristogeitonia
Oldfieldia
Piranhea
TOXICODENDRINAE
PHYLLANTHINAE
Androstachys
Hyaenanche
DISSILIARIINAE
GLOCHIDIINAE
Dissiliaria
Longetia
.
Austrobuxus)
Unci
Mischodon
Figure
Relationships published
1. of subtribes of Phyllantheae, after Pax (1925). Right-hand clade the earliest
is
phylogram of Oldfieldioideae. Generic & Hoffmann (1941*
composition of subtribes, where indicated, follows Pax
oldneldioid taxa are in bold italics. Other than Pseudantheae, Podocalyx subgenus of Richer* [Ann-
(treated as a
desrmnae])
is the only element of Oldfieldioideae known Pax and phylogram.
to not included in the
Description,
Circumscription, and y
Distribution
un
generally
of two are
to five carpels; styles
Plants of Oldfieldioideae are woody, ranging from vided; each locule nouses a pair of pendulous ana
low depressed shrubs a
to tall trees. Latex is absent ropous ovules. Seeds are often carunculate
and
vestiture Repor
consists of unicellular or uniseriate usually possess copious endosperm.
~
J*
trichomes. Leaves =
are
alternate, opposite, or whorled 12 Pseudanthus
for (Hassall,
and may 1976),= J
be simple or palmately compound; n
petiolar for Picrodendron (Fritsch, 1972),
and laminar an
glands are absent; stipules may be Mischodon and Tetracoccus (Hans, 1973),
W*
present or absent. Leaves may = = &
be strongly reduced 48 Mischodon (Sarkar
24, 2/i for
in xerophytic number of x
species. Plants are
typically dioe- 1980) collectively suggest a base
cious. Flowers are apetalous, and most
lack discs. 12 for the subfamily.
n,.
Stamens
range from three many. o
to Pollen The comprehensive classification
is bi- first
nucleate, with four to many brevicolporate to po- fieldioideae (Webster, 1975) included 21 gener
rorate apertures; the exine lacks a foot layer, bears but several changes in composition have occ
a discontinuous
interstitium and thick perforate since then. Two newly discovered genera,
tectum
with prominent V['h\nnbee
supratectal spines (Levin malo Capuron ex Bosser (1976) and
&
Simpson, na
1 994, this volume). Gynoecia consist Airy Shaw & B. Hyland (Airy Shaw, 1976),
Hayden 183
Number 2
Volume 81
,
Euphorbiaceae Subfamily Oldfieldioideae
1994
Two Appendix 2 has been incorporated
been referred to the subfamily. enigmatic gen- cations listed in
Croizatia Steyerm. (Webster et al., 1987) into the following account as appropriate.
era,
have been
Paradrypetes (Levin, 1992), as-
and
signed to Oldfieldioideae following discovery of di-
Leaf Architecture
flowering material and, especially, after
agnostic
Kairothamnus For the most part, leaves are either simple or
study of pollen from these plants.
compound; however, interesting transi-
Shaw (1980a) and Scagea McPherson (1985) palmately
Airy
morphologies do exist. Leaves of Oldfieldia
have been segregated from Austrobuxus sensu lato, tional
3-8
Canaca are palmately compound, with leaflets; those
as has Guillaumin, if only tentatively
Picrodendron, and Piranhea
Celaenodendron,
(Webster, 1994). In recent years Radcliffe-Smith of
compound and sim-
both palmately
are trifoliolate;
has redefined generic limits for several Oldfieldioi-
occur Aristogeitonia; and leaves of
deae. He has included Paragelonium Leandri with- ple leaves in
Parodiodendron Hunz. are unifolioate, as evi-
in Aristogeitonia (Radcliffe-Smith, 1988), he seg-
denced by the minute stipels at the apex of the petiole
regated Stachyandra R.-Sm. from Androstachys
1969) and by the frequent disarticu-
(Hunziker,
to accommodate several Madagascan species bear-
same
compound and lation of the lamina at the point (Fig. 4).
ing leaves (Radcliffe-Smith, 990),
1
Micrantheum, which occur in
leaves of
he expanded Pseudanthus include Stachyste- Finally,
to
have been
in-
groups of three (to five),
mon (Radcliffe-Smith, 1993). The number of gen- alternate
compound
palmately
of a
now terpreted as the leaflets
era stands at 27. (Solely because of the re-
common are
petiole,
by of their
which, loss
cency of Stachystemon herein leaf
its reclassification, is
1858); alternatively,
on the stem (Baillon,
treated as a distinct genus.) sessile
has been attributed to foliate
unique phyllotaxy
Genera mostly mono- or this
of Oldfieldioideae are
&
1913; Webster Miller, 1963).
