ISSN 0085-4417

NUYTSIA

Volume 3 Number 1
1980

WESTERN AUSTRALIAN HERBARIUM— DEPARTMENT OF AGRICULTURE

Cover

A representation of Nuytsia floribunda (Labill.) R.Br. ex FenzI — the Western Australian
Christmas Tree. The journal is named after the plant, which in turn commemorates Pieter
Nuijts, an ambassador of the Dutch East India Company, who in 1627 accompanied the
“Guide Zeepard” on one of the first explorations along the south coast of Australia.

ISSN 0085-4417

NUYTSIA

VOLUME 3 NUMBER 1
1980

WESTERN AUSTRALIAN HERBARIUM,
DEPARTMENT OF AGRICULTURE,
SOUTH PERTH, WESTERN AUSTRALIA

CONTENTS

A revision of the Australian species of Salicornieae (Chenopodiaceae). By Paul G.
Wilson 3

Publication date of Volume 2, Number 6 154

Editor

A. S. George

Western Australian Herbarium, George St., South Perth,
Western Australia 6151.

Nuytsia Vol. 3 (1) 1980

3

A revision of the Australian species of Salicornieae (Chenopodiaceae)

By Paul G. Wilson
Western Australian Herbarium

Abstract

Wilson, Paul G. A revision of the Australian species of Salicornieae (Chenopodiaceae).
Nuytsia 3, 1 : 1-154 (1980).

A taxonomic revision is presented for the Australian representatives of five genera
in the tribe Salicornieae (Chenopodiaceae). Three of the genera, Pachycornia Hook.f.,
Sderostegia P. G. Wils. gen. nov., and Tegicornia P. G. Wits. gen. nov. are endemic to Aus-
tralia. Hahsarcia P. G. Wils. gen. nov. is found in countries bordering the Indian Ocean
while Sarcocornia A. J. Scott is widely distributed in both the northern and southern hemis-
pheres. The genera Pachycornia (which is monotypic) and Sderostegia (with five species)
have their flowers arranged in axillary 3-flowercd cymules in which the lateral flowers are
male and the central hermaphrodite. Tegicornia (with one species) is dioecious and has
solitary flowers. Hahsarcia (with 23 species) has either hermaphrodite or female flowers
and differs from the genus Arthrocnemum (from which it is here segregated) in having only
one stamen in each flower and in lacking sclereids in the chlorenchymous tissue. The genera
Hahsarcia, Tegicornia, Sderostegia, Pachycornia, and Tecticornia appear to be more closely
related to each other than they are to Sarcocornia or to any of the extra-Australian members
of the Salicornieae.

Introduction

The articulate members of the Chenopodiaceae with fleshy stems and reduced flowers
have been variously separated from the rest of the family at the levels of tribe or sub-
family; they have even been treated as a separate family. While the Salicornioideae
have been generally recognised as a discrete assemblage, the circumscription of the genera
has been the cause of considerable debate. This has been partly due to the poor material
that the ta.xonomists have had available to them, and partly due to the paucity of characters
on which to base both the genera and species. With the availability of fresh or pickled
material of some representatives, and good dried specimens of others, it has been possible
to more clearly delineate the genera and recognise many of their constituent taxa.

In this paper five of the six genera recognised as being present in Australia are revised.
A revision of the genus Tecticornia has already been published (Wilson 1972).

Morphology

Stem: The articulate branches characteristic of the Salicornioideae have generally been
assumed to be made up of a stem axis surrounded by the decurrent abaxial portions of
leaf bases (Tolken, 1967). An alternative structure has been proposed in which it is
assumed that the succulent sheath represents a true cortex and that the leaves are reduced
to small lobes (Fahn, 1963). Whichever view is correct it is apparent that there is no
boundary of demarcation in the sheath that could suggest a (division into material of
foliar and stem origin.

4

Nuytsia Voi. 3 (I) 1980

No detailed investigation of the anatomy has been undertaken in the present study;
it is apparent, however, that there are considerable differences in the morphology of the
stem between different members of the Salicornioideae. In all species the young internodes
have a central stele surrounded by a water tissue which consists of thin-walled parenchyma
cells tilled with a (usually) saline sap. Around the water tissue are one or more layers
of chlorenchymous palisade cells covered by an epidermis. The epidermis contains
numerous stomata which are easily visible with a hand lens; these always have their pores
aligned transversely to the stem axis. A single leaf trace arises on each side of the node;
it divides into three, the central branch passing upwards tow'ards the ‘leaf’ tip while the
lateral branches curve downwards into the water tissue below the node. Each trace
branches repeatedly to form an anastomosing network in the aqueous tissue, the degree
of anastomosis varying between the different species. In all Australian genera of the
Salicornioideae the decurrent branches from one leaf trace also anastomose w'ith those
of the opposite trace so as to form a complete network around the stem; this union takes
place even in Pcichycomia triandni where the chlorenchyma is absent in two opposite
vertical strips which delineate the apparent decurrent foliar tissue. The ultimate branches
of the vascular network terminate either as simple vascular strands or as club-shaped
clusters of tracheids (tracheoids) situated just beneath the chlorenchyma. The former
is the case in Pachycornia, Sarcocornia, Scleroslegia tenuis, and Halosarcia iiuiica, while
the latter occurs in Sclerostegia arhuscula, S. ciisarliculata, Tegicornia and most species
of Halosarcia. De Fraine (1912) stated that in Sa/icornia the vascular network arising
from one leaf-trace does not anastomose with that of the opposite leaf-trace, a situation
which is contrary to that observed in Australian members of the Salicornioideae and
is also contrary to the observations of Fahn and Arzee (1959) on species of Artlirocnemiiin,
Sarcocornia. and Salicornia. In the branches of all three traces the phloem tissue faces
towards the outside of the stem and the xylem the inside.

The palisade tissue normally consists of two or more layers of cells which may be
sharply differentiated from the aqueous tissue (as in Sarcocornia) or may merge into it
through a zone of cells of an intermediate type (as in H. halocnemoides). In one species
complex both the stele and the tissue of the palisade layer differ markedly from that found

Figure 1. Halosarcia indica: longitudinal section through outer tissues (a— epidermis, b~palisade tissue
with clear ‘passage’ cells between groups of chlorenchymous cells, c— thick-walled chlorenchymous iso-
diametric cells, d — parenchymatous cells with vascular strands).

Paul G. Wilson, Australian Salicornieae

5

in all other Australian members of the Salicornieae; this complex consists of H. inclica sensu
lato, which is here taken to include Arthrocnemum ciliolatiim Ung.-Sternb., A. hidens
Nees, and A. leiostaebyum (Benth.) Paulsen. In this species the palisade tissue is of
a single layer and is made up of compact clusters of chlorenchymous cells set in a reticulum
of cylindrical colourless cells of similar shape to (but somewhat thicker than) the chlor-
enchymous. A cluster of chlorenchymous cells is situated beneath each of the stomata.
Within the palisade tissue is a single layer of close-fitting thick-walled isodiametric chlor-
enchymous cells which form a continuous sheath between the palisade and aqueous tissues;
this layer frequently becomes lignified with age (Fig. 1). Although this type of palisade
tissue appears to be unique in the Salicornieae it is found in other tribes of the Cheno-
podiaceae, for instance the Salsoleae. In Salsola kali the palisade tissue of both the
leaf and stem consists of a single row of cells within which is a single row of thick-walled
isodiametric cells. This is underlain by parenchymous tissue which, in the leaf, has a
reticulate network of vascular strands beneath the isodiametric layer; in the stem there
is no cortical vasculature. (See also Carolin el al. 1975.)*

A reticulum of colourless cells in the palisade tissue is also found in many other species
of Halosarcia; these are called ‘passage cells’ by Tolken (1967) but their function is not
clear.

Figure 2. Sclereids in chlorenchyma: A — Sarcocornia qtdnqiiefiora showing evenly thickened sclereid;
B — Sarcocornia hlackiaiia, showing spirally thickened sclereid (a — epidermis, b — chlorenchyma, c — sclereid,
d — aqueous tissue).

* This is a typical Kranz type structure— R. C. Carolin and S. W. G. Jacobs pers. comm. A more
detailed account will be published by these authors,

6

Nuytsia Vo). 3 (1) 1980

c

bract

trace

\\

Yf

i

o

o

stele 1 V '

o

'P

O

abaxial '
trace

V

o o
o o

Q O

\

c

o

o

O

E

/

/

O

o

cymule

stele

O

Figure 3. Halosarcia lepidosperma, vasculature of inflorescence: A — Abaxial view of vasculature of triad
of flowers; B — L.S, (medial) of flower; C — G — T.S. at levels indicated in figure B. (all diagramatic).

Paul G. Wilson, Australian Saiicornieae

7

Figure 4. Tegiconua uniflora, vasculature of female flower: A — Optical longitudinal section; B — Abaxial
view of vasculature of a one-flowered cymule; C and D — T.S, through flowers at different levels (a
bract trace b — stele of stem, c — stele of axillary one-flowered cymule, d— vestigial lateral traces, e— upper
bract, f— perianth, g— ovary, h— lower bract, i— trace to vestigial staminode, j— trace to abaxial base of
perianth, k — ovule).

8

Nuytsia Vol. 3 (I) 1980

In Sarcocornki, sclereids occur in the palisade tissue (Fig. 2): these may be evenly
thickened or they may have a spiral thickening. Sclereids have not been observed in other
Australian Salicornieae, but they do occur in both species of true Arihrocnenmm, i.e.,
A. gkuiciini (Delile) Ung.-Sternb. and in A. suhterminak (Parish) Standley which are
respectively Mediterranean and North American in their distribution.

Inflon’scence: A detailed discussion of the inflorescence in the Salicornieae will, it is hoped,
be the subject of a future paper; for the present it is sufficient to indicate that it consists
of decussate axillary clusters of flowers, each cluster being sessile within a common bract;
there are no secondary bracts (or bracteoles). The axillary clusters are in most species
arranged in a spike-like structure and the whole then called a pseudo-spike (although
actually a thyrse). In several species of both Halosarcia and Sclerostegia the spike very
soon becomes intercalary in position through vegetative growth at its apex; it may then
eventually have the appearance of a normal branch. A few species, e.g. Tegicornia uniflora
and Sclerostegia mecitfliosa, have the flowers (or flower clusters) in the axils of “foliage”
leaves and no obvious compound inflorescence is formed.

In the majority of the Salicornieae the axillary clusters are three-flowered. Each cluster
has generally been accepted as constituting a cymule (a dichasium); due to the congested
arrangement of the flowers, however, and the fact that they are sessile on the inflorescence
axis, the vasculature of the cymules is difficult to determine. In Halosarcia kpidosperma
(in which the verticils are relatively widely separated) it is apparent that an axillary stele
composed of many vascular .strands forms a dorsiventrally compressed, somewhat globular
reticulum and terminates in the central flower of the cymule (Fig. 3). It gives rise to a
pair of smaller lateral steles which terminate in the lateral flowers of the cymule. Each
of the steles gives rise to a (bract?) trace on the abaxial side of their respective flowers,
and also to a pair of lateral (bracteole?) traces, while the traces themselves divide to form
a pair of club-shaped spurs or tracheoids (the possible nature of these traces is discussed
later). Since the vasculature of the axillary flower cluster consists of a central and a pair
of lateral steles the classical view' of its structure being that of a cymule is accepted.

In Tegicornia uniflora only one flower is present within each of the leaf-like bracts (Fig.
4). This flower is served by an inflorescence stele which terminates in the flower and
gives rise to a pair of small globose lateral steles that are embedded in the axial tissue
below the base of the perianth. These lateral steles are presumably homologous to the
lateral steles of the cymule vasculature in Halosarcia and therefore it is probable that
the solitary flower in Tegicornia is the result of reduction from a three-flowered dichasium.

In Sarcocornia hlackiana and 5. quinqueflora five or more flowers are found in the axil
of a bract. This cymule is similar to that found in Halosarcia except that each of the
lateral branches of the cymule stele gives rise to a monochasium (in fact a drepanium)
composed of from two to several flowers.

TIowers: The flowers are surrounded by succulent bracts or leaves and in some cases
are completely hidden by these. The perianth is gamotepalous, at least in the bud, and
bears 2-4 terminal lobes. The number and position of these lobes is normally constant
within a genus. In Sarcocornia there are two lateral lobes, an outer adaxial lobe and
usually an abaxial lobe (which may be very reduced); in Halosarcia there are two lateral
lobes and an abaxial lobe which may be either inside or outside the two laterals; in Tecti-
cornia there are two lateral lobes; in Tegicornia there are two lateral lobes and an inner
abaxial lobe; in Pachycornia there are two lateral lobes; while in Sclerostegia there are
two lateral lobes with or without an inner abaxial lobe.

The imbrication of the lateral lobes follows a regular pattern; the situation can be most
readily observed in Halosarcia where the flowers are always in triads and, depending on
the species, the outer florets always have one or other of two possible types of imbrication.
In some species, e.g. H. hahcnenioides and H. pergramdata the outer florets usually have
the outer lateral lobe overlapping the lobe nearest to the central floret (Fig. 5C-D), in
others, e.g. H. indica the reverse is the case (Fig. 5A-B). The central floret has its imbric-

Paul G. Wilson, Australian Salicornieae

9

Figure 5. Imbrication of perianth lobes showing opposite triads: A and B — Halosarcia prtiiiiosa; C and
D — Halosarcia pergraniilata.

ation in either symmetry (i.e. overlapping to the right or to the left) and varies up the
spike in a manner which has not yet been determined; however, the opposite triads have
the imbrication of all three florets as mirror images of each other (Fig. 5). The lobes
of the perianth are best observed in the bud since in some species the orifice is small and
the emergence of the relatively massive anther may cause the perianth to split or the lobes
to be displaced. The type of imbrication is somewhat correlated with the shape and
texture of the perianth and the shape of the seed; thus those species in which the perianth
has a rounded apex, remains soft, and in fruit tears away from its sunken base, typically
have the imbrication as in H. halocnemoides, while those in which the perianth is obtuse
to acute, becomes variously hardened in fruit, and falls entire from the spike, usually
have the imbrication as in H. iiidica.

The va.sculature of the perianth has not been fully clarified and is itself involved in
the larger problem of the vasculature of the axillary cyme. However, certain general
principles are apparent. In Sarcocornia qidnqiieflora and S. hlackiana there are four
traces which separately pass from the cymule stele to the base of the perianth, one to
each of the four sides (Fig. 6). They terminate in the tissue made up of the fused portions
of the perianth base, the upper or lower bracts, and the radial plates which extend between
the flowers from the inflore.scence axis (see paragraph on Fruit); as there is here no differ-
entiation between the floral and axial (or bracteal) tissues it is not possible to determine
whether the traces penetrate the perianth itself or remain in the axial or bracteal tissues.
In Halosarcia lepidospenna there are three traces, one of which passes to the abaxial
margin of the perianth base and the others to its sides; again the traces terminate within
the indefinite axial-perianth tissue (Fig. 3). In all ca.ses the traces bifurcate at their ends
to form club-shaped spurs (tracheoids). The three traces in Halosarcia flowers may
indicate the presence of vestiges of bract and bracteoles, especially as the traces do not
actually penetrate the free perianth tissue; however, since four traces are present in Sarco-
cornia the probability is that in both genera they are perianth traces, related in number
and position to the perianth lobes (four in Sarcocornia, three in Halosarcia). In Tegicornia
uniflora only one trace has been observed and this is abaxial; in this species, however,

JO

Nuytsia Vol. 3 (I) 1980

the inflorescence stele is extremely condensed and remains of the other traces may have
been lost (Fig. 4). Perianth traces have not been observed in such species as H. imiica
and H. pniinosa where the flower is not ‘sunk’ into the inflorescence axis and this possibly
provides support for a bracteole origin of the traces since in these species there is no non-
floral axial tissue which might represent vestigial prophylls.

Figure 6. Sarcocornia qiiimiueflora, vasculature of inflorescence; A— L.S. through central flower- B to
F— T.S. through central flower at different levels (a— bract traces, b— stele of axillary cymule c— trace
to adaxial side of flower, d— trace to abaxial side of flower, e— traces to lateral side of flower f— ovarv
g -perianth, h— upper bract, i— lower bract, j— stele of axis).

Paul G. Wilson. Australian Salicornieae

The perianth at anthesis may be succulent or membranous; at the fruiting stage it may
have changed in texture to become spongy, brittle or pithy. The ovary wall may initially
be membranous or fleshy, while in fruit it can be brittle or woody; in some of those species
with a membranous pericarp this may fuse with the perianth and is very difficult to discern.

Fruit: In Sarcoconiia the fruitlets (perianth with pericarp and seed) tear away from the
inflorescence leaving a number of deep depressions in its broad axis; the.se apparent
depressions appear to be caused by the intrusion of the axial tissue between the flowers
to form plate-like septa to which the basal sides of the perianth are fused. These septa
remain on the spike (with a portion of adherent perianth) since the abscission zone develops
between the free perianth and the plates. The axial portion of the perianth is here ex-
tremely thin and breaks down with the ripening of the seed; thus, when the fruitlets fall,
the seed protrudes from their base w ith the remains of the pericarp adherent to it, or the
seed may for a short time remain attached to the axis while the perianth and pericarp
fall away.

In some species of Halosarcia (e.g. H. halocnciiioiiles) the fruitlets also tear away from
the radiating plates of the spike axis, exposing the seed and leaving behind a portion of
the perianth and pericarp; but in the majority of species (e.g. //. pniinosa) the fruitlets
drop entire leaving only a small scar or depression on a comparatively slender axis; in
these species there are no interfloral axial septa but the flowers are attached along their
narrow base to the inflorescence axis. In several species of Halosarcia (e.g. //. syncarpa)
the fruitlets are not immediately shed when mature but remain attached to the axis and
enclosed within the bracts while the spike apex continues growth vegetatively; the fruitlets
are eventually cast off, along with the surrounding bracts and cortical tissue, on the form-
ation of periderm and the onset of secondary thickening in the spike axis (which has now
become part of the branch).

In Pachycornia the seed is surrounded by the bract bases and the broad inflorescence
axis both of which become lignified; it presumably germinates only on the decay of the
infructescence.

In Sclerostcyia the seed is always surrounded by a hard pericarp w'hich may be free
from the axis (as in S. disarticulata), embedded in it (as in 5. tenuis), or occupy an inter-
mediate state.

Seed: In the Salicornieae the seed provides diagnostic characters for the discrimination
of both genera and species. At the species level it is of particular importance since the
seed is one of the few organs not materially altered on drying. The seed coat is composed
of two layers, an inner membranous layer (the tegumen) which is closely applied to the
embryo and perisperm, and an outer layer (the testa proper); this outer layer is mem-
branous in Sarcocornia, Pachycornia, Sclerostcgia. and some species of Halosarcia, while
in other species of Halosarcia it is hard and brittle. Generally speaking, the hardness
of the testa in Halosarcia is inversely proportionate to the hardness of the pericarp or
fruiting perianth. The Australian species in the genus Arthrocnemum (i.e. Halosarcia)
were segregated into two sections by J. M. Black (1919) on characters based entirely
on the hardness of the pericarp and the testa; while this produced a neat distinction in
the material available to him, now' that far more species are known, some of which exhibit
a variety of intermediate characters, the use of these criteria for an infrageneric classification
is no longer tenable.

The embryo is horse-shoe shaped in Pachycornia and Sarcocornia but is straight or
curved in the other genera; other characters, however, suggest that relationships of Pachy-
cornia are more with Sclerostegia and Halosarcia than w'ith Sarcocornia.

The albumen in the seed of Chenopodiaceae is largely stored as perisperm (Mohabale
& Solanky, 1954; Hindmarsh. 1966; Corner, 1976). This is present in the mature seed
of all genera of Australian Salicornieae except Sarcocornia. The endosperm is largely
absorbed by the developing embryo and normally only a few layers remain around the
radicle.

12

Nuytsia Vol. 3 (1) 1980

Sex: The flowers in most genera of the SaJicornicae are basically hermaphrodite but they
may be functionally female through the failure of the stamens to fully develop: the latter
state is especially noticeable in flowers borne towards the apex of an inflorescence.

In Tegicornia the plants are strictly dioecious while in both Pac/iycornia and Sclerostegia
the lateral flowers of a triad are male and the central hermaphrodite. In Hafosarcia
inclica only plants bearing female flowers have been observed in India and Africa*; in
Australia at some localities only female plants have been found whereas in others the
species is hermaphrodite. In spite of the apparent absence of either male or hermaphrodite
flowers, plants of H. imiica which were observed by the author at Townsville. Queensland
set seed in abundance. In several other species of Halosarcia at some localities only
female plants have been noted and in these the style and stigmas are often folded back
on top of the ovary and remain within the perianth; seed is formed, and apparently without
fertilization being possible. (See also comments under H. pergramilata ssp. queenslamlica
and H. leptocicula.)

Both apomixis and dioecy are probably far more widespread in the Australian Sa/i-
cornieae than is reported here. Herbarium material, however, does not provide a good
guide to the prevalance of these phenomena since abortion of stamens is of frequent
occurrence and appears to be often related to environmental factors.

Taxonomy

When Linnaeus wrote the first edition of his Species Plantarum (1753) he recognised
only one genus, SaUcornia, in the group here referred to the Salicornioideae. Robert
Brown (1810) was the first person to describe a member of this subfamily from Australian
material (SaUcornia arhuscida)\ at the same time he recorded S. imiica Willd. as being
present in Australia although the specimens he referred to this name in fact belonged to
other species.

In the first half of the 19th century several genera were described as segregates, or as
close relatives of the genus SaUcornia, and into three of these were placed species based
on Australian types, or species that had been recorded from Australia. The genera with
Australian representatives were Halocnenmm Bieb. (1819), Sarcafliria Raf. (1837), and
Arthrocnemum Moq. (1840). A non-specialist, working with very imperfect material,
has difficulty in assessing the merits of the dift'erent genera, and it is not surprising that
Bentham (1870), when writing up the Salicornieac for the Flora Australiensis, chose to
recognise only one genus, SaUcornia; he noted that the material available to him was
unsatisfactory and that living plants should be examined in order to obtain a proper
appreciation of taxonomic affinities. Subsequently J. D. Hooker (1880) described two
further genera, Tecticornia and Pachycornia, in both cases basing them on a single Aus-
tralian species; he also recognised the presence in Australia of the genus Arthrocnemum,
in which decision he was largely following the revisions of the Salicornicae by Ungern-
Slernberg (1866, 1876). The circumscription adopted by J. D. Hooker for Australian
taxa was accepted by Volkens (1892) and in a modified form by Ulbrich (1934). The
only person after Bentham to revise the Australian Salicornicae as a whole was J. M.
Black (1919) who distinguished the same genera as those recognised by Hooker. Black
noted the importance of the seed characters when circumscribing species but unfortunately
many type specimens were not available to him and the material he studied was seldom
in a condition suitable for critical analysis.

* Tolken (1967) records having observed a stamen in some flowers in material from Lourenco Marques.

Paul G. Wilson, Australian Salicornieae

13

The genera within the subfamily Salicornioideae were reviewed by A. J. Scott (1978)
who described a new genus Sarcoconiia into which he placed the perennial species of
Salicornia (including the two Australian members). He also re-established the lecto-
typification of the genus Arthrocnemum by the species now correctly known as A. glaucum
(Delile) Ung.-Sternb. ;* this action will be further discussed since it affects the names of
many Australian taxa.

During the present investigation of the Australian Saliconueae it became apparent
that the accepted generic concepts, even after the paper by Scott, inadequately catered
for the variation found within that tribe. It was further apparent that, except for the
species here placed in Sarcocornia, the Australian species (irrespective of genus) showed
greater affinity to each other than they did to any non-indigenous species. The Salicornieae
of Australia have therefore, except for Sarcocornia, probably evolved from a common
ancestor within that country.

The Australian Salicornieae can be divided into two very distinct groups. In the
first group is found Sarcocornia which is distinctive in possessing the following characters:

(1) Sclereids present in chlorenchyma tissue.

(2) Flowers with an adaxial perianth lobe.

(3) Flowers with both an adaxial and an abaxial stamen.

(4) Absence of albumen (perisperm) in the seed.

The remaining members are fairly uniform in flower morphology and it is only by
a combination of characters that they may be discriminated. They have in common the
following characters:

(1) Absence of sclereids in chlorenchyma tissue.

(2) Flowers without an adaxial perianth lobe.

(3) Flowers with a single (abaxial) stamen.

(4) Abundant albumen (perisperm) in the seed.

Within this second group a number of endemic (or nearly endemic) genera have
been segregated out leaving the bulk of the species in the genus usually known as Arthroc-
nemum. As recognised by Scott (1978) this genus, in addition to the Australian species,
contains one species in California and another in the Mediterranean-lndian area. The
two extra-Australian species {A. siihlerminale (Parish) Standley and A. glaucum (Delile)
Ung.-Sternb.) are distinctive in having plain sclereids in the chlorenchyma tissue and in
having two stamens to each flower, one adaxial and the other abaxial in position; they
are therefore intermediate in character between the genus Sarcocornia and the Australian
members of "Arthrocnemum" . The two groups of species 1 consider to be sufficiently
distinct to warrant separation as separate genera. This then rai.ses the question as to
which group should bear the name Arthrocnemum.

The lectotypification of the name Arthrocnemum was discussed by Scott (1978).
The genus as originally defined by Moquin-Tandon (1840) included species here considered
to belong to five genera; e.g.:

(1) Nulostachys (i.e. A. hekingerianum Moq.).

(2) Sclerostegia (i.e. A. arhuscula (R.Br.) Moq.).

(3) Sarcocornia (i.e. A. frulicosiim (L.) Moq. and A. fruticosum var. radicans
(Sm.) Moq.

(4) ‘Extra-Australian’ Arthrocnemum (i.e. A. fruticoaum var. macrostachyum
(Moric.) Moq. and var. glaucum (Delile) Moq.)

(5) ‘Australian’ Arthrocnemum (i.e. A. indicum (Willd.) Moq.)

See note added in proof p. 152.

14

Nuytsia Vol. 3 (I) 1980

Moquin later (1849) modified his original circumscription of the genus and it was
further defined by Ungern-Sternberg (1866, 1876) who retained of the original species
only A. imiiciim, A. arhiiscula. and A. macrostachyum (Moric.) C. Koch (which had been
treated by Moquin as a variety of A. frulicosum). The genus was first lectotypified by
Standley (1916) on “A. fruticosum Moq.” by which he evidently meant A. frulicosum
(non (L.) Moq.) sensu Moq, (i.e. A. macrostachyum) since he included Salicornia frulicosa
(L.) L. (which is based on the same name as A. fruticosum) in Salicornia (see also Standley,
1914). This lectotypification was re-iterated by Ulbrich (1934) and accords generally
with Moquin’s concept of the genus. Unfortunately, in a posthumous paper by Moss
(1954), the type species is stated to ht A. fruticosum (L.) Moq., which statement was repeated
by Tolken (1967), and by the Inde.x Nominum Genericorum where, however, the typification
is incorrectly referred back to Ulbrich. ft is apparent that the lectotype has to be selected
from one of the three species retained in the genus by Ungern-Sternberg of those originally
included by Moquin; the choice is therefore between A. itulicum, A. arhuscuia, and A.
macrostachyum ( A. glaucum (Del.) Ung.-Sternb.).

The lectotypification of the genus Arthrocnemum is critical for the nomenclature of
Australian Salicornieae, since if, as stated by Moss (1954) and Tolken (1967), the type is
A. frulicosum (L.) Moq. then those species now included in Sarcocornia would have to
be transferred to Arthrocnemum and the species at present placed in the latter genus would
require a new generic name. If A. itulicum is chosen then the Australian species currently
referred to Arthrocnemum would remain in that genus but the two non-indigenous species
would require a new generic name. If A. arhuscuia is selected then the name Arthrocnemum
would become synonymous with (and would replace) the name Sclerostegia. The remain-
ing alternative is that adopted, in effect, by Standley (1916) and Ulbrich (1934), and which
has been recently re-iterated by Scott (1978): to choose A. glaucum as the lectotype.

Of the four alternatives only three can be seriously considered since the species A.
fruticosum had already been excluded from the genus by Ungern-Sternberg (1866 and
1876) and his circumscription was followed by Hooker (1880), Volkens (1892), Standley
(1916) and Ulbrich (1934). The two names A. iiulicum and A. arhuscuia have never been
proposed as types. It therefore remains only to accept A. glaucum as the type. Arthroc-
nemum glaucum. with A. suhterminale (Parish) Standley, is here considered to be generically
distinct from the Australian species currently included in that genus; the Australian taxa
are therefore here placed in a new genus Halosarcia.

Systematic Position

It is not intended in this paper to critically discuss the rank to which the group col-
loquially called samphires should be accorded, e.xcept to remark that it is considered by
all recent workers to be a natural group clearly separable from, although obviously closely
related to, the rest of the Chenopodiaceae. The group of genera that developed around
the name Salicornia was first recognised as belonging to a distinct tribe, the ‘"Salicornieae",
by Dumortier (1827); subsequently most taxonomists have accepted the circumscription
and rank of this taxon. However, J. D. Agardh (1858) recognised it as a family (“Sali-
cornieae") and Ulbrich (1934) as a subfitmily (Salicornioideae) of the Chenopodiaceae.
Ulbrich divided the Salicornioideae into two tribes, the Haplopepikleae and the Salicornieae,
the latter being subdivided into the subtribes Halosiaehyinae and Salicorniinae-, all the
Australian genera were referred to the tribe Salicornieae with Teeticornia in the Halo-
staehyinae and the rest in the Salicorniinae. The distinction made by Ulbrich in recognising
these two subtribes was basically as to whether the opposite bracts of the inflorescence
articles were united or free. That this is not a natural character for use in subdividing
the Australian representatives is apparent from the lack of correlation between this and
other characters; in fact in this present treatment the genera Halosarcia and Sclerostegia
both contain species with free bracts and species with united bracts. In some species

Paul G. Wilson, Australian Salicornieae

15

spikes on the same plant may have both free and united bracts, while one species (Halos-
arcia pterygosperma) exhibits a transition in its morphology from spikes with clearly united
bracts (in southern Western Australia) to those in which they are quite free (in the vicinity
of Carnarvon).

Recently Scott (1978) reinstated the family Salicorniaceae. He supported this action
largely on the unique morphology of the stem and leaves since the floral characters by
themselves do not permit a clear separation from the rest of the Chenopodiaceae. This
does not appear to be sufficiently substantial to warrant recognition of the samphires as
representing a distinct family and 1 have retained the broad concept adopted by Ulbrich
(1934) and most other workers. Both Scott and Ulbrich recognised the tribe Salicornieae,
into which they place all the samphire genera found in Australia.

No attempt is here made to erect a new generic classification within the Australian
Salicornieae. The genus Tecticornia, which seemed at first to be so distinct (Wilson,
1972) is clearly linked to the genus Halosarcia through H. fiaheiliformis. The genus
Sclerostegia approaches on the one hand Pachycornia in its floral morphology, and on
the other (through S. arinisciila) members of the genus Halosarcia. The genus Sarcocornia
is clearly distinct from the other Australian genera although closely related to Salicornia.

The name ‘Samphire’

The members of the Salicornieae are, in Australia, generally referred to as ‘samphires’,
although in some recent floras the name ‘glasswort’ is preferred. The word samphire is
derived from the earlier English name ‘sampere’ which itself comes from the French name
‘herbe de St. Pierre’; this name was initially applied to Crithnnun maritimum, a coastal
member of the Apiaceae, which was (and still is) eaten either fresh or as a pickle. The
English coastal plant Salicornia eiiropaca is also used in this way and presumably for
this reason was sometimes also called samphire; the name subsequently became applied
to all the succulent articulate chenopods.

The common name ‘glasswort’ is another early English name for species of Salicornia
and arose from the use of the ashes of these plants in the manufacture of glass.

Citation of Specimens

Only a selection of herbarium specimens of each species is cited except for those
taxa which have been seldom collected. An attempt has been made to map all records
which indicate a significant extension of the range of a species.

Subfamily Salicornioideae, Tribe Salicornieae

Shrubs or herbs with succulent apparently articulate branches. Leaves opposite,
connate (alternate and amplexicaule in one extra-Australian genus), forming an insig-
nificant, fieshy, bilobed rim at the apex of each article (internode). Flowers hermaphrodite
or unisexual, sessile, arranged in a spike-like thyrse usually made up of axillary 3-fiowered
cymules; secondary bracts or bracteoles absent. Perianth gamotepalous, (2) 3-4 lobed,
frequently succulent. Stamens 1 or 2, free, placed in anterior-posterior position; anthers
4-locular (tetrasporangiate). Ovary sessile; stigmas 2 (3). Fruiting perianth and pericarp
membranous to spongy or crustaceous. Seed vertical; testa membranous to crustaceous,
smooth or variously ornamented; perisperm abundant or absent; embryo straight, curved,
annular, or hippocrepiform and incumbent.

Type genus; Salicornia L.

16

Nuytsia Vol. 3 (1) 1980

Key to Genera

I. Perianth with truncate or rounded apex (outer face); stamens two; seeds orbicular.

embryo horseshoe-shaped, perisperm absent 5. Sarcocornia

1 *. Perianth various; stamens solitary, seeds with perisperm 2

2. Flowers solitary in axil of foliage leaves 3. Tegicornia

2*. Flowers in opposite 3-flowered cymules (triads), usually in spike-like inflorescences 3

3. Lateral flowers of triad male, central hermaphrodite 4

3*. Flowers of triad hermaphrodite or female 5

4. Leaf lobes prominent, fleshy, divaricate; embryo annular 1. Pachycornia

4*. Leaf lobes insignificant, scarious; embryo ± straight 2. Sclerostegia

5. Opposite bracts united 4. Halosarcia

5*. Opposite bracts free from each other 6

6. Plants woody perennials or shrubs; perianth gamotepalous 4. Halosarcia

6*. Plants annuals or short-lived perennials; perianth of two succulent (eventually

free) laterally placed tepals Tecticorniaf

I. Pachycornia Flook.f.

Hook.f. in Benth. et Hook.f., Gen. PI. 3:65 (1880); Volkens in Engler et PrantI, Nat. Pfianzenfam. III.
la;77 (1892); .1. M. Black, Trans. Roy. Soc. S. Austral. 43:363 (1919); Ulbrich in Engler et PrantI, op.cit.
ed. 2. 16c:549 (1934); A. J. Scott, Bot. J. Linn. Soc. 75:369 (1978).

Dwarf shrubs with succulent stems. Leaves opposite, of short fleshy spurs apparently
decurrent along internode. Inflorescence a spike-like thyrse consisting of triads of flowers
borne in the axils of opposite decurrent bracts. Flowers dimorphic, the lateral male and
the central hermaphrodite. Perianth gamotepalous, membranous, unchanged in fruit.
Stamens solitary (abaxial in central flower). Ovary with a solitary ovule attached to base;
stigmas two, narrowly triangular, papillose. Fruit: pericarp hard, fused to and sunken
into the woody spike axis. Seed orbicular; testa thin; embryo annular; perisperm central.

Type species: P. triandra (F. Muell.) Black
A single species endemic to Australia.

Origin of name: from a Greek word pachy ^ thick and a Latin word cornu == a horn, but
also evidently cognate with the name Salicornia which is a latinization of the French word
‘salicoF.

1. Pachycornia triandra (F. Muell.) J. M. Black, FI. S. Austral. 206 (1924). Fig. 11.

Arthrocnemum triandrum F. Muell., Fragm. 1:139 (1859 ). — Salicornia robusta F. Muell., Fragm. 6:251
(1868) nom. superfl., based on preceding; Benth., FI. Austral. 5:202 (1870).— P. robusta (F. Muell.) Bail.,
Hist. PI. 9:184 (1888) comb, illeg.; A. J. Ewart, PI. Indig. Victoria 2: tab. 83 (1910); J. M. Black, Trans.
Roy. Soc. S. Austral. 43:363 (1919). — S. triandra (F. Muell.) Druce, Rep. Bot. Exch. Cl. Brit. Isles 1916:644
(1917).

Lectotype: “Near above the junction of the Darling on the Murray and on the lower Darling
as far down the Murray as Lake Victoria”, Dec. 1853, F. Mueller (holo; MEL 71400)
lecto. nov.

Divaricately branched bright green subshrub ca. 50 cm high. Internodes cylindrical
1-2 cm long with thick rigid acute lobes ca. 3 mnr long that are ciliolate towards their
base; at union of lobes arise opposite ciliolate grooves which run for length of internode.
Spike broadly ovoid, 1-2 cm long, 0-8-1 -2 cm wide, bracts congested, the lobes acicular
and strongly divaricate forming a decurrent groove at their union. Flowers steeply ascend-
ing, ca. 4 mm long, free (except near their base) from each other and from upper and
lower bracts. Perianth membranous, dorsiventrally compressed, irregularly fimbriate

+ See P. G. Wilson (1972).

Paul G. Wilson, Australian Salicornieae

17

at the terminal aperture; central floret with a pair of abaxial imbricate lobes (lateral florets
with indefinite lobing). Anther oblong, ca. 3 mm long, filament stout. Ovary vertical,
succulent, fused to spike axis; style firm, narrowly oblong (dorsivenlrally flattened),
not exserted; stigmas narrowly triangular. Fruiting spike scarcely expanded; axis and
base of bracts thick and woody; bracts pithy but firm: perianth membranous; pericarp
thick, horny, fused to and apparently sunken into the woody axis. Seed orbicular, ca.
2 mm diameter; testa membranous, smooth, brown; embryo annular; perisperm central.
Seed released by decay of spike.

Habitat: Usually in heavy somewhat saline soil; it has been recorded from gravelly or
sandy situations but in these, presumably, the soil is underlain by clay.

Distribution: Eremaean areas of inland Australia. Map 1.

New South Wales: Wentworth, Sept. 1912. H. B. Wittiamsoii (MEL); Junction of Darling and Murray
Rivers, May 1887, W.J. /to/r//w (MEL).

Victoria: Merbein, May 1937, J. H. Willis (MEL); 1 mi E of Berribee Tank, 31 Aug. 1948, J. H. Willis
(MEL).

South Australia: Charlotte Waters, 1889, W. Schwartz (MEL); Mildura, 1894, C. French (MEL); Lake
Gardner, J. Z. Weber 3330 (PERTH).

Western Australia: Balgo Downs. H. Demarz 5230 (PERTH); Mongers Lake, M. Koch 1374 (PERTH);
Zanthus, R. D. Royce 5359 (PERTH); 4 mi E of Kalgoorlie, 5 Aug. 1971, ./. Verschuer (PERTH).

Northern Territory: 2 mi W of Maryvale HS., 4 July 1955, G. Chippendale (NT); 2 mi N of Rodinga, P. K.
Latz 1853 (PERTH); 6-5 mi S of Alice Springs, D . ./. Nelson 505 (PERTH).

Pachycornia triandra is remarkable in having very prominently lobed articles which
possess opposite grooves decurrent from the junction of the lobes and beneath which the
palisade tissue is lacking; the general appearance being similar to that of a Salsola species.
The circular seed with its horseshoe-shaped embryo and central perisperm is also distinctive.
Secondary thickening of the spike axis due to cambial activity does not take place although
there is a slight increase in thickness with the maturation of the seed. However, the base
of the bracts, the pericarp, and the broad axis of the spike, become lignified, which is the
cause of the eventual solidity of the whole structure. This condition, then, is fundamentally
different from that in Sclerostegia tenuis where the woody spike axis undergoes secondary
growth.

Since the infructescence does not continue growth it dies after the maturation of the
seed. However, the structure remains on the plant for some time, possibly several years,
until the decay of the axis causes it to break off. In some cases this would appear not to
take place until the death of the whole plant.

The genus Pachycornia was enlarged by Black (1919) so as to include Sclerostegia
tenuis, and Scott (1978) logically added the closely related species S. arhuscula. Each
of these species (and the newly described species in Sclerostegia) has a similar type of
inflorescence in which the lateral flowers are male and the central hermaphrodite; they
also have similar perianths and stigmata. The unique characters which distinguish
P. triandra, and which 1 consider to be of generic significance, are the vegetative morphology
(large leaf-lobes and decurrent grooves), the annular embryo, and the lignification of the
inflorescence axis and bract bases.

2. Sclerostegia P. G. Wilson, gen. nov.

Fruticulus quasi aphyllus, articulatus, glaber. Folia succulenta opposita cauli adnata, apice modice
dilatata sub-2-loba. Inflorescentia; spicoidea congestae, floribus lateralibus masculis, centralibus herm-
aphroditis. Flores: perianthium gamotepalum breviler 2-3-lobum; stamen solitarium, abaxiale; ovarium
ovoideum, apice in stylo robusto attenuatum. Fructus: perianthium persistens utriculo includens; peri-
carpium durum semine omnino includens. Semen erectum, ovoideum; testa tenuiter Crustacea, i; laevia;
embryo rectus vel leviter curvatus; perispermium copiosum, laterale.

91670 — ( 2 )

18

Nuytsia Vol. 3 (I) 1980

Type species: Sclcroslci;ia tennis (Benth.) P. G. Wilson {Scilieornia tenuis Benth.)

Dwarf shrubs, seemingly leafless, glabrous; branches when young of articulated
cylindrical to spherical segments (internodes) which are cup-shaped to two-lobed at apex;
segments succulent, the fleshy outer portion ultimately shrivelling and falling away from
the woody axis. Infioreseence either a spike-like thyrse consisting of triads of flowers
(cymes) immersed in the axils of each fleshy bract, or of triads dispersed among the branch
segments; bracts free from each other or united in opposite pairs. Flowers dimorphic,
horizontal to almost vertical in position, the central one of the triad hermaphrodite and
the lateral ones male; perianth gamotepalous, membranous or fleshy, with a pair of laterally
placed imbricate lobes on the anterior (abaxial) side of the orifice and sometimes a small
abaxial inner lobe; stamen solitary (abaxial in hermaphrodite flower); style short and
stout, passing into the ovary; stigmas 2, narrowly triangular, papillose. Fruit: perianth
remaining membranous or. when originally succulent, becoming shrivelled; pericarp
firmly crustaceous or woody. Seed ovoid; testa thin, smooth or faintly granular over the
embryo; embryo straight or slightly curved with the radicle inferior; cotyledons con-
duplicate; perisperm copious, lateral.

Five species, all endemic to Australia (including Tasmania).

Origin of the generie name: from the Greek words seleros (hard) and stegos (shelter);
referring to the hard pericarp.

The genus Selerostegia may be distinguished from the other members of the Sali-
cornieae by a combination of two characters, (I) the dimorphic flowers, the central one
of the triad being hermaphrodite and the lateral ones male, and (2) the straight or only
slightly curved embryo with copious lateral perisperm. The first character by itself clearly
differentiates it from all other members of the tribe except for Paehyeornia: this genus is
distinguished by its annular embryo and by its unique vegetative characters.

Selerostegia tenuis. S. rneduUo.sa. and S. inoniliformis possess membranous perianths
which persist in this condition even when in fruit. In these three closely related species the
flowers are nearly vertical in position, with the apertures (which are dorsiventrally flattened)
at their summit. The aestivation of the perianth lobes is not clear; in some flower buds
studied there was a pair of laterally placed conduplicate lobes (with each of the lobes
over-lapping on one of its margins and under-lapping on its other); in other flowers only
the imbrication on the abaxial side of the perianth could be distinguished, the adaxial
lobing being either absent or not apparent. In all cases only the aestivation of the central
hermaphrodite florets was noted since that of the lateral florets is difficult to observe.
In S. arhuseula the perianth is succulent and fused to the upper bract but the flowers
are otherwise similar to those of the preceding three species. In S. disarticulata the flowers
are almost horizontal, the perianth succulent, and the aperture on the outer truncate
face. In this case there is a pair of large lateral imbricate lobes and a smaller inner abaxial
lobe which links the lateral pair together.

In all species of Selerostegia the pericarp becomes hard and completely encloses the
seed which has only a thin testa. In S. nwniliforniis, S. niedidlosa, and occasionally in
S. arhuseula. the 'spike' (or fertile branch) continues growth vegetatively and the shrivelled
bracts eventually become sloughed off along with the old flowers and fruit. In S. tenuis
the spike-axis always undergoes thickening and lignification. The stele of this axis consists
(in the mid internodal position) of two sets of vascular strands, one set on each side of
the transversely placed axis (see Fig. 14F). Lignification and eventual secondary thickening
of the stele results in the formation of two opposite vertical bands between "the pairs of
triads (the bands being therefore opposite and decussate, changing their position at each
node) causing the fruit to become more and more deeply imbed'ded in the spike axis. As
the stele undergoes this thickening the woody pericarp also grows in thickness although
not so markedly.