(Griming,
20 stipules
oligotypic. Austrobuxus, with perhaps species,
few
the
margins are entire,
For the most part, leaf
is the most speciose, followed by Pseudanthus (sev-
cuneatus
Austrobuxus
being
species
en The exceptional
species) and Petalo stigma species).
(six &
Beuzev.
Shaw, A. swainii (de
Shaw) Airy
number (Airy
total of species referable to the subfamily
Shaw, Choriceras tricorne
White) Airy
is estimated at 86, approximately percent of the C. T.
1
muel-
Shaw Dissiliaria
Airy (Figs. 7, 8),
number (Benth.)
total of species in Euphorbiaceae.
Paradrypetes ilicifoha
ex Benth.,
Oldfieldioideae largely a southern hemisphere leri (Baill.)
is
G. Levin
subintegrifolia
1986), P.
Kuhlm. (Levin,
group. Only three genera occur areas not derived
in
Parry (only
dioicus
Tetracoccus
from 1992),
(Levin,
the breakup of Gondwanaland; these are Ce-
&
Oilman.
Coville
and ilicifolius
laenodendron, from western Mexico, Picroden- some leaves), T.
spaced spinose
irregularly
Paradrypetes possesses
dron, from the Bahamas and Greater Antilles, and
Tetracoccus
Teeth of
1986, 1992).
(Levin,
tetracoccus from Mexico and the southwest United teeth
median
subfamily:
the
are the largest in
States. Eight genera and 16 species occur in the ilicifolius
from the looped
derived
are
ew of these teeth
World, six genera and 16 species occur in veins
brochidod-
prominent
are
there
secondary veins;
Africa and
Madagascar, one monotypic genus oc-
median
and
the tooth itself;
within
curs romouslike loops
in southern and Lanka, and 13
India Sri
which the
in
the apices,
extend nearly to
genera and 52 No genus veins
species are Australasian.
caps,
appear possess apical
examined to
°f specimens
Oldfieldioideae has a bicontinental distribution,
remaining
Teeth of the
young leaves.
although Androstachys and Aristogeitonia occur at least in
end-
of a vein
much consisting
hth smaller,
are
species
in Madagascar and
Africa.
Significantly,
of leaf tissue.
protrusion
a small
ing in
«
caps are also visible
apical
deciduous
however,
Mater
IALS Choriceras tricorne (Figs.
and
swainii
Austrobuxus
thu
Oldfieldioideae
morphology in
Tooth
This report based on examination 7 8)
is first-hand &
Wolfe,
(Hickey
type
theoid
91 conforms with the
°« leaf and 87 wood specimens of 61 species
spinose teeth,
noninformative
Excluding
1975).
Oldfieldioideae. Specimens examined are listed
m
Phy
found lan-
m type
only
are the
Appendix teeth
1 The specimens available represent theoid
. genera Drypetes \ ahl
the
SOme specifically in
*° degree except thoideae,
genera of the subfamily
all ^duced
theoid
and Bischofia;
Wall.,
roizatia Putranjiva
and Paradrypetes; should be noted
it 1986).
Aporuseae <L.-vin.
present in
at ,eaves of Voatamalo and wood of Kairotham- teeth are also
most frequent fes-
n" Sca Venation always pinnate, y
s gea, and Stachystemon are lacking in this is
' ~ addihon
In
h,dy. i^'^ b«•r»ochidodromous (Figs. 2, 3).
tooned -
Previous concerning
anatomical
literature
. v„e onff Loonnl*ietia
however, leaves \
***ra brochidodromy,
of to
Oldfieldioideae provided by the publi-
184 Annals
of the
Missouri Botanical Garden
v
-
* * I -
..
I . I
ii^^lii
-
L
§«&!
liBH?
JMN
$
— *
{BaumannBode
FIGURES 2-8.
Leaf
architectural features of Oldfieldioideae. 2. Austrobuxus rubiginosus ^
—
- - c
15010), cleared leaf, bar 1 cm. 3. Celaenodendron mexicanum {Ortega 6367), cleared leaf, bar
4. Parodiodendron marginivillosum {Hueck from base ° \
— 469), cleared leaf, note disarticulation of petiole o.5
= - -
mm. bar
bar 5 Picrodendron baccatum
5. (Gillis 6963), note lack of vein orders beyond tertiaries,
Hayden 185
Number
2
Volume 81
,
Euphorbiaceae Subfamily Oldfieldioideae
1994
New
tendency toward eucamptodromy whereas Picrodendron (Fig. 5), and Piranhea (all
exhibit a
An-
Petalostigma, Picrodendron, World genera with trifoliolate leaves) plus
those of Dissiliaria,
extremely rare
and Tetracoccus tend toward reticulodromy. Leaves drostachys; this character state is
comm.)
whole Hickey, pers.