Paul G. Wilson, Australian Sallcornieae

19

Key to the species of Sclerostegia

1. Spikes with the opposite bracts free from each other I. S. disarticulata

1*. Spikes with the opposite bracts clearly united, or the flowers not in spikes but in

the axils of the leaves along the branches ,, 2

2. Perianth fleshy; style hard and protruding in fruit (especially when dry)

2. S. arbuscula

2*. Perianth membranous; style not protruding . .. . , 3

3. Fertile and vegetative articles alike, cylindrical; leaf lobes markedly denticulate;

pericarp not embedded in a woody axis 5. S. medullosa

3*. Fertile and vegetative articles either dissimilar or the former not cylindrical 4

4. Fertile articles obovoid, becoming dry, globular to obovoid and pithy in fruit;

pericarp crustaceous, not embedded in a woody axis 3. S. moniliformis

4*. Fertile articles obovoid to cylindrical, becoming thick and woody in fruit; pericarp
woody and embedded in the woody axis 4. S. tenuis

1. Sclerostegia disarticulata P. G. Wilson, sp. nov. Fig. 12, 34.

Anicidi doleilbrnies vcl obovoklei 3-6 miii long!, apice brevitcr ct obtuse lobato, margine integro
(raro denticulalo). In/loirsceiilicie lenninales, vel lalerales cl pcdunculalae. anguste cylindriceae, 5-10
(20) mm longac, 3 4 mm diam.; bracteae discretae, lade exteriore semidrculari. Flores horizontales,
connali, apice truncali; llores lalerales parvi, centrali magni, Pcrunnhiiim succulentum, orifido in facie
exteriore. Frucliis late obovoideum, ca. 1-7 mm longus. in perianthio succulento (demum areolato)
obtectus, ad tlores lalerales adnalus; pericarpium obovalum, oninino corneum. Infnictcsceiitia demum
in parte discreta seeedens.

Type- N end of Mongers Lake, 28 46'S, 117 22'E, Western Australia, 6 Aug. 1969,
P. G. Wilson 8603 (holo: PERTH, iso: CANB, K).

Divaricately branched rounded suhshnih 0'3-l-5 mm high. Branches slender;
articles fleshy, doleiform to obovoid 3-6 mm long, dull to bright green (often black when
dry), apex very shortly and bluntly lobed, margin entire (or rarely denticulate). Spikes
terminal, or lateral and then shortly pedunculate, narrowly cylindrical, 5-10 (20) mm
long. 3-4 mm wide (2-5-3 mm in dried state); fertile articles 5-10 (20); opposite bracts
free (or almost free) from each other, outer surface hemispherical, upper margin horizontal
or slightly convex. Flowers in triads at right angles to spike axis, intimately fused to each
other, at first adherent in proximal half to the upper bract; apex (outer exposed surface)
of triad truncate and triangular, only partially obscured by subtending bract; lateral
(male) flowers small, central (hermaphrodite) flower large. Perianth at iirst membranous
at apex but eventually entirely lleshy with the aperture along the outer surface; lobes
(of central floret) 3, a pair of long lateral and imbricate lobes and a small inner anterior
(abaxial) lobe. Anther ca. 1 mm long. Ovary thick and fleshy, adherent (at first) to
perianth and fused to spike axis along vertical length of flower; style not exserted; stigmas
very narrowly triangular. Fruit: pericarp broadly obovoid, vertical, ca. 1-7 mm long,
horny all over, surrounded by the thin fleshy (eventually areolate) perianth and fused
to the lateral flowers. Seed vertical (parallel to a.xis of spike), ovoid, ca. 1 -5 mm long;
testa membranous, smooth, pale brown; embryo slightly curved. The spike, when mature,
breaking up into separate bracts and triads, each of the latter consisting of an obovoid
horny pericarp surrounded by the dry perianth and fused laterally to the small shrivelled
male flowers; pericarp eventually dividing longitudinally to release seed.

Habitat: Many of the herbarium specimens (especially those from New South Wales)
record the habitat as stony, or as a gravelly hillside, but the species is widespread along the
southern margin of the Nullarbor Plain where it grows over limestone, while collections
made in Western Australia from the more northern limits of its distribution appear to
have come from clay pans. One specimen is stated to have been collected from the coast
near Adelaide (C. M. Eardley, 19 April 1936, ADW 3208), but the environment here is
so different from that of the other collections that the data are open to doubt.

20

Nuytsia Vol. 3 (I) 1980

Distrihiition: Central and southern Australia, in Eremaean and suberemaean areas,
south of 26 lat. and east of 144° longitude. Map 2.

New South iVales: 18 mi W of White CliflFs, E. F. Constable 4600 (NSW); 16 km NW of Milparinka, G.
M. Ciiiwingham &nd P. L. Milthorpe 1097 (NSW); Tero Ck. Stn„ 30°3'S, 142°50'E, 2 Dec. 1968, P. Martemz
(AD, CANB. NSW).

South Australia: 3 km S of Leigh Ck.. Hj. Eich/er 12982 (AD); Evelyn Downs, 8 Ocl. 1953, E. H. Ising
(AD); 80 mi W of Penang, 6 Sept. 1962, M. E. Phillips (AD); 8 mi S of White Wells, 28 Aug. 1947, J. H.
Willis (MEL).

Western Australia: 5 mi E of Kalannie, K. M. Allan 48 (PERTH); 9 mi N of Carnarvon, T. E. H. Aplin
1565 (PERTH); 2 mi E of Norseman, J. S. Beard 5204 (PERTH); 3 mi N of Coolgardie, E. M. Bennett
3257 (PERTH); 6 mi W of Hamelin Stn., J. W. Green 1409 (PERTH); .limba Jimba, Gascoyne R., C. A.
Gardner 6141 (PERTH); Windimurra Stn., 19 Aug., 1976, V. Semeniuk (PERTH); Jimba Jimba Stn.,
D. O. Wilcox 158 (PERTH).

Northern Territory: Henbury Stn., T. S. Henshall 789 (NT); Finke R., 1889, W. F. Schwarz (MEL); Glen
Helen-Alice Springs Rd., 1 mi W of Ormiston Gorge turn-off, A. C. Beauglehole 24247 (PERTH).

Sclerostegia disarticulata exhibits little regional variation apart from that which
might be expected from the same species growing in different environments. However,
at Jimba Jimba, in the Gascoyne River area of Western Australia, is found a variant
with more deeply lobed articles which, with the bracts, are prominently denticulate on
their margins (Wilcox 158 and Gardner 6141).

The species may be recognised by its slender spikes with free opposite bracts, and
by the manner in which the spike breaks up when mature into single bracts and compact
triads each of which contains a single seed. The specific epithet has reference to the
general disintegration of the mature infructescence.

2. Sclerostegia arbuscula (R.Br.) P. G. Wilson, comb. nov. Fig. 13A-D, 35.

Salicornia arbuscula R.Br., Prod. 411 (1810). — Arthrocnemum arbuscula (R.Br.) Moq., Chenop. Enum.
113 (1840).— Halocnemum arbuscula (R.Br.) F. M. Bailey, Synopsis Qld. Flora 409 (\881)).—Pachycornia
arbuscula (R.Br.) A. J. Scott, Bot. J. Linn. Soc. 75:369 (1978).

Lectotype: Port Dalrymple, Jan. 1804, R. Brown (3074) (holo; BM) lecto. nov.

Much branched shrub to 2 m high with ascending branches. Articles slightly glaucous
or dull green, obovoid, ca. 5 mm long, shortly lobed, entire. Spikes terminal to both
main and lateral branchlets (sometimes sessile), of 2-6 articles, slightly torulose (articles
laterally compressed), succulent; bracts shortly lobed, united in opposite pairs and almost
enclosing the flowers. Flowers ascending, intimately fused to each other and to the upper
bract, free from the lower; lateral flowers (these occasionally aborted) male and smaller
than the central hermaphrodite flower. Perianth fleshy at lateral margins and apex,
membranous abaxially with the aperture at or just below the apex on the outer face;
lobes 2, lateral, rounded and overlapping but becoming obscure with emergence of style
and anther, sometimes also with a small abaxial lobe. Ovary fused along v^ertical length
of spike a.xis, passing distally into a short stout style; stigmas'narrowly triangular. Fruit:
perianth shrivelled, weak; pericarp thick and hard, deltoid in lateral view and passing into
the thick hard persistent style which is exserted from the perianth. Seed vertical, elliptic,
ca. 1-5 mm long; testa pale brown, thin, smooth except for faint granulation over radicle-
embryo slightly curved, perisperm lateral.

Habilal: Sclerostegia arbuscula (in its typical form) is coastal or near coastal in its dis-
tribution and frequently occupies marshes subject to occasional inundation by the sea.

Distribution: Principally in coastal areas of southern New South Wales, Victoria, Tasmania,
South Australia, and south east Western Australia. Map 3.

New South Wales: Currarong, 20 June 1970, L. ,4. S. John.son (NSW).

Victoria: Griffith Is., A. C. Beauglehole 19487 (PERTH); Goat Is., A. C. Beauglehole J-’ISS (PFRTHt-
Hastings, Jan. 1937, A. Meebold (NSW); Port Phillip, 1848, F. Mueller (MEL). *” ’

Ralph Bay, F. H. Lang 138 (HO); Double Creek,
28 Feb. 1976, J. W. Parham (PERTH); S arm of R. Derwent, F. A. Rodway 7154 (NSW).

Paul G. Wilson. Australian Salicornieae

21

Soiilh Australia: Point Riley, B. Ctipley 1300 (AD); I -5 km N of Prospect Hill, Kangaroo Is., Hj. Eich/er
15479 (AD); Outer Harbour, Pt. Adelaide, M. A. Fagg 120 (AD); Coobowie Inlet, Yorke Pen., D. N.
Kraehenhuehl 1978 (AD); Fowlers Bay, 29 Sept. 1959, D, E. Symon (ADW).

Western Australia: 2 km NW of Pt. Charles, Fitzgerald R., K. Newbev 402& (PERTH); Israelite Bay, P. G.
fPf/iw; 10079 (PERTH).

Sclerostegia arhuscuUi may be readily distinguished from other species of the genus,
and from those of related genera, by the presence of a hard protruding style when in fruit;
this character is even more pronounced in the dried state.

In Sclerostegia arhiisciila the spikes typically terminate a branchlet although occasion-
ally one may resume growth to form a vegetative branch. Normally the bracts and
perianths become dry and the spike breaks up into .separate articles, each of which contains
a pair of opposite fruitlets. Further disintegration presumably occurs on the decay of
the articles. Where the spike has continued growth to form a branch the fruitlets may
remain attached to the axis for up to at least one year and then fall away with the decay
of the bract tissue.

Bentham (1870) confused Sclerostegia arhitseii/a with H. Iialoctiemoides and H. syn-
carpa (a species which at that time w'as undescribed); much of the material cited by him
(under Salicoriiia arhiiscii/a) belongs in fact to these latter species.

In the eastern States and in South Australia the fertile articles are somewhat laterally
compressed so as to produce an uneven appearance to the spike, whereas in Western
Australia the articles are circular in cross-section.

At Israelite Bay on the south coast of Western Australia S. arhuscula and S. monili-
formis grow together. Some plants intermediate in form between these two species have
been collected in that area and the fact that these plants are apparently sterile suggests
that they are of hybrid origin. Examples are as follows:

Western Australia: Israelite Bay, /?. Hnatiuk 761206, 761233 (PERTH); ibid., R. D. Royce 6302 (PERTH).

A plant with similar flowers to those of 5. arhuscula, but with a very different in-
florescence and habit, has been collected in South-western Western Australia (12 km W
of K.oninin, P. G. Wilson 10240. PERTH and Mt. Caroline. 1886. Mrs. G. R. Sewell,
MEL 71557). In this plant the spikes are sessile, ellipsoidal in form (circular in cross
section), and are divaricately spreading, while the pithy bracts retain their shape on drying.
No material with mature .seed has been collected and (he nature of this taxon is at the
moment obscure.

In South Australia, along the coast near Adelaide, S. arhuscula and several Halosarcia
species grow in the same locality and collections have been made of a plant apparently
intermediate in habit between S. arhuscula and members of the genus Halosarcia; this
plant is seemingly abnormal in its floral morphology. The 'intermediate' plant has a
fruiting spike which is pithy in te,xture and does not contract around the fruit on drying
as is usual for S. arhuscula. The sex of the flowers is as in Sclerostegia; the perianth has
a small but obvious inner abaxial lobe, membranous in texture, an organ lacking in 5.
arhuscula. Field observations are necessary to ascertain whether this plant is of hybrid
origin or whether it is a discrete taxon. Some collections are as follows (all AD):

South Australia: Ethelton. 13 Jan. 1921,./. M. Black; Gleneig, 14 Dec. 1919 and 21 Jan. 1920,7. M. Black;
Port Gawler, J. P. Jessnp 2118.

From the south coast of Victoria come three collections which appear to be inter-
mediate between S. arhuscula and H. pergranulata. These specimens have hermaphrodite
flowers but have the style and stigmas of S. arhu.scula. In one collection (Morrison
s.n.) the florets are fused together and the perianth has simply a pair of lateral lobes set
in a fleshy apex; the seed is deformed and has faint porcate ribbing. In another collection
(Williamson s.n.) the florets are partially free, the imbrication of the three perianth lobes
is variable, and the immature seed again has faint porcate ribbing. Both 5. arhuscula
and H. pergranulata have been collected in the same general area as that in which the
apparently intermediate plants w'ere found. Records of this plant are as follows:

Victoria; Mouth of Yarra, Port Melbourne, 2 Apr. 1885, A. Morrison 336 (CANB); North Williamstown,
16 Feb. 1895, A. Morrison (BRI, CANB, NSW); Geelong, M. B. Williamson (AD 97526144 p.p.).

22

Nuytsia Vol. 3 (1) 1980

3. Sclerostegia moniliformis P. G. Wilson, sp. nov. Fig. I3E-H.

Ai ticiili obovoidei ca. 5 mm longi, lobis apiculati.s inlegris. liiflorescentiae terminales demum inter-
calares, 1-2 (6) articidis, bracteis inclusi. dimidio superiore dorsi-ventraliler applanati, orificio terminal!.
Perianthium membranaceum; stylus robustus in perianthio inclusus. Infnicte.sceiitici sicca interdum monili-
formis; verticilli bractearum obovoidei vel sphaerici. medullosi, 5-6 mm diam.; perianthium membran-
aceum; pericarpium crustaceum ad a.xem adnatum.

Type: W. bank of Lake King near causeway, Western Australia, margin of salt lake,
subshrub 0-3 m high, articles dull green and slightly glaucous, 29 Sept. 1970, P. G. Wilson
9983 (holo: PERTH, iso; AD, BRI, CANB, K, MEL, NSW).

Much-branched shrub ca. I m high with slender branchlets. Articles dull green,
fleshy, obovoid, ca. 5 mm long: apex lobed, apiciilate; margin scarious, entire. Spikes
terminal (becoming intercalary with age) of 1-2 (6) articles; bracts similar to the vegetative
articles and enclosing the flowers. Flowers almost erect, ca. 3 mm high, dorsiventrally
flattened distally, the aperture terminal, coalescent in lower two-thirds to each other and
to upper bract, free from lower bract except near base. Perianth membranous; central
floret with a pair of abaxial imbricate lobes reaching half-way to base of flower; lateral
flowers irregularly lobed. Anther ca. 2 mm long. Orary narrow, fused to spike axis
along vertical length of flower, continuing outwards as a stout style; stigmas narrowly
triangular. Fruiting spike dry, sometimes moniliform, the bracts broadly obovoid to
spherical, pithy, 5-6 mm diameter, completely obscuring the flowers; perianth i erect,
membranous; pericarp crustaceous, adhering to spike axis; style weak, not exserted.
Seed ovoid, ca. 1-5 mm long; testa thin, smooth, pale brown; embryo slightly curved;
perisperm lateral.

Habitat: Moderately saline areas in periodically waterlogged situations.

Distribution: South-western Western Australia from Bullfinch south to Lake Grace and
east to Israelite Bay. Map 4.

tVesrmi Australia: Lake King, A. S. George 9355 (PERTH); 7-2 mi E of Pt. Malcolm, R. Hnatiuk 761203
(PERTH); 23 mi N of Bullfinch. P. O'. Wilson 8736a (PERTH); Mt. Le Grand to Luckv Bav, P. G. Wilson
10014 (PERTH); tsraelite Bay, P. G. Wilson 10081 (PERTH); Pine Hill, 20 km N of'Mt. 'Ragged, P. G.
Wilson 10101 (PERTH).

Scleroslegia moniliformis resembles the western variant of S. tenuis which was described
by Ewart and White as a distinct species, Salicornia donaldsonii. In S. moniliformis
the habit is more branched and the articles are obovate rather than cylindrical; the scarious
margin of the lobes is also less prominent. The main distinction betw'een the two species,
however, lies in the fruit: in S. tenuis the pericarp becomes woody, fuses with, and becomes
immersed in, the enlarged woody spike axis, while in S. moniliforn.is the pericarp is thinly
crustaceous and does not fuse with the spike axis which at first remains slender. The
bracts of the latter species do not shrivel but become pithy, retain their spherical shape,
and then remain around the fruit while the spike tip continues to make vegetative growth.
The bracts disintegrate with age and are eventually sloughed off with the enclosed fruits.

Scleroslegia arbuscula is also similar to 5. moniliformis but may be distinguished by
its fleshy perianth which is totally fused to the upper bract. Intergradation between
these two species is discussed under S. arbuscula.

4. Sclerostegia tenuis (Benth.) P. G. Wilson, comb. nov. Fig. 14.

Salicornia tennis Benth., FI. Amstral. 5: 204 W10).~^Pachycornia tennis (Benth.) Black, Trans. Roy. Soc.

5. A. 43! 363 (1919).

Lectotype: “Howitt’s Expedition” (holo: MEL 71381; iso; K) cf. J. M. Black, op. cit.
364, see also comment below.

Salicornia donaldsonii Ewart et White, ,1. Roy. Soc. N.S.W. 42;
(Ewart et White) C.A. Gardn., Enuni. PI, Austr. Occ.'39 (1930), ’

194 (1909 ), — Arthrocnemam donaldsonii

Paul G. Wilson, Australian Salicornieae

23

Lectotype: Cowcowing, Sept. 1904, M. Koch 1147 (holo: MEL 71457; iso: K), lecto nov.

Divaricately branched sith-shruh to 0-6 m high. Braiichicis slender: articles pale
green narrowly turbinate to cylindrical, ca. 10 mm long; ape.x shortly lobed (the lobes
rounded to obtuse, sometimes apiculate) with a broad scarious margin, entire (rarely
slightly denticulate). Spikes slender, ca. 2 cm long, narrowed towards the apex, of 5-10
articles, usually continuing growth vegetativcly for at least 1-2 cm: bracts united in opposite
pairs, slightly lobed, ca. 4 mm long, enclosing the flowers, similar at first to the vegetative
articles. Flowers almost erect, ca. 4 mm high, dorsivcntrally flattened distally. the aperture
terminal, coalescent in proximal two-thirds to each other and, except near apex, fused to
the upper bract. Perianth membranous; central floret with a pair of abaxial imbricate
lobes: lateral florets irregularly lobed. Ovary narrow, fused to spike axis along vertical
length of flower, passing into a stout style: stigmas narrowly triangular. Fruiting spike
narrowly ellipsoidal (sometimes slightly moniliform), woody, covered by the shrivelled
remains of the bracts, or these soft, pithy and persistent; perianth membranous: pericarp
thick and woody and eventually embedded laterally in the enlarged woody spike axis;
style not e.xserted. Seed narrowly ovoid. 2-5-3 mm long; testa thin, smooth, pale brown;
embryo straight; perispenn lateral.

Habitat: Found in a variety of saline soils but usually with underlying clay. A collection
made in New South Wales ca. 38 km N of Hay (M. A. Wilson 34) was growing in a heavy
clay with 1-2% salt made up of almost equal quantities of common salt (NaCl) and
gypsum (Na^SOj).

Distribution: Eremaean areas of Australia. Map 4.

Queensland: Between the Bareoo and the Roma, 1871, Birch (MEL): Dynevor Lakes, M. E. Phdlips 475
(PERTH).

New South Wales: 15 mi SW of Yantabulla. D. H. Benson and ,/. Pickard 1825 (NSW); Corona, M. Collins
43 (NSW); One Tree, Margaret Wilson 34 (PERTH); Junction of Darling and Murray River. 1896, R. R.
Harvey (MEL): ibid.. 1889. Mrs. Holding (MEL).

Victoria: 3 mi NE of Lake Wallawalla. A. C. Beaugleholc 40677 (PERTH): W side ol Lake Walla-Walla,
31 Aug. 1948, J. H. Willis (MEL); Kulkyne Nat. Park. 16 Oct. 1960, ./. H. Willis (MEL); near Mildura,
20 Oct. 1937, W.J. Zinmwr(WWt).

South Australia: 5 mi SW of Pt. Augusta. C. R. Alcock 2148 (ADW, CANB); Yudnapinna Stn., ./. G.
Davies (ADW); 80 km S of Kingoonya, A/. Fagg 371 (AD); 2 km NW of Mt. Lyndhurst, M. Lazarides
8258 (PERTH); 10 mi NE of Woomera, A7. Lazarides 8405 (PERTH).

Western Australia: W end of Hopkins Lake, D. E. Svinon 2348 (ADW): Mongers Lake, Ruth Watson
5 (PERTH).

Northern Territory: Alice River, 1884, C. W. Birch (MEL); Finkc R., 11. G. Briggs 1258 (NSW); 12-4 mi
N of Andado H.S.. 6 Sept. 1956, G. Chippendale (NT).

In Sclerostegia tenuis the bracts of the fruiting spike normally shrivel to form a thin
covering to the woody axis. In some collections, however, the bracts have become firm
and spongy and assume a moniliform appearance similar to that found in S. ntoniUfonnis
but w'ith the seed embedded in the spike axis. The spikes sometimes become intercalary
due to the continued vegetative growth at their apex; in most cases thi.s vegetative growth
is short-lived producing only a few centimetres of branch. In the variant found at Lake
Cow'cowing, Western Australia, (which was described as a distinct species, Salicornia
dotiahlsoniik however, vegetative growth sometimes continues indefinitely with the result
that the spike becomes embedded in thick branches several years old. In those species
of Halosareia where the spike becomes intercalary and the branch continues growth for
several years, the fruit (seed with surrounding pericarp) is on the outside of the stele and
is eventually shed with the rest of the bract structure; wdiereas in S. tenuis the stele of the
young spikij consists of two .sets of vascular strands, one set on each side of the transversely
placed axis (Fig. 14F); with the lignification and eventual secondary thickening of the
stele the ovary is enclosed in the spike, which may become a portion of a branch, and
the seed is only shed on the death and eventual decay of the spike or branch.