Australian genera of Pseudanthinae (Cale- in the dicots as a (L. J.
of the
synapomorphy
tieae sensu stricto) exhibit a continuum of increas- and is thus a potentially robust link-
presumably a consequence of ing these genera. Areoles are well developed in the
ing disorganization,
xeromorphy: Neoreopera leaves are weakly fes- genera with compound leaves (Figs. 5, 12) plus
in
Parodiodendron
tooned brochidrodromous; in Micrantheum one can Dissiliaria, Mischodon, (Fig. 6),
and Podocalyx; otherwise, areoles are imperfectly
brochidodromous venation as well
find vestiges of
arranged randomly,
as secondary veins that recurve and ramify or form developed. Areoles are usually
Celaenodendron and somewhat
a weak reticulum; in Stachystemon the range is but are oriented in
Piranhea, and Dissilia-
Picrodendron,
from reticulodromous to kladodromous; and in oriented in
and mostly simple
Pseudanthus leaves are kladodromous (Fig. 9). ria. Veinlets are usually present
all
branching varies widely,
degree of
branched; the
Primary veins are mostly of moderate size, but, or
branched leaves with
may be most highly in
especially in xeromorphic leaves, they range tending to
and higher order veins.
Hyaenanche, low regularity of tertiary
to stout and massive, those of
e.g.,
Androstachys but
(Fig. 12),
are absent in
Tetracoccus, and the Australian Pseudanthinae (Fig. Veinlets
upon
may appear to be present superficial
they
9). Intersecondary veins are usually absent, al-
examination because of the columnar sclereids lo-
though one or two simple intersecondaries per in-
each areole (see below).
tercostal area occur occasionally in Austrobuxus cated in
and Mischodon, and routinely in Croizatia (Levin,
1986), Oldfieldia, Paradrypetes (Levin, 1986),
Anatomy
Leaf
Piranhea, and Podocalyx. Intersecondary veins
about two-
have been observed in
Trichomes
are frequent in several primitive Phyllanthoideae,
Many
spe-
genera of Oldfieldioideae.
namely, Amanoa Aubl., Blotia Leandri, Hey- thirds of the
mature
although
with age,
glabrescent
woodia and are
Sim, Petalodiscus Savia Willd., cies
Baill.,
and
Parodiodendron,
Androstachys,
Wielandia presence leaves of
Baill. (Levin, 1986); their is
example, are clearly
Petalostigma, for
thus species of
postulated to be primitive for Oldfieldioideae.
simple unicellular
Trichomes are either
Tertiary vein patterns are usually random reticu- pubescent.
than
consisting of less six
generally
'ate; ramified patterns occur Hyaenanche, spe- or uniseriate,
in
emer-
epidermal
Pseudanthinae,
Within
cies of Austrobuxus, and those species of Austra- cells.
four-celled
from small papillae to
lian Pseudanthinae gences range
that possess distinguishable
packed
The densely tri-
trichomes.
tertiaries; there is a slight tendency toward trans- uniseriate
which
johnsonii Prain,
Androstachys
verse chomes of
Paradrypetes
tertiaries Parodiodendron,
in
terminal
and long curly
Levin, 1986), Petalostigma, and Podocalyx; and consist of a short basal cell
I
by Alvin (1987) to func-
he have been suggested
l tertiary veins of Piranhea have a slight ten- cell,
moisture (mists
atmospheric
dency absorption of
to form orthogonal patterns. Marginal ve- tion in
and
extremely en-
otherwise
the
nation and drizzles) in
is incomplete (Fig. or looped (Fig. 6) in
1 1)
possesses
Oldfieldia
southern Africa.
rnany vironment of
genera, "hemmed" with a fimbrial vein in
Androstachys, ar-
those of
the genera trichomes similar to
with compound leaves (Figs. 10, 12)
between these
some degree of relationship
P'us Dissiliaria and Mischodon, or dominated by guing for
d.fTerences in
manifest
mass two genera despite their
ive intramarginal vein most Australian
p in
Depending on the spe-
morphology.
Pseudanthinae reproductive
A
(Fig. distinctive furcate-fla-
9).
simple
Austrobuxus are either
!ate form Hyae- trichomes of
of marginal venation occurs in cies,
Parodiodendron
is
nanche. (malpighiaceous).
or bifurcate
strongly
t|. possessing
, subfamily in
"ign-order unique within the
venation varies considerably within
Rao & Raju (1985)
margins (Fig. 6).
Oldfieldioideae. leaf
Perhaps most notable the oc- ciliate
is
and glandular hairs in
currence uniseriate, stellate,
reported
of order
ultimate which vein
reticula in
un.ovu-
reminiscent of
not characteristics
Dissiliaria,
distinguishable, Celaenodendron,
a feature of
^^^jjf^
Chonrcras
7.
fnm.
°- Parodi«>den dron margl nivillosum (Hueck 469)—margir of V man
i " {r or s.n.
I.
lrir°rne {*-F-o.r.m,.a„n„ s»..n„.\/, tiooootihn aait mmaarrggiinn ooif ctliewairceud l»e«a•f,, bar = 25- 0 urn. *• 10
•
dermal bar = 100 Mm.
apex,
glandular
section through margin, note
tooth at leaf
186 Annals of the
Garden
Missouri Botanical
^^
Figures 9-12. Leaf architecture of Oldfieldioideae Pseudanthus orientalis {Clemens 44092), klad
9.
(Ortega
fl
= mexicanum
venation, massive primary and 250 Celaenodendron
intramarginal veins, bar /im. 10.
™ »ith
= jul— nched
6367), fimbrial vein (far right), bar 250 11. Petalostigma banksii {Perry 1981), b JfLaton
= Lao
Research
swollen ultimate tracheids, bar 100 /im.- 12. Androstachys johnsonii {Wellcome Chemical
=
s.n.), well-developed areoles with columnar mm.
sclereids, veinlets absent, bar
1
!®*^
late euphorbs that would be unique to Oldfieldioide- or chamber thus formed contain*
cell
Expa
^
ae; however, neither trichomes nor their bases were which common in other genera as well.
is
detected Australian
in the leaves available to me. of the mucilaginous layer in
The epidermis
epidermis is fundamentally uniseriate. How- thinae greatly distends the adaxial [
mucJag
ever, a single layer of hypodermis has been re- 18, 19); while the grossly expanded
^
^
Paradry arat
ported for petes (Milanez, 1935) and some layer may well be an artifact of P re P !T
ied
v%i(
su
of
or all epidermal cells of the Australasian genera derived solely from the lower portions ^
^j
ar
multicellu
have a horizontal partition resulting in a poorly to epidermal and not thickly ^
cells is
erroneo
well-defined hypodermis (Figs. 17-19). The inner depicted by Gaucher (1902), whose
Number Hayden 187
2
Volume 81
,
Euphorbiaceae Subfamily Oldfieldioideae
1994
^^^^^—^JrJ^^
Figure, 13-16. -13. Longetia
Leaf anatomy
of Oldfieldioideae.
^^ « (McKee 2352) surface view-of ^stom"ate t^rorn
from - v*
cross section, bar 20 ^n.-14. Scagea oligostemon
Z,
through
—late
.T* falUof subsidy cells
anticHnal
\7}:™\
9
15. Austrob subdivided
!r
base
SawajM*
R and
ichena,
*;pre.a, n Phyllanthus L.,
10 has been reiterated by others (e.g., lem, «*«
>""£*%£
& &
'^alfe & Chalk, 1950). In
So
Chalk, 1950; Raju Rao, 1977). calfe
gh
F^gs.
ar are general! s,r
-.no mucilage waUs of ep.dermal eells y
has been detected in Aristogei-
^"'^T
be.ng
£«. except.ons
Cefawodend™, Won, fWAeo. the only
Afcc and 1 4, 1 5),
fl^a,
memte™
J»
***«/>*. Cehenodendron, (all
- MucUaginous known
epidermis in '
is DtofcW wbch^hese*al
»
e-l A« V and
crodendreae),
Phyllanthoideae, example, Atf«
for
rmal sec
»"
.ntngu.ngly, ,,,,,,.
4n„We wavy
Sma L ^ BU Lanoa, «ac have a outline;
., Cdamodm**n
reveal
""'* and
Uur., Aristogeitoni,,
of
flridefia WilU., /fyeromma Fr. Al- tions
188
Annals
of the
Missouri Botanical Garden
—
Figures 17-21. (Bau
Leaf anatomy
and wood parenchyma buxoides
of Oldfieldioideae. 17. Longetia
Bodenheim Pseuda
5605), leaf cross section, note chambered mucilaginous epidermal cells, bar = 100 /mi.- 18. " y
orientalis {Wilson prominent
679), leaf cross section, note grossly expanded mucilaginous hypodermis and
P
marginal veins, bar = 100 Mm.-19. Micrantheum hexandrum (McGillivray 3196), cross section of Pr,m* n
,,
^
note grossly expanded = JT^
mucilaginous
hypodermis, bar 100 m.-20. Hyaenanche globosa (Godfrey
M
cham^
-
cross section of leaf, bar 100 Mm.-21. Piranhea USw normal and
{Ducke 31485),
trifoliala
s.n.,
crystalhferous axial xylem parenchyma = =
PF
cells, bar 15 phloem
urn. fibers.