Another noticeable variant is found in northern New South Wales near Yantabulla
{Benson and Pickard 1825). This plant is rather larger in all its parts than in the typical
S. tenuis and has firm spreading acute apices to the bracts.

24

Nuytsia Vol. 3 (1) 1980

J. M. Black (1919) noticed that the seeds in one plant he examined had rotated 90°
relative to the inflorescence axis. Material I have studied suggests that while rotation
sometimes does take place this is not usually as much as 90 , and is presumably a passive
response to the thickening of the axis.

Lectoiypification: Bentham (1870) cited four collections with his description of Salicornia
tenuis as follows: (1) Victorian Expedition: (2) Mitchell; (3) Howitt s Expedition, and
(4) McDouall Stuart's Expedition. The lectotype was selected by J. M. Black (1919)
as being the specimen collected on the Howitt Expedition (MEL 71381). The specimen
(in herb. K) from the Victorian Expedition is conspecific with the lectotype. The collection
from the Major Mitchell Expedition (“Interior of New Holland, 1838 , herb. K) is Saico-
cornia quinqueffoni (Ung.-Sternb.) A. J. Scott. The sheet at K of the collection made
on McDouair Stuart's Expedition contains three specimens, one being Halosarcia hciloc-
nemoides subsp. longispicatci P. G. Wils., a second H. pergrainilata ssp. elongala P. G. Wils.,
and the third a vegetative fragment of another (unidentifiable) Halosarcia species. The
description drawn up by Bentham combined in it the features of the various collections
seen by him. The specimens from the Howitt and Victorian Expeditions (S. tenuis sensu
lectotypico) Bentham considered to be of male plants, and those from the other expeditions
to be of female, but he expressed doubts as to whether the plants of the ‘two sexes were
conspecific.

5. Sclerostegia medullosa P. G. Wilson, sp. nov. Fig. 15.

Arlicu/i (internodia) cylindracei. 4-5 mni longi, 2-3 mm diam., demum medullosi, lobis acutis margini-
bus denliculaiis. Flores in axilla foliorum positi, fere verticale, ca. 2 mm longi, parte superiore dorsi-
ventraliter applanalo, orificio ciliatis; stylus robustus, in perianthio inclusus. Articuhis fructificans medul-
losus cylindracetis; perianthium mcmbranaceum: pericarpium crustaceum, ad stelam adnatum.

Type: Theldarpa-Yandama Stn. (ca. 30 mi W of Milparinka); bushy. 1-2 ft.; locally
frequent along small water course on gibber hill; 2 June 1955, L. A. S. Johnson and E. F.
Constable (holo: NSW 40039, iso. PERTH).

Much-branched subshrub to I m high. Articles cylindrical, 4-5 mm long, 2-3 mm
diam., papillose (at least when dry), becoming pithy with age; apex acutely lobed; margin
denticulate. Flowering articles interspersed among the vegetative and not differentiated
from them. Flowers almost vertical, ca. 2 mm long, dorsiventrally flattened distally
with the aperture terminal, united to each other and in proximal part to upper bract,
adherent to lower bract in proximal tw'o thirds; perianth membranous: anther ca. 2 mm
long: ovary narrow, membranous, fused to axis along vertical length of flower; style
stout, included in perianth: stigmas narrowly triangular. Fruiting articles pithy, cylindrical:
perianth membranous; pericarp hard and brittle ca. 2-5 mm long, adherent to stele of
axis. Seed vertical, ellipsoidal, ca. 2-5 mm long; testa membranous, colourless; embryo
slightly curved; perisperm copious, lateral.

Habitat: The only indication of habitat is that given on three herbarium specimens, of
which one, the type, is cited above; the second, from Whynot Stn. Queensland (Smith
6069), provides the note “growing among good grass on Borie (Acacia sp.) slope, near
but not along drainage line", while the third from near Woomera, South Australia (Lazor/r/ci
8401) indicates that it was “co-dominant with Atriple.x vesicaria and Frankenia on gently
sloping, stony floodplain".

Distribution: Southern Queensland, western New South Wales, and north of Port Augusta
in South Australia. The localities are too few and too scattered to obtain a clear idea
of the plant's distribution. Map 20.

Queensland: Whynot Stn. (38 km SW of Quilpic), L. 5. Smith 6069 (BRI); Whynot. 5. L. Everts! 5904 (K).

New South Wales: Wilcannia, Darling River. 1885, Mrs. Kennedy (MEL); Grey Ranges, {Baeuerlenl)
409 (MEL): “Koorningbirry" ( Koonenberry), W. Baeuerlen 220 (MEL); 5 km S of Binerah Downs —
Fort Grey road junction, S. Jacobs 3434 (NSW).

South Australia: "Near Spender's Gulf", 1881, Lattorf (MEL); Mt. Gunson, 31 30'S, 137°15'E, M. N.
Cole5\ (K); 7 mi NE of Woomera, M. Lazaridesm\ (PERTH); NE of William Creek, 28 21 'S, 136°17'£,
D. E. Synton 11313 (ADW),

Paul G. Wilson, Australian Salicornieae

25

Sclerostegia meduUosa is distinctive in having the triads of flowers scattered incon-
spicuously along the branches, the fertile articles being identical in outward appearance
to the vegetative. The divaricately branched habit is also a characteristic feature of this
species. The nature of the fruiting article is similar to that of S', moniliformis, for in
both species the pericarp is attached to the stele of the inflorescence axis and surrounded
by pithy bracts. S. mecliillosa differs from S. moniliformis in the shape of the articles
(and in their denticulate margins), the erect flowers and the thicker pericarp. The bracts,
with the enclosed seeds, remain attached to the stem for some time after the formation
of the fruit (probably for more than a year), but they eventually decay and allow the
pericarp with seed to fall away.

The specific epithet has reference to the pithy nature of the articles.

3. Tegicornia P. G. Wilson, gen. nov.

Herha prostrata perennis quasi aphylla. articulata, glabra. Folio succulenta opposita cauii adnata,
apice modice dilatata lobis minutis. Fiores erect! dioecii solitarii in axillis foliorum cauiinorum. Perian-
thium succulentum gamopetalum, ad apicem 3-lobum, ab foliis discretis, Flos masculiis: stamen solitarium
abaxiale ovarium vestigiale. Flos feniineiis: staminodium absens; ovarium erectum, stigmatibus anguste
lanceolatis, papillosis. Semen verticale; testa tenuiter Crustacea; embryo curvatus: perispermium copiosum,
laterale.

Type species: Tegicornia imiflora P. G. Wilson.

Perennial seemingly leafless succulent herb. Branches apparently jointed due to
the opposite leaves enveloping the stem in a succulent sheath; leaf-blade minute. Flowers
dioecious solitary in the axils of the stem leaves. Perianth fleshy, free from bracts, united
to summit; orifice terminal, 3 lobed. Male flower: stamen solitary abaxial; ovary vestigial,
on adaxial side of stamen. Female flower: staminode absent, ovary erect, attached to
base of perianth; stigmas 2, narrowly lanceolate, prominently papillose. Seed vertical,
testa thinly crustaceous; embryo curved; perisperm copious, lateral.

Origin of name: from the Greek word teges. a mat, and cornia, alluding to its relationship
to the plant Salicornia. The type species has a mat-like appearance.

One species endemic to southern Western Australia.

The genus Tegicornia differs from other members of the Australian Salicornieae in
the shape and manner of insertion of the perianth, and in the solitary unisexual flowers.
It is also unusual (although not unique) in having a prostrate habit and in bearing the
flowers on the undifferentiated stem articles. Its affinities appear to be nearest to the
genus Halosarcia as is suggested by the following characters; —

(1) Perianth 3-lobed with the median lobe in the inferior (abaxial) position.

(2) Stamen solitary and abaxial in position.

(3) Testa crustaceous; embryo curved; perisperm copious and lateral.

These characters in combination, agree with the genus Halosarcia but not with any
other member of the Salicornieae. The presence of a short rigid style and narrowly
triangular stigmas, as well as the position of the flowers, suggest a particular relationship
with those species of Halosarcia centred around H. priiinosa (Paulsen) P. G. Wilson.

1. Tegicornia uniflora P. G. Wilson, sp. nov. Figs. 4. 7, 16, 17. 36, 37.

Herha proslrata dioecia. Artieiili nitidi. anguste obovali, ad 15 mm longi. 6 mm lati (ad apicem),
versus basem angustali, lobis apicalibus rotundatis integris. Flores unisexuales, solitarii. Perianthinm
erectum. dorsi-ventraliter compressum. translucens. 3 mm altiim, in margine laterale succulentum abler
membranaceum: apice rotundatus, lobis parvis. Flos masculiis: antherum 2 mm longum; pistillodium
minutum, ca. I mm longum, stylo tenui ca. 0-2 mm longo. Flos femiueus: staminodium abaxiaie minutissi-
mum ovarium ellipsoideum succulentum, translucens, ca. 1-5 mm longum; stylus brevissimus stigmatibus
ca. 1 mm longis. Friictiis: perianthium di pateliiforme, ca. 3 mm altum, marginibus succulentis; peri-
carpium succulentum, ad perianthium adnatum. Semen late ovatum. ca. 2 mm longum; testa rufa, Crust-
acea, prominente concentrice pluricostata.

26

Nuylsia Vol. 3(1) 1980

Tvpe: Between Stirling Ra. and Porongurnp Ra.. Western Australia, 34 33'S. 11H4'E,
30 Oct. 1975. P. G. Wilson 11626. female plant (holo: PERTH; iso; AD. BRI, CANB,
K, MEL, NSW).

Prostrate perennial herb, dioecious. Arlicles on young branches glossy, succulent,
very narrowly obovoid, up to 15 mm long, 6 mm wide at apex, narrowed to a slender
base; apex with rounded lobes, entire. Flowers solitary, unisexual. Perianth erect,
strongly dorsiventrally compressed (transversely narrowly elliptical in cross section),
oblong in male flower, broadly obovate in female, succulent in lateral wing-like margins,
thin in centre, translucent, 3 mm high; apex rounded, with a small terminal aperture
surrounded by 3 small lobes consisting of two semicircular posterio-lateral outer lobes
and one anterior (abaxial) inner lobe. Male flower: anther 2 mm long: pistillode minute,
ca. I mm long, with a single slender style ca. 0-2 mm long. Female flower: staminode
a minute fleshy tube ca. 0- 1 mm long on abaxial side; ovary ellipsoidal, succulent, trans-
lucent, ca. 1-5 mm long; style very short; stigmas ca. I mm long, orange or yellow.
Fruiting perianth -t- patelliform (circular in face view) with solid succulent margins, ca.
3 mm high; pericarp succulent, soft, fused to perianth; seed vertical, usually radially
orientated, broadly ovate, ca. 2 mm long; testa reddish brown, crustaceous, with prominent
concentric porcate ribs over the embryo, smooth elsewhere; embryo curved; perisperm
lateral.

Figure 7. Tegiconiia iiiiiflora; on margin of salt lake between the Stirling and Porongurup Ranges Western
Australia. Photo by A. S. George.

Paul G. Wilson, Australian Salicornieae

27

Habitat: Growing in slightly saline loam periodically subject to inundation.

Distribution: Southern Western Australia, between the Porongurup Ra. and Ongerup.
Map 5.

tVesteni Australia: Salt lake 40 km NE of Ongerup, 16 Dec. 1973, K. Newhev 4014 (PERTH); Chillinup
Rd., S of Stirling Ra., Oct. 1975, N. G. Marchaui s.n. (PERTH); ibid., B. R. Masliu 4017 (PERTH); ibid.
A. S. George 14238 (PERTH).

The prostrate habit of this plant, and insignificant flowers, are probably reasons
for it being overlooked by collectors, for all the herbarium specimens seen, apart from
that collected by K. Newbey. came from the margin of the same salt lake sittiated between
the Stirling and Porongurup Ranges.

This species differs from all other members of the Salicornieae in being strictly
dioecious and in the flowers being solitary (not aggregated into small axillary clusters).
It is also unusual in having the flowers in the leaf axils and not arranged in an inflorescence.

The very short stele which serves the solitary flower of Tegicornia uniflora is similar
to that found in Halosarciu although much abbreviated in form. The stele divides to
form three prominent traces of which the central one passes into the funicle while the
two lateral traces pass into the base of the perianth where they terminate in globular
clumps of vascular tissue (Fig, 2A-B). The lateral traces arise in a manner similar to
that of the traces serving the lateral flowers in Halosurcia; if the traces in the two genera
are homologous it woufd indicate that the solitary flower in Tegicornia is derived from
a cymose inflorescence. The short ‘inflorescence’ stele also gives rise on its abaxial side
to a minute trace w'hich passes into the base of the flower.

The flowers of both sexes bear vestiges of the male or female organ. In the male
flower the pistillode is readily apparent even though small. In the female flower the
staminode is minute and consists simply of a fleshy scale-like disc on the abaxial side of
the ovary base; it is about 0- 1 mm in length (Fig, 1 1 D). That this is a rudimentary stamen
is suggested by its position and by the fact that no such scale-like structure is present
in the male flower.

Flalosarcia sp. x Tegicornia uniflora Fig. 18

Recently several collections have been made near Truslove (60 km N of Esperance)
of a plant which has the habit of Tegicornia uniflora but in inflorescence and floral mor-
phology is intermediate between that species and a generalised species of Halosarcia.
This plant exhibits considerable variation within the one population as may be apparent
from the following description;

Prostrate perennial herb, polygamo-monoecious. forming a mat-like cover up to
1-5 m diameter. Ultimate branchleis ascending, ca. 3 cm long: articles obovoid, ca.
5 mm long, 3 mm wide, somewhat laterally compressed, glossy, apex deeply lobed, the
lobes obtuse to acute or shortly acuminate. Inflorescence a terminal subglobular spike
of 2-4 bract pairs; lowest bract-pair united and similar to the vegetative articles; bracts
of the uppermost bract-pairs free, and of the middle bract-pairs slightly united or free
(sometimes very shortly 3-lobed); bracts when free deeply concave adaxially to infolded
(somewhat involute) and attached to axis by a small base (not decurrent). Flowers solitary
or in triads; lowest bracts 3-flowered. the central flower hermaphrodite, the lateral female
and often sterile (or rarely all three hermaphrodite); middle bracts 1-3 flowered, the
central flower female or hermaphrodite and the lateral female and usually sterile, or much
reduced in size and represented only by a vestigial perianth; upper bracts 1 -flowered,
or the terminal pair flowerless. Flowers free from bracts and from each other, steeply
ascending and attached by a small base. Perianth of central flower urceolate (but dorsi-
ventrally compressed); lateral margins fleshy; orifice terminal; lobes 3, the medial abaxial
lobe broad, usually within the lateral pair which are united on the adaxial side. Stamen

28

Nuytsia Vol. 3 (1) 1980

abaxial (in lateral flowers usually represented by a mintite staminode). Ovary ellipsoidal,
succulent, not appreciably changing in fruit, attached to base of perianth; style short and
stout; stigmas narrowly triangular. S'cer/ broadly elliptic, ca. 1-3 mm long, usually abort-
ing before reaching maturity: testa pale reddish brown (almost white when semi-mature)
with 2-3 prominent porcately sculptured ribs on the margins otherwise smooth.

Western Aiislralia: Truslove Reserve 27985 (ca. 60 km N of Esperance), 18 .Ian. 1978, G. J. Keigherv 1829
(PERTH); ibid., 16 Jan. 1979, C. J. Robinson and J. M. Koch (PERTH), from several plants. Map 5.

Habitat: Saline sand on margin of salt lake.

This taxon is recorded from a single population that is stated by one of the collectors
(C. J. Robinson, pers. comm.) to cover many hectares of saline soil. Robinson also
observed what he considered to be the same plant in a neighbouring area, but no collections
were made. Within the population there is variation in vegetative morphology, in in-
florescence shape and structure, and in floral morphology. The plant is anomalous in
several ways in that it does not conform to the structure found in any of the recognised
samphire genera. In particular the following characters make it unique within the Sali-
cornioideae; (1) The terminal (and often the penultimate) pair of bracts are free from
each other, their margins are often infolded, and they are attached to the axis by a small
base; (2) The spike often ends abruptly in this terminal pair of often flowerless bracts;
(3) The bracts are sometimes shortly 3-lobed; (4) The axillary cymules vary in their flower
number from 1-3; (5) The lateral flowers of a cymule are usually female and sterile, or
they may be vestigial and represented by a very reduced perianth. A further peculiarity
which has not been observed in other species is that the spike sometimes appears to end
in a terminal flower which is presumably axillary to one of the terminal bracts (while the
other bract is sterile) and becomes displaced to occupy a terminal and axial position
(Fig. 18-Ac).

In its general morphology, its flower structure and in the sculpturing of its testa,
this plant is similar to Tegicornia iiniflora, while the cymule character, the sexuality of
the flowers, and the nature of the inflorescence are reminiscent of the genus Halosarcia.
The variation in general morphology found within the population suggests that this plant
is of hybrid origin, a suggestion which is supported by the irregularity of some spikes and
the high degree of sterility of the flowers. In addition, most of the seed observed appeared
to be immature and to have stopped growing at an early stage; this is apparent from the
slightly decomposed nature of the surrounding tissue or of the seed itself.

Further field work is required before a conclusion can be reached as to what species
might be involved in producing this population but, as suggested earlier, it seems probable
that the plant is a hybrid between Tegicornia unifiora and a species of Halosarcia. At
present it seems unwise to give the putative hybrid a name, since its parentage has still
to be investigated; I therefore refer to it by a formula ; Halosarcia sp. x Tegicornia unifiora.

4. Halosarcia P. G. Wilson, gen. nov.

Arthrocnemnm sect. Leiusperma J. M. Black, Trans. Roy. Soc. S. Austral. 43: 358 (1919). Lectotype:
A. teiostachynm (Benth.) Paulsen ( H. indica subsp. leiostachya (Benth.) P. G. Wilson), lecto. nov.

Arthrocnemnm sect. Trachysperma J. M. Black, op. cit. 357. Lectotype: A. halocnemoides Nees H.
hatocnemoides (Nees) P. G. Wilson), lecto. nov.

Arthrocnemnm subgen. Angianthemnm Moss, J.S. Afr. Bot. 20: 5 (1954). Lectotype: A. indicnm (Willd.)
Moq. ( H. indica (Willd.) P. G. Wilson), lecto. nov.

[Arthrocnemnm auct. non Moq. s.str.: Ungern-Sternberg, Salic. Syn. 281 (1876); J. D. Hooker in Bentham
et Hooker. Gen. Plant. 3: 65 (1880) p.p.; Volkens in Engler et PrantI, Nat. Pflanzenf. Ill, la: 76 (1892);
J. M. Black, Trans. Roy. Soc. S. Austral. 43: 357 (1919); Ulbrich. in Engler et PrantI, op. cit ed 2 16c-
548 (1934) p.p. I

[Saiicornia auct. non L.: Bentham, FI. Austral. 5: 201 (1870) p.p.]

Frnticnii glabri; ramuli succulent!, internodiis ut videtur articulatis; chlorenchyma absque scleridibus.
Foiia opposita. in rima connata lobata redacta. Inflorescentia spicoidea; bracteae oppositae, connatae
vel liberae; cymulae sessiles, 3-florae. Fiores hermaphroditi (aut ovario vel stamine abortivi); perianthium

Paul G. Wilson, Australian Salicornieae

29

gamophyllum urceolatum vel varie compressum, lobis 3, duobus laleralibus et uno abaxiali; stamen solit-
arium abaxiale; ovarium sessile. Fnicliciili: perianthium membranaceum aut spongiosum aut medullosum
vel crustaceum; pericarpium membranaceum ad crustaceum. Semen erectum, ovoideum ad circulare;
testa membranacea ad crustaceam; embryo curvatus. radicula inferiori: perispermium copiosum laterale,

Arthroenemnm Moq. affine, a quo chlorenchymis absque scleribidus, floribus unistamineis differt.
Type species; Halosarcia halocnemohles (Nees) P. G. Wilson.

Dwarf glabrous shrubs. Branchlcis composed of succulent apparently articulate
internodes; sclereids absent from peripheral chlorenchyma. Leaves opposite forming
an insignificant fleshy bilobed rim (scarious on the margin) at the apex of each internode.
Inflorescence a spike-like cyme (thyrse). usually terminal to the branchlets, consisting of
three-flowered cymules each in the axil of a bract. Opposite bracts united or rarely
free, succulent. Flowers sessile, usually hermaphrodite (but the stamen or ovary some-
times failing to develop) or in some species female; perianth gamophyllous. initially
membranous or succulent, lobes 3 (two lateral and usually a medial abaxial); stamen
solitary abaxial, lilament initially turgid and terete, eventually lorate, sometimes indurated
towards the base: ovary vertical, membranous or succulent; style slender, 2 (3) lobed.
Fruiting perianth membranous, succulent, spongy, pithy, crustaceous, or corneous. Seed
erect, ovoid to circular; testa membranous to crustaceous; embryo curved; radicle inferior;
perisperm abundant, starchy, lateral.

In this genus 23 species are recognised of which all except one are endemic to Aus-
tralia; H. indica is also found in Malaysia and in those countries bordering the Indian
Ocean.

Origin of name: from the Greek words halos (salt) and sarx (fleshy).

The genus Halosarcia shows greater morphological variability than any other member
of the Salicornieae, and possibly any other member of the Chenopodiaceae. This vari-
ability is most obviously shown in the inflorescence, flower, and fruit, but is also apparent
in the anatomy of the vegetative parts. In particular there is a sharp distinction between
the anatomy of //. indica and of that of the rest of the genus. In H. indica the palisade
tissue of the articles forms a single layer and consists of groups of chlorenchyma cells
(around the stomata) interspersed with colourless cells; beneath the palisade layer is a
continuous sheath of small isodiametric chlorenchymous cells rich in starch (Fig. I).
In all other species of Halosarcia (and of the other members of the Australian Salicornieae)
the palisade tissue is of several layers which may either gradually pass into the aqueous
tissue or be sharply delimited from it.

The inflorescence varies from being contined to two or three intercalary articles
identical in appearance to the vegetative ones, to forming a terminal well-defined spike-
like thyrse. The flowers may be sunk into the succulent inflorescence axis or may merely
abut onto a relatively slender axis; they may be truncate or dorsiventrally compressed
at apex; and in fruit the perianth and pericarp vary in texture from membranous to pithy
through to crustaceous or horny.

In spite of this great variability it has not seemed practical to provide a formal sectional
treatment since there is no clear segregation of the species. Although in its stem anatomy
H. indica differs from the other species it shows less evidence in flower and fruit characters
of being sectionally distinct, and its seed is very similar in appearance to that found in
other Halosarcia species with a hard pericarp.

Several Halosarcia species are very variable within themselves and this is particularly
obvious when examining fresh material. Unfortunately the characters which can be used
in practice for discriminating the species do not allow full justice to be given to all those
characters which are obvious in the living plant, both in gro.ss appearance and in the
fine structure of the flower. For this reason the species have generally been broadly
defined and it is apparent that further work will enable many more to be discriminated;
this is important, since, as delimited here, a confused situation is presented as to the
ecological requirements of several of the broadly based species.

30

Nuylsia Vol. 3 (I) 1980

In addition to those 'incipient' species which remain hidden through a too broad
delimitation of otiters, there are many more which have not been recognised because of
the imperfect material which is available in herbaria; furthermore it is apparent from
recent collecting activities that some species are very restricted in their distribution and
it is probable that many of these remain to be discovered.

Hahsarda and oilier genera

The two characters which most obviously distinguish Halosarcia from Arthrocnemum
(i.e. absence of both sclereids and the adaxially placed stamen) are of tundamenlal import-
ance and suggest that these genera are not closely related. However, the characters
which distinguish Halosarcia from Sclerostegia, Padiycornia, Tegicornia and Teclicornia,
although relatively obvious, relate to organs which tend to be plastic in their morphology.
For this reason I consider that these five genera are closely related and, because of their
virtual endemism, that they have evolved from a common ancestor within Australia.

2 .

2*^

3.

3*

4.

4 =!=

5.
5*

6 .

6 *

7.
7 *.

8.
8 *
9.

9 #

10 .

10 *.

11 .

12 *.

13.

13*.

14.
14*.

Key to the species of Halosarcia

Opposite bracts free from each other (Fig. 24F) 2

Opposite bracts united (Fig. 24A) 4

Flowers intimately fused to each other and to upper bract (Fig. 26) 13. H. peltata

Flowers free from each other and from bracts 3

Seed pale brown, concentrically tuberculate over embryo (Fig. 52)

12. H. flabelliformis

Seed white with prominent uneven scale-like ribs covering whole of surface (Fig.

46) 9. H. pterygosperma

Perianth rounded to truncate at apex 5

Perianth dorsiventrally fiattened at apex . 14

Seed white; testa raised into scale-like ribs 6

Seed brown, reddish brown, or black, tuberculate or granular 7

Spikes long and slender, even in outline; seeds with smooth ribs around circum-
ference (Fig. 44) 8. H. lepidospernia

Spikes relatively short, undulate in outline; seeds with uneven ribs covering whole

of surface (Fig. 46) 9. H. pterygosperma

Seed dark reddish brown to black, covered with concentric porcate ribs (Fig. 42);

abaxial perianth lobe often prominent, external 8

Seed brown, variously tuberculate, abaxial perianth lobe small, usually inside

lateral lobes (external in H. fonlinalis) 9

Spikes even in outline (smooth); bracts ciliolate (Fig. 22F)

Spikes even or undulate in outline: bracts entire (Fig. 22A)

Bracts denticulate or ciliolate; axillary cymules 3-7 flowered

Bracts entire; axillary cymules 3-flowered

Axillary cymules 3-(1owered; seed concentrically tuberculate over embryo (Fig.

38-41) 1. H. halocnemoides

Axillary cymules 3-5 (7)-flowered ; seed concentrically porcate over embryo

(Fig. 67) 5. H. pluriflora

Perianth crustaceous in fruit; seed smooth (Fig. 50) 11. H. leptoclada

Perianth spongy to membranous in fruit; seed ribbed or tuberculate over embryo 12
Branches and .spikes very slender and smooth in outline and remaining so on
drying (resembling Casuarinu branchlets); fruit apex mammillate when mature

(Fig. 23) 10. H. lylei

Branches undulate in outline |3

Fruitlets failing entire (not tearing at base so as to expose seed); abaxial perianth

lobe external (Fig. 24C-E) 4. H. fontinalis

Fruitlets tearing at the base to expose seed; abaxial perianth lobe internal
(Fig. 19B) .... .. 1. H. halocnemoides

Margin of articles and bracts denticulate or ciliolate 15

Margin of articles and bracts entire 18

7. H. doleiformis
6. H. pergranulata

10

1 1

Paul G. Wilson. Australian Salicornieae

31

15.
15 ^

16.
16*.

17.
17*.

18 .

18 *.

19.

19*.

20 .
20 *.
21 .
21 *.
22 .

22 *.

23.

23*.

24.

24*.

25.

25*

Perianth in Iruit pithy; pericarp horny all over; fruitlets entire (not torn at base);

chlorenchyma tissue of stem as in Fig. 1 23. H. indica

Perianth thin in fruit, either mcmbranotis or cartilaginous; pericarp membranous

(at least towards base); chlorenchyma tissue of 2-3 rows of palisade cells 16

Articles with broad rounded lobes; fruitlets falling entire; perianth cartilaginous
(Fig. 31) 19 H. entrichoma

Articles with acuminate to caudate lobes; fruitlets tearing at base to expose

seed ; perianth thin, weak 17

Articles glossy, margin entire or crenulate; perianth entire at apical margin

3. H. nitida

Articles dull or glossy, margin fimbriate; perianth fimbriate on margin

2. H. finibriata

Perianth pithy in fruit, pericarp horny (at least towards apex); chlorenchyma

tissue of stem as in Fig. 1 23. H. indica

Perianth various (but not pithy) in fruit; chlorenchyma tissue of stem of 2-3

rows of palisade cells

Perianth papery or soft; fruitlets free from each other

Perianth crustaceous; fruitlets free or united

Articles shortly acuminate; testa tuberculate over embryo 21.

Articles truncate; testa smooth

Fruitlets laterally united

Fruitlets eventually free from each other and from bracts

Fruitlets persisting, released only on decay of perianth and bracts;

forming a hard cap to .seed (Fig. 25) , 22.

Fruitlets eventually splitting in medial sagittal plane to release seed;

not forming a persistent cap

Spikes terminating branches; articles glaucous, barrel-shaped (Fig. 27) .

15. H. auriculata

Spikes becoming intercalary; articles dull green, obovoid (Fig. 28) 16. H. syncarpa

Bracts truncate or slightly undulate on margin, cup-shaped at fruiting stage;
fruitlets muricate outside; spikes usually terminal to branchlets (Fig. 27)

14.

Bracts undulate, shrivelled at fruiting stage; spikes usually sessile
Fruitlets smooth and glossy outside; vegetative articles ca. 7 mm Iona

"^ 17 .

Fruitlets prominently muricate-areolate outside; vegetative articles ca.

19

20

21

H. cupuliformis
20. H. chartacea

22

24

pericarp

H. calyptrata

pericarp
23

long

H. pruinosa

.... 25

H. undulata

ca. 15 mm

18. H. bulbosa

1. Halosarcia halocnemoides (Nees) P. G. Wilson, comb. nov.
Arthrocnemum hulociieinoides Nees in Lehmann, PI. Preiss. 1 : 632 (1845).

Type: Fremantle, Preiss 1910 (iso; AD, K, MEL).

Spreading to erect suhsbruh to 0-5 m high. Branchlets slender; articles glossy to
dull (rarely glaucous), globular, doleiform or obovoid, mostly 3-5 mm long. 2-3 mm
diam., lobes rounded or attenuate, entire (denticulate in subsp. caudata). Spikes terminal
to main and lateral branchlets, 2-3 mm diam., the articles circular or laterally compressed
(when producing an uneven appearance to spike), opposite bracts united, undulate to
truncate, entire (denticulate in subsp. caudata). Flowers free from each other and from
bracts, exposed at apex, upper and lower surfaces vertical to spike axis (or the lower
slightly ascending). Perianth succulent; apex truncate to rounded; lateral lobes large
extending over length of apex; medial abaxial lobe small, hemispherical to triangular,
situated within lateral lobes. Fruiting perianth either soft and becoming shrivelled on
drying, or firm, chartaceous and translucent, free from bracts; pericarp thin, weak and
translucent, intimately fused to perianth. Seed discoid, ovate to elliptic or circular,
ca. I mm long; testa reddish brown, crustaceous, with small to prominent tubercles in
concentric rows over embryo, otherwise smooth to granular; embryo curved. Fruitlets
eventually breaking away from spike due to an abscission zone around proximal portion
of perianth where fused to radial septa of spike axis; seed protruding from torn base.

32

Nuytsia Vol. 3 (I) 1980

Distribution: Australia (excluding Tasmania).

Halosarcia iuilocnenwides is widespread in Australia and exhibits considerable vari-
ability. Those variants which are most clearly recognizable are here segregated as sub-
species, but there still remains much material which is far from homogeneous. In addition,
none of the subspecies is clearly demarcated since specimens can be found which are
intermediate in morphology between each of them and the heterogeneous “subsp. haloc-
nemaides" . The classification proposed here will therefore prove unsatisfactory if it is
desired to distinguish by name many of the regional variants, but it has not seemed profit-
able, with the presently available knowledge, to recognise further taxa, nor has it seemed
possible to provide critical characters for their differentiation. The most satisfactory
organ on which to base di.scriminating characters is the seed and only in a small proportion
of the specimens examined is this present in a mature state; therefore, within H. Iialoc-
nemoides (which is probably a species aggregate), infraspecific taxa have only been recog-
nised in those cases where morphological characters, additional to those found in the
seed, can be recognised and described.

The name Arthrocnennon halocnenwides was included by Bentham (1870) in the
synonymy of Salicornia arhiisctda ( - Sclerostegia arbuscida) and most of the specimens
he saw of the former species were also included here. His description of S. arbuscula
consisted of a combination of characters drawn from both species. J. M. Black (1919)
included as a variety of A. halocnenwides the plant here recognised as a distinct species
H. pergranidata, and later (1936) also the plant here recognised as H. pterygosperma.
Apart from considerable differences in habit and in vegetative and seed morphology,
//. halocnenwides diflers markedly from the latter two species in its perianth; the lower
lobe being small and included within the large lateral lobes in H. halocnenwides, whereas
in both H. pergranidata and H. pterygosperma it is large and outside the lateral lobes.

Key to Subspecies

1. Articles dull green, denticulate, lobes caudate Ib. subsp. caudata

1*. Articles dull or glossy rarely glaucous, margin entire, lobes rounded or obtuse

(rarely apiculate) 2

2. Spikes long and slender (up to 8 cm), even in outline; seed semicircular, smooth

on sides Ic. subsp. longispicata

2*. Spikes up to 2 cm long (rarely more), undulate or even in outline; seed smooth
or granular on sides . ... 3

3. Branchlets moniliform, slender; .seed ovate with smooth sides Id. subsp. catenulata
3*. Branchlets with articles narrowly cylindrical, doleiform, or obovoid; seed sub-

orbicular, granular on sides 4

4. Articles narrowly cylindrical to narrowly doleiform; spikes slender, even in outline

le. subsp. tenuis

4*. Articles obovoid (often broadly so), glossy; spikes undulate in outline

l.a subsp. halocnemoides

la. subsp. halocnemoides Fig. 19A-B, 38-41.

Erect much branched subshrub 10-15 cm high; branchlets soon losing the succulent
cortical tissue. Articles glossy (to dull), narrowly obovoid to sub-globular, 2-5 mm long,
apex obtusely 2-lobed to truncate. Spikes terminal, 0-5-2 -5 cm long, 2-3 mm diam., of
2 to 14 articles, the articles somewhat laterally compressed producing an uneven appearance
in spike; bracts undulate to truncate, entire: cymules disjunct or contiguous. Fruitlets
frequently protruding beyond bracts; perianth 'soft and spongy when dry (chartaceous
in some Eremaean material). Seed flattened, broadly ovate to suborbicular, ca. 11 mm
long; testa concentrically tuberculate over embryo, somewhat granular on sides.

Habitat: A wide range of saline habitats, each of which appears to be occupied by a different
variant. It is found in tidal salt marshes, around inland salt (NaCl) and gypsum lakes,
and brackish seepages. ’

Paul G. Wilson, Australian Salicornieae

33

Distribution: North-western Victoria; subtropical South and Western Australia. Map 6.

Victoria: Lake Tyrell, A. C. Beaiiglehole 55466 (PERTH); Pink Lakes, 10 mi NNW of Underbool, A. C.
Beaiiglehoie 40376 (PERTH).

Soulh Aasiralia: Port Pirie, H. W. Andrew 23 (AD); Ethelton, J. M. Black (AD); Stenhouse Bay, B. J.
Brock 14 (AD); Colona, H. W. Caidfield 149 (AD); Wilgena Stn., 26 Nov. 1925, J. B. Cleland (AD); Ooldea
Soak, 27 April 1951, y. B. Cleland (AD).

Western Australia: 2 mi S of Beenong, K. M. Allan 146 (PERTH), 2 mi E of Meckering, T. E. H. Aplin
664 (PERTH); Lake Ninan, T. E. H. Aplin 734 (PERTH); Murgoo, A. M. Ashhv 5364 (AD); Lake Anneen,
A. C. Beaiiglehole 49068 (PERTH); Mandurah, R. A. Saffrey 875 (PERTH);' Alfred Cove, Swan River,
P. C. Wilson 8698 (PERTH).

The considerable variation in the appearance of this subspecies would suggest that
further taxa should be distinguished. The organ which is least responsive to environ-
mental conditions, and therefore most suited for use in discriminating taxa, is the seed;
within the subspecies hulaawmoides can be recognised several seed variants but it has
not proved possible to clearly correlate (in a manner which can be described) these variants
with other morphological characters (although there is considerable geographical corre-
lation). The problem is increased because only a small minority of the herbarium
specimens studied had mature seed and were thus in a suitable state for making critical
comparisons. It would seem therefore that for the present it is preferable to apply a
broad taxonomic concept to the typical subspecies of H. haiocnemoidcs.

In the field subsp. hcdocnenwides may often be readily distinguished from other species
by its small size, compact habit and glossy appearance. Some of its variants appear
to be able to withstand periods of extreme salinity and it is frequently the only plant
present at the innermost zone of vegetation of an inland salt lake; here it may be repre-
sented by small non-flowering subshrubs with reddish sub-globular articles.

The type of H. halocnemoides was collected near Fremantle in Western Australia;
it consists of flowering material with very immature seeds but these do indicate the sub-
species group to which it belongs and allow comparison to be made with more recent
collections from the same area. The seed has a reddish brown testa w'ith concentric
rows of tubercles over the embryo and a .slight granulation over the perisperm (Figs.
40-41); this agrees with the seed of some recent collections from near Fremantle (e.g.
Saffrey 875) while other collections (e.g. Wilson 8698) have practically no tubercles (Figs.
38-39), or are intermediate in character.

In Victoria and in the southern part of South Australia the seed is ovate and similar
in ornamentation throughout the area although the plants themselves vary considerably
in vegetative morphology. They agree fairly closely with plants from the type locality
(Swan River, W.A.). Plants from the Eremaean region of South Australia, while having
similar seed to those from coastal situations, have a fruiting perianth which is firm, trans-
lucent, and deltoid in outline, whereas in plants from more equable climatic regions the
perianth shrivels at maturity. The variation in texture of the fruiting perianth is probably
an environmental response and not due to a genetic difference since there is some in-
consistency in plants collected from the same area but in different years.

The fertile articles in subsp. halocnemoides arc slightly laterally compressed which,
due to the decussate arrangement of the bracts, causes the spike to have an uneven appear-
ance. This character is most pronounced in specimens coming from the more southern
region of its distribution and helps to distinguish this subspecies from subsp. lenuis and
subsp. longispicata.

Hybridization between H. halocnemoides and H. pergranidala probably occurs since
in areas where the two species grow together may be found plants intermediate in habit
and with somewhat deformed flowers. The immature seed which is sometimes found
in these 'intermediate' plants appears to be developing characters of the testa which
cannot clearly be attributed to either species.

91670 — 0 )

34

Nuytsia Vol. 3 (I) 1980

lb. SLibsp. caudata P. G. Wilson, subsp. nov.

Articuli doleilbrmes vel obovoidi, ca. 3 mm longi et lati, opaci, papillosi; lobi caudati denticLilati.
Spicae 5-10 mm longae apicem versus angustatae: bracteae caudatae, denticulatae. Semen late ovatum,
lateribus laevibus.

Type: 9 mi S of Pin^rtip, Western Australia: salt loam; plant 12” to 18” high, dull green;
23 Nov. 1969. K. Newhey 3075 (holo; PERTH, iso: CANB, K).

Suhshmh ca. 0-3 m high; densely branched. Articles doleiform to obovoid, ca.
3 mm long and wide, dull green, papillose; lobes caudate, denticulate. Spikes 5-10 mm
long, ca. 3-5 mm diam., narrowing towards the apex, circular in cross-section; articles
deeply concave over cymules, caudate, denticulate. Seed broadly ovate, 1-0 mm long,
strongly tuberculate over embryo, smooth on sides over perisperm.

Habitat: Margins of inland salt-lakes.

Distribution: Lake Grace-Norseman area of Western Australia Map 8.

Western Australia: Lake King, A. S. Georse9357 (PERTH); Lake King, 22 Sept. 1976, R. Hnatiiik (PERTH)-,
Lake Cobham, K. Newhey 3315 (PERTH); Lake Grace, P. G. Wilson 8273, 8276 (PERTH); 93 km N of
Esperance, P. G. Wilson 10125 (PERTH); 6 km N of Lake Grace township, P. G. Wilson 10241 (PERTH).

This subspecies may be recognised by its dull appearance and by the caudate and
denticulate margins of its articles. Material intermediate between subspecies caudata
and the typical subspecies has been collected in two areas where the two subspecies are
found growing together. The intermediate collections are as follow's; Lake Cobham,
K. Nen'bey 3313, 3314, 3316 (PERTH); 9 mi S of Pingrup. A'. Newhey 3077 (PERTH).

A plant very similar in appearance to subsp. caudata has been collected at Lake
Miranda (ca. 130 km S of Wiluna, Western Australia): it differs, however, in its seed which
is noticeably granular over the perisperm. Eor the present this must be considered as
one of the indefinite members of the H. halocnemoides complex.

Ic. subsp. longispieata P. G. Wilson, subsp. nov.

SaUcornia tennis [non Benth.] Benth., FI. Austral. 5: 204 (1870) p.pte., not as to lectotype.

Artienli tloleiformes ca. 5 mm longi, 2-5 mm diam., apicem leviter rotunde lobati. Spicae tenues
aequatae, anguste cylindraceae, 2-8 cm longae, 3 mm diam.; bracteae leviter rotunde lobatae Frncticiili-
perianthium chartaceum, firmum, spiceum transverse oblongum. Semen semicirculare, pallido brunneum
supra embryonem tuberculatum aliter laeve. ’

Type: Finke River near .lunction with Palm Creek, 24 06'S, 132 52 'E. Northern Territory;
on shingle of river bed; stems semi-prostrate; 24 Aug. 1967. B. G. Briggs 1257 (holo:

Subshrub ca. 0-3 m high. Articles barrel-shaped, ca. 5 mm long, 2-5 mm diam
apex slightly and obtusely lobed, entire. Spike slender, even and cylindrical, 2-8 cm long’
3 mm diam., bracts very slightly lobed. Fruit/ets: perianth chartaceous but firm when
mature, transversely oblong at apex (central fruitlet). Seed semi-circular, I -0 mm long
pale brown, tuberculate over embryo, smooth on sides. ’

Habitat: Poorly drained somewhat saline situations.

Distribution: Central Atislralia. Map 7.

Queensland: Pulchera Waterhole, 30 mi WNW of .Sandringham, P. K. Latz 5\9 (PERTH).

New South Wales: " Western interior of New South Wales", 1880 C. Moore (MEL 71387).

South Australia: Coward Springs, ,3 1 Dec. 1926,7. B. Clelaiul (\E»-, Beresford, 25 Apr. 1950 7 B Cleinnd
(AD); Moolawatana Bore. R. H. Ai/c/ie/ 2672 (AD); N.W, interior of South Australia 7 McDouall Stnnrt
(K, MEL 71379); Dalhousie Springs. 5. A. White 117 (AD). Me Douall Stuai t

Northern Territory: Erldunda Stn.. T. S. HenshaU 55 (NT); Napperbv Stn.
Finke River, ex herb, F. Mueller (AD 97543201 p.pte.).

T. S. HenshaU 1015 (NT);

The collection HenshaU 1015 cited above differs from the others in having shorter
and thicker fruiting spikes; in seed characters it closely resembles the type.

Paul G. Wilson, Australian Salicornieae

35

A plant which is very similar to, and possibly sympatric with, subsp. loii^ispicata
should probably be included under that subspecies. In the dried state the only obvious
differences between the two are found in the seed which in the variant is reddish brown
and broadly ovate. From the few ecological notes accompanying the specimens it would
appear that this variant is found in very saline situations. The following specimens
have been seen;

Northern Territory: Napperby Salt Lake. C. Diiiilop 2353 ( PERTH ) ; Lake Bennet, C. Diwhp 2435 (PERTH ) ;
6 mi S of Mongrel Downs, J. R. Maeniiochie 952 (PERTH); I mi W of Central Ml. Wedge, A. O. Nichol/s
819 (PERTH); near MacDonnell Ras., 1889, 5c/;n'arr (MEL).

A variant of H. hulocnemohles subsp. temds has been collected in the same localities
as subsp. longispicata. They differ from each other in habit (subsp. tenuis is a small
open plant with divaricate branches and thick articles) and in seed (circular with granular
sides in subsp. tenuis). The few collections seen do not indicate whether there is hybridiz-
ation between the two subspecies.

Herbarium material of subsp. longispicata is, at the flowering stage, very similar to
that of H. pergranulcita subsp. elongatu. The two may be readily distinguished by their
perianths: in H. pergranulata the perianth has a large semi-circular outer lower (abaxial)
lobe while in subsp. longispicata it has a small inner lower lobe. In the fresh state they
are easily recognisable since H. pergraintlala is bluish-green and glaucous in colour while
H. halocneinoicics subsp. htngispicata is glossy green (or red). In seed characters, the two
plants are very different.

The syntype material of SuHcornia tenuis Benth. includes representatives of four
different species. The name was lectotypified by .1. M. Black (1919) on a How'itt collection
which is now referred to .Selerostegia tenuis (Benth.) P. G. Wilson. One of the specimens
among those syntypes of ,S'. /«»//.? collected on .1. McDouall Stuart's expedition (1859-1862)
is of H. hahenenwides subsp. longispicata: duplicates are in Herb. K and MEL.

In the vicinity of Port Pirie, South Australia, is found a plant which in the dried
state somewhat resembles subsp. longispicata and should possibly be included in it. The
plant at Port Pirie is found in tidal marshes, conditions very different to those found in
Central Australia, and this may account for some of the morphological variation; in seed
characters the maritime and inland forms show close similarities.

Id. subsp. catenulata P. G. Wilson, subsp. nov. Fig. I9C-D.

Articuli nitidi (vel glauci), doieiformes, 2-3 mm longi, 3 mm diam., ad nodes constricti, margine
integro, truncati vel leviter undulati. Spicue tenues, 5-10 mm longae, apicem versus leviter angustatae.
Semen ovatum, lateribus laevibus.

Type: 6 km S of Lake Barlee HS., 29 ID'S, II9°6'E, 26 Aug. 1970, P. G. Wilson 8849
(liolo: PERTH).

Compact suhsliruh to 0-5 m high. Branchlets slender, articles glossy (or glaucous),
doleiform, 2-3 mm long. 3 mm diam., constricted at the nodes, margin entire, undulate
or truncate. Spike terminal to short branchlets, slender with a smooth outline, narrowing
slightly towards the apex, 5-10 mm long, of ca. 6 articles. Seed ovate, I mm long, reddish
brown, smooth on sides over perisperm.

Habitat: in heavy soil, not strongly saline.

Distribution: Mt. Magnet-Meekatharra (south to Lake King, glaucous variant); Map 7.

Western Australia: Yandil, Sepi. 1939, W. E. Blackall (PERTH); Lake View Stn., C. A. Gardner 7811
(PERTH); 15 mi SW of Nannine, N. H. Speck 729 (CANB. PERTH); 48 km N of Mt. Magnet, P. O.
Wilson 8564 (PERTH); 12 km N of Glen Stn., P. G. Wilson 8584, 8585 (PERTH); 1 km S of Yuin H S,
P. G. Wilson 9951 (PERTH).

This subspecies may be recognised by its slender branchlets (whose articles retain a
barrel-shaped appearance even in the dried state), by the slender, smooth spikes, and
by the ovate seeds with smooth sides.

36

Nuytsia Vol. 3 (I) 1980

The typical variant of subsp. cateiuilata is glossy; however a similar but strongly
glaucous plant has been collected at Lake Barlee and Lake King. This variant represents
the only case in H. halocnemoides where the plant is glaucous; in all other variants of
this species the articles are either glossy or merely dull. The flowers and seed of both
typical subsp. catenukita and the glaucous variant appear to be nearly identical. The
following three collections are of this variant;

Western Australia: S side of Mollerin Lake, /i. G. Muir 550 (PERTH); Lake King, east bank near Causeway,
P. C. Wilson 9991 (PERTH); southern margin of Lake Barlee, P. G. Wilson 8814 (PERTH),

The herbarium material of the glaucous variant of subsp. calenulata cited above has
a distinctive appearance. Other collections, however, suggest that this plant integrades
with other variants of H. halocnemoides which are also found in the Lake King district
of Western Australia.

The subspecific epithet refers to the resemblance of the articles to the links of a chain.

le. subsp. tenuis P. G. Wilson, subsp. nov. Fig. I9E-F.

anguste doleiformes, ca. 5 x 2 mm, opaci vel nitidi, apicem, leviter lobati interdum apiculati,
margine integro. Spicae fere cylindraceae 10-25 (70) mm longae, 2-3 mm diam., apicem rotundatae vel
oblusae; bracteae leviter undulalae, super cymulas profunde concavae, margine integro. Friiciiculi:
perianthium tirme chartaceum, leviter exsertum. Semen sub-orbiculare, porphyreum, lateribus granul-
aribus.

Type: Near aerodrome, Wyndham Township, Western Australia; salt flat; spreading
succulent plant 9” high; 20 July 1949; R. A. Perry 2558 (holo: CANB, iso: AD, BRF,
NSW, NT, PERTH).

Compact to open subshrub ca. 0-3 m high. Branchlets slender, spreading; articles
narrowly barrel-shaped, ca. 5 x 2 mm, dull green (not glaucous) to glossy, apex slightly
lobed, margin entire. Spikes almost cylindrical, 10-25 (-70) mm long, 2-3 mm diam.,
apex rounded or obtuse, bracts slightly undulate, deeply concave over cymules, margin
entire. Florets slightly protruding beyond bracts. Fruiting perianth firmly chartaceous
outside, pithy within, slightly protruding from between bracts. Seed suborbicular with
prominent radicle, 0-8-1 mm long, finely tuberculate over embryo, granular on the sides,
reddish brown.

Habitat: Coastal or near coastal, frequently on tidal flats, (the central Australia variant
on “semi-saline flats” and around salt springs).

Distribution: Coastal region of tropical Australia from Carnarvon (Western Australia)
north and east to Wide Bay district (Queensland); also (Central Australian variant) north-
ern South Australia and southern portion of the Northern Territory. Map 8.

Queensland: Alligator Creek near Townsville, W. Macnae 12.1 (BRI); Shell Ridge, 'Wernadinaa’ S
Jacobs 1386, 1387 (NSW); Elliott Heads, L. S. Smith 489 (BRI); Karumba, G. Trapnell 199 (BRI).

Western Australia: Broome, A. C. Beatigiehole 48201 p.p. (PERTH); Nobbys Well, W. V. Fitzgerald i'HS.Vy')-
Derby, R. D. Royce 3335 (PERTH); Wyndham, C. A. Gardner 7316 (PERTH).

Northern Territory: 10 mi NE of Leguna Stn., Victoria R., R. A. Perry 2590 (AD, BRI, CANB, MEL).
Central Australian Variant

South Australia: Cootanoorina, 5 May 1891, R. Helms (AD, MEL).

Northern Territory: Tanami Desert, A. C. Beauglehole 50927 (PERTH); Tanami Sanctuary T. S Hcnshall
1232 (PERTH); 1 mi W of Central Mt. Wedge, A. O. Nicholls 821, 823 (PERTH).

This subspecies is most distinct in north-eastern Queensland where it is a sprawling
subshrub of coastal mud-flats. There it has terete branches and slender spikes 2-3 cm
long; the seed is sub-orbicular and is ornamented with small tubercles over the embryo
which grade into the granulations on the sides. The plant found in the coastal swamps
of the Northern Territory and at Wyndham in north-east Western Australia is very similar
to the Queensland plant but further south along the Western Australian coast, e g at Derby

Paul G. Wilson, Australian Salicornieae

37

and Broome, the position becomes complicated by the presence of plants which are
evidently the northern variant of subsp. ha/ocnenioic/es. In this region of overlap the
two subspecies are sympatric and, in fact, grow intermixed in the same tidal mudflats
(Fig. 28). They have different (but overlapping) flowering times (that of subsp. tenuis
being earlier in the winter than that of subsp. halocnemokles) but plants have been collected
which are intermediate between the two subspecies in habit, vegetative morphology,
and seed characters, so that it would appear likely that hybridization does occur.

//. halocnemoides subsp. tenuis is the only member of this species recorded from eastern
and northern Queensland, the northern half of the Northern Territory, and from the
north-east Kimberley area of Western Australia; no variability between the different
populations has been observed in plants from these areas, nor is it likely to occur since
they are isolated from the other subspecies.

A plant having an open divaricate habit, similar to that of the type variant of subsp.
tenuis, is found in Central Australia. It has similar seed to the coastal plant and differs
principally in having thicker articles. This Central Australian variant is here included
within subsp. tenuis but further work may show that it should be recognised as a distinct
taxon.

Many of the specimens of this subspecies which come from the north coast of Western
Australia bear spherical galls made up of numerous separate flower-like segments. This
structure has not been observed on any other species, or on any of the other subspecies
of H. halocnemokles: when present it therefore enables one to recognise ssp. tenuis even
when only vegetative material is available.

2. Halosarcia fimbriata P. G. Wilson, sp. nov. Fig. 20A-C.

Fniticiilosiis ad I m altus. Articidi late obovoidei, ca. 3 mm longl, valde lobati, lobis acuminatis
saepe caudatls, margine fimbriato. Spicae breves, in ambito undulatae, 5-10 mm longe; articuli profunde
lobati fimbriati. Flores ad apice expositi ad vicinas liberi, dorsiventraliter applanati truncati fimbriati.
Perianihiiim tenue; lobus abaxialis parvus, lobi laterales prominentes lobos abaxiali superpositus. Spica
fructifera sicca, bracleis tenuibus. floribus occultist perianthium tenue, debile; pericarpium membranaceum
ad perianthium coalescens. Semen late ovatum ca. 1 mm longum; testa tenuiter charlacea, porphyrea,
super radiculam, tuberculata aliter laevis.

Tvpe: 5 km S of Morawa, Western Australia. 4 Sept. 1970, P. G. 9968 (holo:

PERTH, iso: CANB, K).

Suhshruh to I m high with ascending branches. Articles dull pale green or glossy,
broadly obovoid ca. 3 mm long, deeply lobcd, the lobes triangular, acuminate and usually
shortly caudate; margin prominently fimbriate. Spikes short and in outline prominently
undulate due to the lateral compression of the articles, 5-10 mm long with 2-6 articles,
at first terminal but usually becoming intercalary; fertile articles (bracts) deeply lobed
and fimbriate, becoming dry and thin as fruit matures, completely obscuring flowers.
Flowers ascending, to 1-5 mm long, free from each other but coalescent to upper bract
near axis. Perianth thin, apex dorsivemrally flattened and fimbriate, with a pair of
large lateral lobes on abaxial face and a small inner medial abaxial lobe. Ovary mem-
branous. Fruitlets enclosed within bracts, free from each other and from bracts, eventually
tearing away from axis exposing seed at their base; perianth thin, weak; pericarp mem-
branous and coalescent to perianth. Seed broadly ovate, I mm long; testa thinly chart-
aceous, reddish brown, concentrically tuberculate over embryo otherwise smooth; embryo
curved; endosperm lateral. Seed released on breakdown of perianth and pericarp at
spike axis.

Habitat: Margin of salt lakes, either of common salt or gypsum.

Distribution: Lake Annean south-west to Three Springs, Western Australia. Map 5.

Western Australia: N side of Lake Austin, A. S. George 793 (PERTH); S side of Lake Austin, ,•). S. George
769 (PERTH); Lake Annean, R. Hnatink s.n. (PERTH); 1 km S of Three Springs, D. Lowry (PERTH); 3 mi
S of Three Springs, B. R. Muslin 735 (PERTH); Carnamah, A. Morrison 16135 (K); Yarra Yarra Lakes,
M. D. Tindale 1292 (NSW, PERTH); 23 km W of Wubin, P, G. Wilson 6480 (PERTH); 6 km S of Dam-
boring, P. G. Wilson 6492 (PERTH).

38

Nuylsia Vol. 3 (1) 1980

The flowering spikes of this species soon become intercalary due to vegetative growth
at their apex. On the maturation of the seed the bracts, perianth, and pericarp become
dry and thin; eventually the fruitlets tear away from the spike axis to expose the seed
which is released. The vegetative and fertile articles have a noticeably fimbriate margin
Irom which character the specific epithet is derived.

While H. fimhriata in its typical state has a very distinctive appearance, two collections,
that of Maslin 735 and Wilson 6492, are intermediate in morphology between this species
and H. halocnenwides suggesting that introgression does occur between the two taxa.

Of those species of Halosarcia in which the perianth ‘tears' away from the axis, H.
fimhriata is probably most closely related to H. luilocnemoides, for their seeds are very
similar; in addition, in both species the perianth has a small inner abaxial lobe; however,
the apex of the perianth in H. luilocnemoides is truncate whereas in H. fimhriata it is
dorsiventraliy flattened.

3. Halosarcia nitida P. G. W ilson, sp. nov.

Fniticn/iis ca. 0'5 m altus. AnicuH nitida ca, 5 mm longi, margine integro vel liveter crenulato, lobis
breviter acuminatis. Spicae terminales, anguste cylindracei; bracteae cupuliformes. iruncatae. margine in-
tegro vet leviter crenulato, lobis breviter acuminatis. Fiores liberi, valde adscendentes, parte di.stali dorsi-
ventraliter applanatus. Periainhiiiin tenue, margo terminali rotundato vel acuminato, lobis duo, lateralibus,
conduplicatis, imbricatis. FructicuH demum secedentes semine expose; perianthium tenue, nec induratum;
pericarpium chartaceum, ad basim membranaceum; stylus ad basim durus. Semen ovoideum, I mm
longum; testa valde tenuis pallido tusca supra embryo concentrico granularis aliter laevis.

Type: South Australia, between Lake Torrens and Lake Gairdner; at the channel at the
south-eastern end of Pernatty Lagoon; 23 Oct. 1966, HJ. Eicliler 18855 ( AD96650459).

fiiruh ca. 0-5 m high. Branches erect, articles glossy, ca. 5 mm long, margin entire
to slightly crenulate, lobes shortly acuminate. Spikes terminal; bract pairs resembling
the vegetative articles but shorter, ca. 4 mm long, becoming cup-shaped with age. Flowers
hermaphrodite, free from each other and from bracts, ca. 3 mm long, steeply ascending,
distal half dorsiventraliy compressed and slightly winged, apex just exposed. Perianth
thin (at least when dry), terminal margin rounded to acuminate; lobes two, lateral and
conduplicate, imbricate, thin and soft in fruit. Utricle ellipsoidal, papery, membranous
towards the axis: .style with hard base. Seed ovoid, 1 0 mm long; testa very thin, pale
brown, concentrically granular over embryo otherwise smooth. Fruitlets eventually
tearing away from axis at base.

Habitat: Strongly saline or gypseous soil.

Distribution: North-western Victoria, central South Australia. Map 19.

Victoria: Nowingi, NW Victoria, 30 Nov. 1949, A. M. 0'A'c///(MEL 70631).

South Ausnutia: SE corner of Lake Gairdner. J. Pickard 246S. 2469 (NSW); between Lake View Dam and
Lake Frome, Frome Downs, D. E. Symon 8030 (ADW, PERTH); S shore of L Frome 18 Sept 1972
[). A. Webb (CANB), '

The nature of the perianth in H. nitida is interesting in that it is intermediate in
structure between the type found in H. itnditlala and that found in H. halocnemoides.
in the material included in this species may be found flowers with only a pair of terminal
conduplicate lobes, ranging to flowers with imbricate lobes covering the orifice on the
abaxial rounded surface but extending to the adaxial; in the latter case a small inner
abaxial lobe is present. In those specimens with the conduplicate lobes the imbrication
is of the type found in II. itndiilata (the lateral florets having the lobe nearest to the central
floret overlapping the farthest away), while in those specimens with the imbricate lobes
the imbrication is of the type found in H. halocnemoides (the lateral florets having the lobe
nearest the central floret iinderlapped by that farthest away). Some specimens, as might
be expected, have an indefinite imbrication.

Paul G. Wilson, Australian Salicornieae

39

In spite of the variability of the perianth structure I am treating the specimens as
being representatives ot a distinct species because I am unable to suggest from which
taxon (apart from H. halocncmoitlcs), H. niiicla could be derived. In addition, the field
notes suggest that sizeable populations of the taxon were present at two of the sites.

The manner in which the perianth is attached to the spike axis is similar to that found
in H. halocnemoides but the seeds of the two species are quite different, those of H. nitida
being almost identical to the seeds of H. syncarpa.

The specific epithet alludes to the glossy appearance of the plant when fresh.

4. Halosarcia fontinaiis P. G. Wilson, sp. nov. Fig. 24C-E, 66.

Fruticulus at 70 cm altus. Rainiili tenues; artieuli opaci vel glauci, margine obtuse lobato, integro.
Spicae tenues, attenuatae, 3 -8 cm longae, ca. 3-5 mm diam ; artieuli breviler cloleil'ormes. margine undulato
integro. Flores ad vicinas et bracteas liberi. leviler adscendentes, apicc rolundato. Pt’i Uiiilliiiiiit succulen-
tum trilobatum: lobi laterales breves; lobus medius aba.xialis prominens, semieireu laris, lobos iaterales
superposilus. I'niciiis: perianthium medullosum (vel interdum chartaceum ?). ba.sim versus membran-
aceum; pericarpium membranaeeum ad perianthium connalus. Semen obtuse ovatum, ca. 1 mm longus;
tesla Crustacea, atroporphyrea. super radicula cum costis concentricis prominenlibus porcatis ornata,
aliter laevis. Fniciiciili integri, .semene includenti ad pericarpium adhaerenti secedentes.

Type: South Australia, Dalhousie Springs, 24 Sept. 1974, D. E. Srmon 9302 (holo; ADW
47816, iso; PERTH).

Suhshruh to 70 cm high. Braiuiilets slender; articles dull to glaucous, margin bluntly
lobed, entire. Spikes terminal to main and lateral branchlets, erect, slender, attenuate,
3-8 cm long, ca. 3-5 mm diameter; articles shortly barrel-shaped, margin undulate, entire.
Flowers hermaphrodite, free from each other and from bracts, somewhat ascending;
apex rounded, 3-lobed, 2 short lateral and a semicircular outer abaxial medial lobe;
perianth and ovary succulent. Fruiting perianth shrivelled, pithy (or sometimes firm and
chartaceous?) at apex, membranous towards axis, fused to the membranous pericarp.
Seed bluntly ovate, ca. I mm long; testa crustaceous, porcately ribbed over embryo,
smooth on sides, dark reddish brown. Fruitlets shed entire; seed enclosed by and adherent
to pericarp.

Habitat: All collections seen refer to the plant as grow'ing around springs, presumably
saline.

Distribution: northern area of South Australia. Map 9.

South Austratiu: Nilpena, 2 May 1891, R. Helms (AD. MEL, NSW); Dalhousie Springs, P. K. Latz 4792
(NT); ibid., Sept. 1974, Mines Dept. (ADW); 50 mi NE of William Creek, 25 Aug. 1969, A. C. Robinson
(AD).

This species resembles H. pergranutata stibsp. elonguta but when in fruit may be
readily distinguished by its distinctive seed. The fruiting perianth is. in the material
seen, always shrivelled in the dry state but it appears likely that, as with H. pergraniilata,
under the right conditions it becomes firm and chartaceous on the outside. The perianth
when ripe falls entire; it does not tear away from the spike to expose the seed as does
the perianth in both H. halocnemoides and H. pergraniilata.

The specific epithet refers to the association of this plant with springs.

5. Flalosarcia pliiriflora P. G. Wilson, sp. nov. Fig. 67.

Fruticulus ca. I m altus. Rami graciles; artieuli opaci vel glauci, lobis aliquantum acutis, margine
denticulato. Spicae graciles; artieuli breviter doliiformi. lobis aliquantum acutis, margine denticulate.
Cymulue 3-5 (7)-llorae; Itores ad vicinas et bracteas liberi. Perianthium versus apicem succulenlum (sed
lato abaxiali tenui); apex rotundalus. lobis 3, irregularibus, denliculatis; ovarium membranaeeum. Fructi-
culi: perianthium medullosum vel chartaceum; pericarpium membranaeeum ad perianthium conjunctum.
Semen semi-circulare. I mm longum; testa atroporphyrea cum costis concentricis porcatis supra radiculam
ornata.

Type: South Australia, Lake Eyre, May 1951, C. W. Bonython 1194 (holo: ADW).

Subshrub ca. I m high. Branches slender; articles dull to glaucous, lobes somewhat
acute, margin denticulate. Spikes slender, terminal to branches and short lateral branch-
lets, erect; articles shortly doieiform, lobes somewhat acute, margin denticulate. Flowers

40

Nuytsia Vol. 3 (I) 1980

androdioecious, 3-5 (7) in each of the opposite bracts (when more than 3 the lateral ones
reduced in size), somewhat ascending, free from each other and from bracts; perianth
prominently fleshy towards apex on upper (adaxial) and lateral margins, thin below
(abaxially) and towards base; apex rounded, lobes 3 at apex, irregular, denticulate, the
abaxial one small and usually within the lateral lobes; male flower with small pistillode,
the stigmas exerted but minute (ca. 0-2 mm long); hermaphrodite flower with linear
stigmas ca. I mm long, ovary thin, eventually fused to the perianth. Fruitlels with pithy
or chartaceous perianth; pericarp membranous and fused to perianth. Seed semi-circular,
ca. I mm long, porcately ribbed over embryo, smooth on sides, dark reddish brown.
Mature fruitlets eventually breaking away from spike axis to expose seed at their base.

Habitat: Around inland salt lakes and salt springs.

Distribution: Central and northern South Australia; north-eastern New South Wales.
Map 10.

New South Wales: Warroo, via Bourke, 25 Oct. 1936. K. I. Murriss 4 (BRI).

South Australia: Moolawatana Bore. R. M. Kuche! 2669 (AD); Dalhousie Springs, P. K. Latz 4793 (NT);
ibid., D. E. Svnwu 3254 (ADW, CANB); ibid., D. E. Svrnon 9312 (ADW, PERTH); S of Scrubby Peak,
Lake Hart, May 1919, G. Tayhr 19 (AD); Gawler Range, P. G. Wilson 541 (AD).

Halosarcia phtriflora is closely related to H. fontinalis as is evidenced by its flower
structure and seed. It is peculiar in frequently having more than three flowers in each
axillary cymule and in the irregularity of the perianth. This species may be distinguished
from all others by a combination of characters involving its seed, its free florets, and the
denticulate margin to the leaf-lobes and bracts.

One of the specimens examined (Symon 9312) was of a male plant with vestigial
ovaries, but in those other specimens in which the anthers and ovaries could be observed
these appeared to be functional.

The method of separation of the fruitlets from the spike is the same as that found in
H. haiocnemoides, H. pergraniiiata, and H. pterygospenna. The chartaceous perianth
retains its shape and tears away from the axis due to an abscission zone between the axial
septa and the fruitlet; the seed is exposed at the ruptured base.

The Taylor specimen from Lake Hart, which is cited above, is one of the several
syntypes of Artlirocnennim haiocnemoides \aT. pergraniilatum J. M. Black.

The specific epithet refers to the presence of up to 7 flowers in a bract axil.

6. Halosarcia pergranulata (J. M. Black) P. G. Wilson, sp. et stat. nov.

Ar^hrocnemum haiocnemoides var. pergraniilatum J. M. Black, Trans. Roy. Soc. S. Austral. 43: 359, t.33

Lectotype: Railway bridge between Pt. Elliot and Victor Harbour, 12 Feb. 1919, H. W.
Andrew (holo; AD 97614357), lecto. nov.

Subshrub, erect or spreading (sometimes decumbent), ca. 50 cm high. Articles
cylindrical, barrel-shaped, or obovoid; lobes short, rounded or obtuse; margin entire,
dull green or glaucous (never glossy). Spikes terminal to main or lateral branchlets
(rarely intercalary) to 4 cm long; articles cylindrical, barrel-shaped, or obovoid, frequently
laterally compressed, the bract-pairs united, entire; triads of flowers included within bracts
or their apices exposed. Flowers hermaphrodite (or in some variants only female flowers
present), tree from each other and from upper bract or fused to a varying extent; abaxial
surface ascending, apex curved, and -J; continuous with abaxial surface, to truncate;
perianth succulent, lobes 3, 2 lateral and one lower rounded (abaxial) lobe which is either
inside or outside the laterals; pericarp succulent, style slender. Fruitlets free from each
other and from upper bract or ± united; perianth fleshy, spongy, or firmly chartaceous
on the surface with pithy interior; pericarp fused to and similar in texture to perianth.
Seed broadly ovate to circular, ca. 1 mm diameter; testa crustaceous, dark reddish-brown

Paul G, Wilson, Australian Salicornieae

41

to black, concentrically ribbed and porcate all over (in one variant not ribbed in centre);
embryo curved. Perianth and pericarp eventually either breaking down near base, thus
releasing the separate fruitlets from which the seed protrudes, or remaining attached to
upper bract and spike axis.

As circumscribed in this revision H. pergrainilata encompasses numerous variants
some of which are so distinct in habit, in floral morphology, and in gross morphological
appiearance, as to appear deserving of recognition as distinct species. Linking all these
variants together is one constant and readily observable organ, the seed, which in shape
and ornamentation is so distinct as to allow one to discriminate this taxon from all others
(except for the closely related species H. doleiformis). The variants which make up the
species H. pergrcimilala appear to form an intergrading complex and the subspecies as
here delimited merely represent those that are more clearly definable.

Comparison between different specimens is not always easy since the flowers and
fruit change considerably in shape and texture with development, while local conditions
and methods of preservation add to the morphological diversity. A number of variants
have not been collected in flower, or not retained in a condition where the flower structure
has been adequately preserved for critical examination, for this reason the descriptions
of the subspecies do not always permit close comparison. In particular the extent of
fusion between the flowers and the upper bract, the shape and position of the perianth
lobes, and the shape and texture of the mature fruitlets, are not known for each of the
taxa.

The imbrication of the perianth lobes is (basically) constant throughout a subspecies
and provides an easy way of distinguishing subsp. pergrainilata (the most common and
widespread of the subspecies) from //. halociwnioicles. In the western variant of sub-
species pergrainilata the abaxial lobe is large, rounded, and overlaps the lateral lobes,
whereas in H. halocnemokles the lower lobe is small and is within the laterals. The
imbrication of the lateral lobes, in relation to each other, is apparently consistent through-
out the species; the lateral florets of a triad have the lobe nearest to the central floret
overlapped by the outer one while in the central floret itself the imbrication can be of
either type.

Lectotypification: The lectotypification of A. halocnemokles var. pergramdatum is critical
since j. M. Black cited numerous collections of which four appear to have been used in
the production of the illustrations accompanying his description. The collection here
designated as lectotype represents the most widespread of the four subspecies recognised
in H. pergrainilata and it is also the one which best fits the original description. Of the
other syntypes, one (from Cootanoorina, leg. R. Helms) is here referred to H. pergrainilata
subsp. elongata, a second (from Lake Hart, leg. <7. Taylor) belongs to H. pinrijiora, a third
(from River Frome, leg. J. M. Black) is of H. pergrainilata subsp. divciricata, and a fourth
(from Port Alma, leg. L. Hassell) is of H. pergrainilata subsp. qiieenslatulica. The selection
of lectotype has not been made from those syntypes collected near Adelaide (Grange
and Birkenhead) since, as mentioned elsewhere, in this area subsp. pergrainilata differs
slightly from the bulk of the material included in this taxon.

Key to Subspecies

1. Flowers free from each other and from upper bract; lower perianth lobe obvious 2
1 *. Flowers i fused to each other and to upper bract; lower perianth lobe very small

or absent 3

2. Spikes undulate in outline; articles somewhat laterally compressed: lower lobe of
perianth prominent, usually overlapping lateral lobes .... a. subsp. pcrgranulata

2*. Spikes slender, terete, lower lobes of perianth shallow and scarcely overlapping
lateral lobes b. subsp. elongata

3. Plant sprawling or decumbent; spikes slender, erect (coastal plant)

c. subsp. queenslandica

3*. Plant erect; spikes spreading (inland plant) d. subsp. divaricata

42

Nuylsia Voi. 3(1) 1980

6a. SLibsp. pergraniilata Figs. 21, 22A-C, 42, 43.

Erect suhshrub ca. 0-5 m high. Ariicics obovoid, dull green or glaucous; lobes
short and rounded. Spikes terminal to branches or on lateral ascending branchlets,
3-4 mm diameter; articles barrel-shaped to shortly obovoid, either circular in cross section
or laterally compressed, bract-apices convex. Flowers free from each other and from
bracts, predominantly hermaphrodite, ascending; apex curved, lobes 3 with the large
semi-circular abaxial lobe outside (in Western Australia) or inside the lateral lobes.

Habitat: Coastal swamps, brackish swamps, and margins of salt lakes.

Distribution: Western New South Wales, Western Victoria, South Australia, south Western
Australia. Map 15.

New South Wales: Lake Victoria Sin.. 17 Mar. 1959, L. A. S. Johnson & E. f\ Constahie (NSW 48435);
Wentworth, T. M. Whaite 1809 (NSW).

Victoria: NW of Lake Albaculva, Sept. 1887, C. French (MEL); 16 mi SE of Swan Hill, C. W. E. Moore
.3613 (CANB); Nhill, A. C. Beaiiglehole 19975 (PERTH); Tooradin, Oct. 1921. H. B. Williamson (CANB).
South Australia: Wallaroo, B . ./. Blavlock 382 (AD); Redcliff, R. J. Chinnock 21 19 (AD); near Swan Reach,
./. B. Clelaml 37 (NSW); South Pearson Island, 6 .Ian. 1923, T. G. B. Osborn (NSW).

Western Australia: 24 mi N of Karonie. K. M. Allan 292 (AD); Lake Grace, S. L. Everist 9080 (BRl);
near Diindas Rock, B. R. Maslin 2469 (PERTH); 1 km S of Three Springs, R. A. Sajfrey 1568 (PERTH).
Northern Territory: "Near the MacDonnall ranges". 1889, Rev. Schwartz {MW 71544).

This is the most widespread of the different subspecies of H. pergraniilata. It can
usually be recognised by its dull green or glaucous appearance and by the uneven margin
of the spike; on drying it frequently takes on a dark grey colour.

There is considerable regional variation in the floral morphology and in the general
appearance of this subspecies. In Western Australia the lower perianth lobe is large
and obvious, both in the fresh and dried condition; in South Australia it is relatively
small (about one third the width of the perianth) and thin, its position being sometimes
within and sometimes outside the lateral lobes, while in the dry state it may not be immedi-
ately apparent and with age appears to become fused with the underlying perianth tissue.
Around Adelaide is found a variant in which the seed is ovate (rather than circular) with
the central region of either side tin-ribbed. This variant ranges eastwards into Victoria
and grades locally into the typical variant; it is also found in Yorke Peninsula. In New
South Wales is found a variant with slender branches and spikes; this is similar to a plant
found in Yorke Peninsula and in both cases looks different to the typical variant which
is found in the same general area.

The subspecies pergraniilata grades northwards in South Australia into both subsp.
elongata and subsp. Jivaricata, although the latter two subspecies appear sometimes to
retain their distinctive appearance even when growing in areas where the two occur. In
coastal New South Wales it appears to grade northwards into subsp. queenslandica.

Plants which are evidently hybrids between //. Iialoenemoiiles subsp. haloenenioides
and H. pergraniilata subsp. pergraniilata have been collected at various localities through-
out the overlapping ranges of these two taxa. These putative hybrid plants frequently
exhibit deformities and sterility of flowers — the ovules either not developing or the seed
never reaching maturity.

Plants which are probably hybrids between H. pergraniilata and Sclerostegia urbuscida
have been collected in Port Phillip Bay. For further comment see under the latter species.

On the north coast of Western Australia is found a variant in which the flowers are
adherent to each other and to the upper bract. It is an erect plant quite different in appear-
ance from subsp. queenslandica which grows in a similar habitat on the Queensland coast.
In the flow'ering state it is very similar to H. indica subsp. Iciostachya and in fact the two
may be found growing together on the coastal mudflats. Possibly for this reason it has
been very seldom collected. The two collections seen of this north-coast variant are
as follows;

We.stern Australia: Wyndham, A. S. George 14544 (PERTH); Sir Graham Moore Is., P. G. Wil.son 11268
bis. (PERTH),

Paul G. Wilson, Australian Salicornieae

43

A single collection from central Queensland (Rowen Downs, MEL 7061 1) has slender
branchlets, short terete spikes, and flowers fused to the upper bract; the orifice of the
perianth is deltoid in shape, quite different from that found in subsp. qucenskmdica. For
the present this plant and the collections made along the north coast of Western Australia,
can only be treated broadly as being members of the species II. pergrainilata.

The only collection of subsp. pergrunidata from the Northern Territory (MacDonnell
Ranges) comes from a locality well north of the otherwise recorded range of the subspecies.
The specimen is typical of the southern South Australian populations and for this reason
it seems likely that a confusion of labels has occurred.

In Western Australia the typical subspecies usually bears flow'ers in which the stamen
is vestigial and obviously not functional; they are also often cleistogamous. In southern
South Australia and Victoria the flowers are frequently hermaphrodite with apparently
fertile anthers.

6b. subsp. elongata P. G. Wilson, subsp. nov. Fig. 22C-E.

Spiccw erectae. graciles, cylindraceae, ad 6 cm longae, ca. 2 ■ 5 mm diam.. bracteis fere truncatis. Flores
libri. Periuiilluiim ad apicem exposum; lobi tre.s, subaequales, lobo abaxiali semiorbiculari, lobos laterales
leviter superposito.

Type: 7 mi SE of Rabbit Flat, 130 E, 20 I5'S, Northern Territory; salt pan; 26 May
1970, J. R. Maconochie 1032 (holo; PERTH, ex NT 27274, iso: CANB).

Herbaceous to somewhat w'oody. erect to decumbent short-lived perennial, to 25
cm high. Articles on branchlets short cylindrical to doleiform, 3-5 mm long, ca. 2-5 mm
diam., obtusely lobed, glaucous. Spikes terminal to branches or pedunculate on lateral
ascending branchlets, slender, cylindrical, to 6 cm long, equal to branchlets in diameter;
bracts similar to vegetative articles but shorter, slightly lobed, the lower margin deeply
concave over the triads. Flowers hermaphrodite (in some specimens only female seen),
free from each other and from bracts, at first ascending but eventually - vertical to spike
axis. Perianth: apex sub-truncate and exposed with 3 large almost equal lobes, the lower
(abaxial) lobe semi-orbicular and in bud slightly overlapping the lateral lobes.

Habitat: This subspecies appears to be restricted to the strongly saline areas around salt
lakes.

Distribution: Central Australia. Map 16.

1 Queensland: E shore of Lake Buclianan, 10 mi N of Bowie Stn., L. Adams 1 168 (BRI, CANB, NSW) —
specimen in flower only.

South Au.sfralia: Ciirdimiirka, A. C. Beauplehole 20885 (AD); Cootanoorina Creek, 5 May 1891, R. Helms
(AD).

Northern Territory: Napperby Salt Lake, C. Dunlop 2352 (PERTH); Erldunda Sin., T. S. Henshall 53,
81 (NT); Napperby Sin.. T. S. Henshall 101 I, 1021 (NT); 10 mi WSW ol Stuart Ras, G. f. Hill 221 (NSW);
Mt. Dare Stn., P. K. Latz 4787 (NT); Lake White, ./. R. Maconochie 934 (PERTH); 7 mi SE of Rabbit
Flat,/. R. Maconochie 1032 (PERTH).

This subspecies is, in the dried slate, similar in appearance to both //. halocnemoides
subsp. longispicata and H. fontinalis. From the former it may be distinguished by its
glaucous colour, by its circular seed which is concentrically ribbed and porcate all over,
and by the lobing of the perianth (large outer lower lobe as against a small inner lower
lobe). From II. fontiiuilis it is principally distinguished by its .seed.

At King Sound, on the north coast of We.stern Australia, is found a plant which,
in the dried state, is very similar in appearance to specimens of subsp. elongata from Central
Australia. At King Sound it grows in mud on the landward edge of mangrove swamps
and has a sprawling habit. It seems inappropriate at present to give this coastal plant a

44

Nuytsia Vol. 3 (1) 1980

distinct formal status, and yet from its habit and locality it seems incorrect to refer it to
the same subspecies as the Central Australian plant. The following is the only collection
seen of the coastal variant;

Western Aiisiralia: Derby, 20 Aug. 1975, V. Semeniitk 24 (PERTH).

At Lake Buchanan in north-east Queensland has been collected a flowering plant
which is similar to the Central Australian specimens of subsp. elongata; however, fruiting
material is required to permit positive identification.

6c. subsp. queenslandicii P. G. Wilson, subsp. nov.

Spicae erectae, angusle cylindraceae, 2-3 mm diam., bracteis undulatis. Flores exposi, ad bracteas
superus conjuncti vel partibus distalibus iiberis, ad vicinas liberi vel conjuncti. Perianthium lobi laterales
sinui: lobus abaxialis ut videtur absens.

Type: Townsville Common, Queensland, coastal mudflats, 24 Feb. 1977, P. G. Wilson
s.n., (holo: PERTH, iso: BRI, CANB, K, NSW).

Open straggly, often decumbent suhshruh to 25 cm high; branches slender. Articles
cylindrical to barrel-shaped, ca. 5 mm long on branchlets, dull to glaucous, apex very
shortly and bluntly lobed. Spikes narrowly cylindrical, 1-3 cm long, 2-3 mm diameter,
bract pairs with undulate margin. Flowers either totally fused to upper bract or free at
distal edge, either free from or fused to each other. Lower perianth lobe not observed;
lateral lobes sinuous.

Habitat: Coastal mud-flats.

Distribution: Queensland coast at least from Burketown east and south to Brisbane.

Queensland: Cooktown, S. T. Blake 21876 (BRI); Serpentine Ck., L. Diirrinitton 593 (BRI); Whyte Is.,
L. Durrington 1337 (BRI); Marmon, Mar. 1920, W. D. Francis (BRI); 48 km N of Burketown, T. J. Hall
590 (BRI); Port Alma, Oct. 1919, L. Hassell (AT). BRI); Townsville, W. Macnae 12.3 p.p. (BRI); Curtis
Is., 21 Apr. 1962, W. Macnae (BRI); Nudgee Beach, 6 July 1940, 1. S. Smith (BRI); Cribbe Is., Van Raven
9365 (BRI),

This subspecies appears to contain three variants but these, with further collecting,
may be shown to be representatives of a continuous dine running around the east and
north-east coast of Australia, The variants differ in the degree of fusion of the flowers
with eaeh other and with the upper bract; in the variant found near Townsville (which
includes the type) the flowers are free from each other and, at their distal margin, from the
upper bract (at least when young); in the southern variant (Brisbane to Rockhampton),
the flowers are totally fused to the upper bract (the epidermis being continuous over both
organs) and to each other in the distal half, A single collection from Burketown in the
Gulf of Carpentaria has spikes similar to the southern variant but with the flowers totally
fused together. Plants from all localities occupy a similar habitat (coastal mudflats or
mangrove swamps) and have a virtually identical vegetative appearance and habit; for
this reason it does not seem profitable to make formal taxonomic divisions within this
subspecies.

There is no indication that subsp. citieenslatulica grades into the inland subspecies
nor does it appear to hybridize with other Halosarcia species w'hich occur in the same
area and habitat, but it may form a clinal relationship w'ith the coastal variant of subsp
pergrainilatiim which occurs in southern New South Wales, a supposition that again can
only be confirmed or otherwise by further collecting. It may also grade westwards along
the north coast of Australia into the variant of subspecies pergranulata found on the north
coast of Western Australia; here the plants are erect, with the florets somewhat adnate
(but not fused) to each other and the upper bract.

In none of the collections examined have any fertile stamens been seen and, in fact,
a study of material which is at a stage, or in a condition, suited to investigation has in
all cases shown that the flowers bear a staminode which consists of a minute vestige about
0-1 mm long. In some flowers the stigmas become exserted but in others they remain

Paul G. Wilson, Australian Salicornieae

45

within the perianth and in either case seed is formed. It would appear that the plants
are parthenogenic and it is interesting that the variant of //. inilica subsp. indica which
also grows on coastal mudflats likewise has only female flowers. The adoption of an
apomictic mode of reproduction could in these cases, be related to the habitat which
is subject to tidal inundation; however, in other instances (e.g. in H. lepiodada) this mode
of reproduction is obviously not related to a tidal situation.

The precise morphology of the perianth in this subspecies is uncertain; no lower
(abaxial) lobe has been observed although it is possible that this lobe is present but dia-
phanous and that at an early stage it becomes fused with the underlying perianth tissue;
the lateral lobes are long and sinuous. Fresh material at an early stage of anthesis should
be studied to determine the situation.

The two subspecies, queenslandica and divaricata, are evidently closely related but
their distinct appearance suggests that, apart from other characters, they should be recog-
nised as separate taxa.

6d. subsp. divaricata P. G. Wilson, subsp. nov.

Spicae divaricatae, cylindraceae, 1-3 cm longae, ca. 3-5 mm diam., bracteis undulatis. F/o/'ci expos!,
ad vicinas et bractea.s superas conjunct!, apicem truncati. Perianthii \ob\ laierales late imbricati, grandes;
lobus abaxialis semi-circularis, membranaceus lobos laterales superpositus.

Type: Goyders Lagoon, 8 km S of Koonchera Waterhole, 26'’46'S, I39°3I 'E, South Aus-
tralia; sand dunes; 14 Aug. 1975, J. Z. Weber 4497 (holo: AD).

Dense erect shrub to 1 m high. Articles on branchlets cylindrical to obovoid, greyish
green, apex bluntly lobed. Spikes divaricate, usually on very short lateral branchlets,
cylindrical 1-3 cm long, ca. 3-5 mm diam., with up to 20 articles; bract pairs undulate,
i circular in cross-section. Flowers either female or hermaphrodite, exposed at apex,
fused to each other and to upper bract; apex truncate with the centre slightly raised.

Perianth: lateral lobes broadly imbricate, the outer one prominent, semi-circular, extending
length of apex, lower (abaxial) lobe membranous, semi-circular, either outside or inside
the lateral lobes. Seed as in subsp. pergraiuilata. Fruiting perianth remaining attached
to spike and together eventually breaking down to release seed.

Flahitat: Inland claypans; apparently not in strongly saline soils.

Distribution: South-western Queensland, w'estern New South Wales, north-western Victoria,
north-eastern South Australia. Map 16.

Queensland: Nockatunga, 5 Jan. 1935, A. C. Boyle (BRl); Mulligan River, Feb. 1904, H. Clarke (NSW);
S of Eulo, 31 July 1962, anon. (BRl); Birdsville-Betoota Rd,, 9 Aug. 1964, A. Rudd (NSW),

New South Wales: Delalah Downs, O’. M. Cunningham 516 (NSW); Fort Bourke, Oct. 1919, T. Wadded
(NSW).

Victoria: 17 km NW of Kerang, A. C. Beauglehule 57137 (PERTH).

South Australia: 24 km S of William Ck., A. C. Beauglehole 2081 (AtJ); Ml. Dare Stn., P, K. Latz (NT);
Oaken Hills, B . ./. Murray 503/2 (AD).

This subspecies is strictly an inland plant of eremaean situations. It may be recog-
nized by its short cylindrical spikes which are strongly spreading and are borne on short
lateral branchlets. The flowers are intimately fused to the upper bract and to other
members of the cymule; in fact the epidermis is continuous over both bract and flowers.
The lower (abaxial) perianth lobe is probably always present but being very delicate soon
becomes indistinguishable from the underlying perianth tissue into which it appears to
merge with the development of the fruit.

The subspecies divaricata grades to the south into the typical subspecies; plants
intermediate in location have a depression between the flowers and the bract; in plants
even further to the south this depression develops into an absence of cohesion between
the two organs. There is no suggestion of introgression of subsp. divaricata with subsp.
elongata even though the two subspecies are to some extent parapatric.

46

Nuytsia Vol. 3 (I) 1980

7. Halosarcia doleiformis P. G. Wilson, sp. nov. Fig. 22F-G.

Salicoriiia leiosiachya Benth., FI. Austral. 5: 204 (1870) p.pte., as to "Drummond, (Herb. F. Mueller)”,
not as to lectotype.

Fiiiiiciilosiis ad 0-5 m altus. Articiili glaucescentes, doleiformes, ca. 5 mm longi, in statu sicco medul-
losi, apice leviter lobato, margine ciliolato. Spicae terminales, anguste cylindraceae, attenuatae, ad 15
mm longae, laeves et medullosae; bracteae confertim imbricatae (segmentis undulatis). rotundatae, ad
vicinas coalescentes; facies abaxialis valde adscendens. Pmanihium versus apicem succulentum, margine
adaxiali ciliolato: lobos laterales superpositus. Spica fructifera grisea, laevis; bracteae siccae, medullosae,
confertim imbricatae. Iloribus occultis; perianthium medullosum; pericarpium tenue, molle, ad perianthium
adhaerens. Semen fere circularc, ca. I mm diam.; testa Crustacea atroporphyrea, cum costis conccntricis
prominentibus porcalis ornata.

Type: Murchison River, ca. 8 km E of Ml. Narryer, 26 30'S, II6‘'20'E. rounded shrub
0- 3-0-6 m high. 2 Sepl. 1970, P. G. Wilson 9914 (holo: PERTH, iso: CANB, K, NSW).

Low spreading rounded subshrub to 0-5 m high. Articles dull green, doleiform,
ca. 5 mm long, pithy when dry; apex slightly lobed; margin ciliolate. Spikes terminal
to branches, narrowly cylindrical and attenuate, to 15 mm long, grey, smooth, and pithy
when mature; bracts closely sheathing, the opposite members connate and forming an
undulate segment, margin ciliolate. Flowers hermaphrodite, in disjunct triads, steeply
ascending, coalescenl to each other laterally and frequently to upper bract, ape.x rounded,
somewhat exposed. Perianth succulent towards apex with its upper (adaxial) edge forming
a thin ciliolate border: lower (abaxial) lobe prominent, semi-orbicular, ciliolate, over-
lapping the large lateral lobes; anther ca. 1 mm long; ovary membranous, free from
perianth; stigmas slender. Fruitini> spike grey and smooth in outline, the bracts dry,
pithy, closely sheathing so as to obscure the fruitlets; perianth pithy; pericarp thin and
soft, adherent to the perianth. Seed flat, almost circular, ca. I mm diameter; testa
crustaceous, very dark reddish brown, concentrically ribbed all over, the ribs with porcate
sculpturing; embryo slightly curved; perisperm lateral. Seed released on decay of spike.

Distribution: South-western Western Australia from Shark Bay south to Northam and
south-east to the Norseman district. Map 3.

Western Australia: Lake Ninan, T. E. H. Aplin 731 (PERTH); Western Australia, J. Drummontl s.n. (K,
MEL. NSW — syntype of Salicoriiia ieiustachya)-, Dirk Hartog Is., A. S. George I 1609 (PERTH); 16 km
S ol Mt. .lackson, K. F. Keiiiiecilly 4741 (PERTH); 145 km E of Norseman, P. G. Wilson 101 16 (PERTH)-
93 km N of Esperance, P. G. Wilson 10129 (PERTH).

Halosarcia doleiformis is closely related to, and in some situations sympatric with,
H. pergranulata. It may be readily distinguished from the latter species by its barrel-
shaped articles which become pithy with age and retain their form, by its smooth attenuate
spikes, and by the ciliolate margin to the articles and bracts. In the mature fruitlets of
H. pergranulata the perianth either remains fleshy and eventually shrivels or it becomes
firmly chartaceous on the outside and retains its deltoid outline (from above), in either
case the fruitlets fall away from the spike axis. In H. doleiformis the perianth becomes
dry and pithy, similar to the bracts in texture, the whole spike being brittle and readily
fragmenting between the articles. The seeds of the Uvo species are virtually identical.

Specimens which are intermediate in form between H. doleiformis and H. pergranulata
have been collected; examples are as follows: Goomalling, T. E. H. Aplin 714; 8 km S
of Coorow, A. S. George 2310; 15 km E of Wagin, P. G. Wilson 8270 (all herb. PERTH).

The specific epithet refers to the barrel-shaped appearance of the dry articles.

The flower of H. doleiformis resembles the southern Western Australian variant of
H. pergranulata (i.e. in having a large rounded outer abaxial lobe), and therefore, con-
sidering the virtual identity of the seeds of the two species, it seems likely that the two
plants have arisen in this region from a common ancestor in Western Australia. As is
noted elsewhere in the text, the form taken by the flower of H. pergranulata in other areas
of Australia differs considerably from that found in H. doleiformis.

Paul G. Wilson, Australian Salicornieae

47

8. Halosarcia Icpidosperma P. G. Wilson, sp. nov. Fig. 23D F. 44, 45.

Friiliciildsii.K ad I m altus. ramis crcctis. Articnli obovoidei vel doliit'ormes vel cylindracei 5-10 mm
longi leviter glauci. Spicae graciles, aequaliter cylindraceac, ad .50 mm longae 3-5-5 mm diam.; bracteae
truncatae. Flores fere liberl leviter adscendentes, apice truncato. Peruinlliiimi succulentum; lobus medius
abaxialis prominens, scmi-circularis. integer, lobos laterales superpositus. Perianthitim in fructo succulen-
tum. demum spongiosum; periearpium membranaceum. ad perianthitim connatum. Fniciiciili demum
secedentes semine expose. Semen sub-orbiculare, ca. 1-5 mm longum, album: testa in costis prominentibus
squamiformibus ornata; embryo semi-circularis.

Type: Western Australia, Baandee tovvnsite W of Merredin; light brown soil on edge of
salt lake; 21 June 1969. R. A. Saffrey 621b (holo: PERTH, iso: CANB).

Subshrub to I m high; branches slender, erect. Articles obovoid to barrel-shaped
or cylindrical, 5-10 mm long, yellowish green to dull green, slightly glaucous. Spikes
slender, terminal to branches or sub-sessile, evenly cylindrical to 50 mm long. 3-5-5 mm
diam., ape.x T: rounded; bracts truncate, the opposite pairs united; cymules disjunct,
prominently exposed at apex. Flowers almost free from each other, slightly ascending
(upper and lower sides ± parallel), apex truncate and broadly oblong (central flower),
or triangular (lateral flowers); perianth succulent. 3-lohed in centre of apex, the lower
(abaxial) lobe large semi-circular, and overlapping the pair of lateral lobes; ovary mem-
branous. Friiiileis not or scarcely exceeding bracts; perianth succulent, eventually spongy
and becoming shrivelled on drying, or firm and chartaceous on outside: pericarp mem-
branous, fused with perianth which eventually falls from the spike with the seed which
is exposed at the torn perianth base. Seed sub-orbicular, ca. 1 -5 mm long, white or pale
fawn when dry; outer testa raised on the margin to form 5-7 concentric scale-like ribs
with transverse corrugations (sides of seed smooth); embryo semi-circular; perisperm
lateral (somewhat enclosed).

Habitat: Growing in moderately or only slightly saline soil.

Distribution: South-western Western Australia and south-western South Australia. Map 1.

Sunrh Australia: Stenhouse Bay, Yorke Peninsula. D. Svmon 9641 B (ADW); Lake Pillie. 10 km S of Pt.
Lincoln, P. C. Wilson 2678 (AD).

Western Australia: 8 km W of Tammin, A'. M. Allan I (PERTH); 2 mi W of Waeel, T. E. H. Apliii 670
(PERTH); 18 mi S of Wagin, E M. Bennett 3070 (PERTH); Point Pedder. Hardy Inlet, Congdon & McComb
74032b (PERTH); Oldfield River. Hj. Eiehler 20240 (PERTH); Stokes Inlet. Hi. Eichler 20319 (PERTH);
Lake Wagin, MBs Cronin iMEL): 3 mi E of Newdegate, A. S. OVoree 7338 ( PERTH); 4 mi N of Pt. Malcolm,
R. Hnatiuk 761200 (PERTH); Bunswood Is., Swan R.. 14 Apr. 1900. A. Morrison (BRI, CANB, PERTH);
N of Stirling Ra.. Oct. 1867, F. Mueller (MEL 71463); ibid.. 1869. F. Mueller (K); I mi N of Ongerup,
K. Newhe\ 3067 (PERTH); between Miling and Pithara, P. 0. Wil.son 9979 (PERTH).

Halosarcia Icpidosperma may be readily recognised by its long evenly cylindrical
spikes and by its seeds which are white with smooth concentric scale-like ribs over the
embryo. In the held it is usually of a more yellowish green (tending towards chlorotic
in appearance) than other Halosarcia species growing in the same area. In herbaria
it has been confused with H. pierygosperma which also has white seeds; the latter species
may be readily distinguished by its short spikes with undulate outline and by the seed in
which the ribs are rough and cover the entire surface of the testa (Fig. 46). The seed
of H. Icpidosperma is almost identical to that of Tegicornia uniflora but in other respects
the two species have little in common.

Halosarcia Icpidosperma is one of the most common samphire species in the south-
west of Western Australia. It is an early invader of the newly created saline areas which
follow the clearing of woodland and is frequently present along roadside drainage channels
in salt-aflected country. Only two collections made prior to 1900 have been seen (“Margin
of salt lakes north of Stirling Range, Oct. 1867, F. Mueller", which was included under
Salicornia hidens by Bentham, and Lake Wagin. leg. Miss Cronin). This suggests that
the plant was far less common in the nineteenth century. The localities of the few collec-
tions made in South Australia are such as to suggest that it is a recent introduction to that
State.

48

Nuytsia Vol. 3 (1) 1980

This species exhibits no apparent regional variation and does not appear to hybridise
with any other member of the genus.

The specific epithet is derived from the Greek words lepidos (scale) and sperma (seed);
referring to the scale-like ribs on the seed.

9. Halosarcia pterygosperma (J. M. Black) P. G. Wilson, comb, et stat. nov.

Arthrocnemum hahcnemoides var. plerygospermnm J. M. Black, Trans. Roy. Soc. S. Austral. 60: 166 (1936).

T\pe: Mount Victor Station, 24 May 1936, C. M. Eardlev (holo: AD 97614361, iso:
MEL 70586).

Spreading subshrub to 0 ■ 5 m high, with slender divaricate branches. Articles narrowly
to broadly obovoid, ca. 5 mm long, dull green or somewhat glaucous. Spikes terminal
to main and lateral branches, cylindrical to ellipsoidal, to 30 mm long and 5 mm diam.,
with up to 15 articles; bract pairs united or apparently free, di deeply concave over cymules;
opposite cymules contiguous. Flowers free, or adherent to each other and to bracts
at least towards axis, arranged vertical to axis, deltoid in shape from above with both
adaxial and abaxial sides horizontal; apex truncate, quadrate to transversely oblong,

1 -5 mm long and wide, 1-2 mm high; perianth succulent; lobes 3 in centre of apex, entire
or denticulate, the lower lobe semi-circular and overlapping the pair of curved lateral
lobes: anther ca. I mm long. Fruiting spikelets with thin shrivelled bracts forming septa-
like plates between the disjunct whorls of fruitlets. Fruitlets deltoid from above, to 2-5
mm long, considerably exceeding bracts and readily shed when mature (the seed totally
enclosed), perianth chartaceous, glossy, hollow (somewhat mucilaginous around seed
when immature); pericarp membranous or apparently absent. Seed thick, ovoid, ca.
1-5 mm long, pale fawn to cream coloured; outer testa raised into corrugated scale-like
ribs over whole of seed (or the cells of the ribs J; separate and forming curved to uncinate
hairs); embryo slightly curved; perisperm lateral.

This species is distinctive because of its fruiting spikes in which the fruitlets are arranged
in prominent whorls considerably exceeding the bracts. The perianth of the fruitlet is
papery and glossy, which, with the white ovoid seeds with prominent uneven ribs, clearly
distinguishes H. pterygosperma from other species.

The distinction between the two subspecies noted below is not sharp for in collections
of subsp. pterygosperma which come from areas towards the western end of its range the
articles are ciliolate and the ornamentation of the testa is somewhat intermediate between
the typical variants of the two subspecies.

Key to subspecies

1. Vegetative articles with rounded to obtuse lobes; testa raised into scale-like ribs

^ : : , . , . a. subsp. pterygosperma

1*. Vegetative articles with acuminate or apiculate lobes; testa covered with rows of
uncinate hairs which may be united towards their base b. subsp. denticulata

9a. subsp. pterygosperma. Fig. 46, 47.

Articles with rounded to obtuse lobes, margins entire to ciliolate. Spikes ± cylin-
drical; fertile articles undulate. Perianth: lower lobe entire. Seed: testa raised into
scale-like ribs.

Habitat: Margin of salt lakes.

Distribution: Western New South Wales and Victoria, central and south-western South
Australia, south-central and south-eastern Western Australia (from Shark Bay eastwards)
Map 17. J J-

Paul G, Wilson, Australian Salicornieae

49

New South Wales: Donsandel (Broken Hill), 10 Oct. 1974, L. Richley (NSW).

Victoria: 30 mi S of Mildura, J. Ciilliinore 9 (AD, BRl, NSW, PERTH); 8 mi SSE of Ouyen, /. CuHimore
49 (AD, NSW); Lake Wahpool, N. Macfarlane 852 (PERTH); Cowongie, 29 Aug. 1955, J. H. fP/Y/A (MEL).

South Australia: Fowlers Bay, Bay 111, 29 .Ian. 1802, R. Brown (BM); Koonamore, Hj. Eichler 12433 (AD);
Yalata Swamp, A. E. Orchard 3214 (AD).

Western Australia: Quelquelling Wells, T. E, H. Apliii 708, 2217 (PERTH); 5 km NE ol Norseman, A. C.
Beauglehole 49380 (PERTH); Nanga Stn., A. Devitt 71 (PERTH); Dirk Hartog Is., A. S. George 11454a
(PERTH); Lake Anneen, 28 Mar. 1976, R. Hnatiuk (PERTH); Mundrabilla Stn., C. V. Malcolm 652
(PERTH); Nullarbor H.S., A. E. Orc/wrt/ (PERTH); S end of Hamelin Pool, P. G. Wilson 8229 (PERTH);
305 km N of Geraldton, P. G. Wilson 8331 (PERTH); 21 km N of Mt. Ragged, P. G. Wilson lOlOO (PERTH).

9b. subsp. denticulata P, G. Wilson, subsp. nov. Fig. 24A-E.

Lobis arlicularum acuminatis vel apicuiatis, margine ciliolato. Spicae ellipsoideae; bracteae ad
apicem liberae vel fere liberae, facie exteriori transverse triangulari. Perianthium lobo medio abaxiali
denticulate. Testa chartacea, pilis uncinatis seriatis ornatis.

Type: Western Australia, 6 mi SW of Yardie Creek Fl.S., North-West Cape; shrub 20
cm, 1 bright green; open saline Hat, 6 Sept. 1970, A. S. George 1031 1 (holo: PERTH).

Articles with acuminate or apiculate lobes, margins ciliolate. Spikes ellipsoidal with
rounded apices; bracts either united and markedly concave above triads or ! distinct
distally with the outer surface transversely triangular in outline. Perianth: lower lobe
denticulate. Seed: testa covered with rows of uncinate hairs which are sometimes united
towards their base.

Habitat: Coastal (or near coastal), in tidal flats, along tidal creeks, and margins of salt
lakes.

Distribution: Lake Macleod north-east to the Eighty Mile Beach, Western Australia.
Map 17.

Western Australia: 9 mi N of Yardie Ck., K. Allan 454 (PERTH); Wooroo Ck., Eighty Mile Beach,
N. T. Burhidge 1307 (PERTH); Lake Macleod, A. S. George 10196 (PERTH); Cape Keraudren, A. S.
George 14823 (PERTH); ibid., 26 .lune 1974, R. E. Johnstone (PERTH).

10. Halosarcia lylei (Ewart et White) P. G. Wilson, comb. nov. Fig. 23A-C, 48, 49.

Salicornia lylei Ewart et White, J. Roy. Soc. N.S. Wales 42: 195, t.34 (1909). Arthrocnernum lylei
(Ewart et While) J. M. Black, Trans, Roy. Soc. S. Austral. 43:359, t.34 (1919).

Lectotype: Cowcowing, near salt lakes. Sept. 1904, M. Koch 1051 (holo: MEL, '?iso:
NSW), cf. J. M. Black, op cit. 360.

Erect woody perennial to 1 m high, frequently with several fastigiate stems arising
from the rootstock. Branchlels very slender, evenly and narrowly cylindrical; articles
ca. 3 mm long, not swollen, dull green, margin truncate, eciliate, leaf-lobes apiculate.
Spikes terminal, slender, to 20 mm long, 3 ■ 5-4- 5 mm diameter. Bracts short, the opposite
pairs united, lobes deltoid, freely exposing the flowers. Cymules contiguous. Flowers
perpendicular to spike axis, free from each other and from bracts; apex quadrate, truncate.
Perianth fleshy, orifice triangular in centre of apex, abaxial lobe overlapped by the pair
of lateral lobes. Anther ca. I mm long. Ovary at first membranous and free from
perianth; style very short and thickened at base; stigmas narrowly triangular ca. 0-5 mm
long. Fritillels extending beyond bract, 1-5-2 mm long, firm and pithy (not indurated);
pericarp convex and protruding slightly from perianth giving it a mammillate appearance,
crustaceous at apex where fused to perianth, membranous below; style base persistent,
hard. Seed broadly elliptical, reddish brown, 1-1-5 mm long; testa crustaceous, con-
centrically granular over embryo, otherwise T smooth; embryo curved; perisperm lateral.
Fruitlets shed with enclosed pericarp and seed.

Habitat: Usually around salt lakes but also in poorly drained inland localities.
Distribution: Western Victoria, south-western South Australia, south-western Western
Australia. Map 12.

91670 — ( 4 )

50

Nuytsia Vol. 3(1) 1980

Victoria: Landrook Lake. A. C. Bcaitglehole 19814 (PERTH); Tyrrell Ck., 5 Sept. \9(i5,J. H. W-V/Z/.v (MEL);
8 mi W ol' Nowingi, 1936, \V. J. Zimmer (MEL).

Soiit/i Australia: S of Lake Bring. D. E. Svmuii 3391 (ADW, CANB, K): near Maralinga, 5. Warne 20103
(AD).

Western Australia: 2 mi W of Waeei, T. E. H. Aplin 665 (PERTH); between Dundas Hills and Lake Lefroy,
J. D. Balt (MEL); Lake Lefroy, 7 Nov. 1891, /?. Helms (AD. MEL, NSW): Cowcowing, July 1904, AT.
Koch 1051 (NSW, svnlvpe); ibid,, Aug. 1904, AT. Koch 1051 (NSW. syntype); ibid., Dec. 1904, AT. Koch
1051 (K. syntype); Lake Biddy. 22 Sept. 1971, C. V. Malcolm (PERTH).

Halosarcia ly/ei may be readily identified because of its slender branches, which are
similar to the branches of Casuarina: in addition, the mammilliform appearance of the
fruitlets clearly distinguishes it from all other species. The fruitlets are similar in structure
to those of H. luiloaiemoulcs and, as in that species, the seed is shed with the perianth;
however, in H. lylei the seed is almost totally enclosed by that structure since the fruitlets
are only slightly embedded in the spike axis, whereas in H. halocnenmides the prominent
interlloral radial septa of the axis are fused to the proximal portion of the perianth, and an
abscission zone forms well above the base of the flower causing the seed to be exposed
on the release of the fruitlets.

The plant found in South Australia has thicker inflorescences than does material
from Victoria and Western Australia, and in South Australia the seed is 1-5 mm long
whereas in Western Australia it is only I mm.

11. Halosarcia leptoclada P. G. Wilson, sp. nov.

Iriiticulus parvu.s, diffusus ad 30 cm altus. Rami graciles; articuli icres. doliiforme.s vel anguste
obovoidei. apice breviter et obscure lobalo. integro, Spicae anguste ellip.soideae, ad 10 mm longae;
bracteae conlertim imbricatae. - truncatae. Flores occulti, initio ad vicinas coalescentum, basim versus
membranaceum: apice ± truncatus. lobis lateralibus prominentibus, lobo medio abaxiali interno; stamen
sterile vel ahsens; ovarium membranaceum. Fructiculi libri. .seminc inclusum; perianthum crustaceum,
extra muncatum, apice truncate late ovato; pericarpium tenue, basim versus absens. Semen ovatum,
ca 0-8 mm longum; testa membranacea, straminea, laevis.

Type: Western Australia: Mt. Sandiman Sin., north-east of Carnarvon; on saline stony
uplands, Aug. 1969, D. G. Wilcox 53 (holo: PERTH).

Small divaricately branched delicate suhshruh ca. 30 cm high. Brunches slender;
articles terete to doleiform or obovoid, 5-8 mm long, dull to glaucous; apex with small’
rounded, and entire lobes. Spikes terminal to main and lateral branches, narrowly
ellipsoidal, of 4-10 articles, to 10 mm long; bracts closely imbricate, the opposite pair
united with i horizontal margin. Flowers in ; contiguous triads, obscured by bracts,
abaxial surface ascending, at first adherent laterally to neighbouring florets, to upper
bract (except towards apex), and in proximal third to lower bract, ca. I mm long.’ Perianth
succulent at apex, thin towaids base; apex ^ truncate with a pair of prominent lateral
lobes and a small inner abaxial lobe. Stamen absent, or present and sterile. Ovary
membranous. Fruiting spike: bracts shrivelling on drying and eventually falling as plate-
like rings from the slender spike axis. Fruitlets free from each other and from bracts
standing vertical to spike axis, broadly ovoid in lateral view, ca. I -5 mm long, completely
enclosing the seed, perianth forming a thin crustaceous inner layer which is smooth within
but areolate-muricate on outside, apex dr truncate, broad-oval in outline and slightly
conical in centre due to projecting lateral lobes; pericarp thin, membranous or absent
towards the base. Seed ovate, ca. 0-8 mm long; testa membranous, very pale fawn
coloured, smooth or with very fine longitudinal lines over embryo; embryo somewhat
curved; perisperm lateral, prominent.

The epithet leptoclada is derived from the two Greek words deptos' slender and
‘dados' branch; it refers to the slender branches.

The species consists of two variants which are morphologically very distinct and
geographically disjunct. Due to their having almost identical fruit I am treating them as
subspecies since I consider it probable that further collecting will indicate that thev are
cimally related to each other. ^

Paul G. Wilson, Australian Salicornieae

51

Key to subspecies

Articles cylindrical to barrel-shaped; perianth (in flower) dorsiventrally flattened

towards apex I la. subsp. leptoclada

Articles narrowly obovoid; perianth (in flower) rounded to truncate at apex

1 lb. subsp. inclusa

11a. subsp. leptoclada. Fig. 25D E.

Branclilels slender and terete, or the articles doleiform. Flowers with perianth lobes
thin and forming a beak-like ascending conical apex; stamen sterile, the anther ca. 0-6
mm long; stigmas exserted, slender. Fruitlets with truncate broadly elliptic apex, the
perianth lobes projecting in centre.

Habitat: Stony areas, frequently saline.

Distribution: Gascoyne River area of Western Australia. Map 18.

Western Australia: Wanna, J. S. Beard 6064 (NSW, PERTH): Mt. Vernon, A. Mitchell 221 (PERTH);
Jimba Jimba Sin., D. G. Wilcox 10 (PERTH); Dalgety Downs Stn., D. G. Wilcox 137 (PERTH).

11b. subsp. inclusa P. G. Wilson, subsp. nov. Fig. 25F-], 60.

/)/•?«■»/;■ obovoidei vel anguste obovoidei; perianthium (in (tore) ad apicem rotundatum vel truncatum.
Type: 3 km E of Meckering on Mortlock R. flats. Western Australia; sandy saline soil;
open subshrub 20 cm high, articles dull green; 4 Nov. 1977; P. G. Wilson 11692 (holo:
PERTH, iso: CANB, K, MEL).

Branchlets slender; articles narrowly obovoid. Perianth ± truncate at apex in flower,
the lateral lobes eventually prominent in fruit as a vertical beak. Style frequently remaining
within the perianth.

Habitat: Sandy or loamy saline areas.

Distribution: South-western Western Australia. Map 18.

Western Australia: 34 km N ot Weslonia, K. M. Alia/? (PERTH); 5 mi W of Pingaring, A. S. George 9361
(PERTH); One Mile Rocks Reserve, A. S. George 10505 (PERTH); 2 km N of Mt. Stirling, B. R. Maslin
182 (PERTH); I mi S of King Rocks, K. Newhey 3244 (PERTH); Hines Hill, R. A. Saffrey 619 (PERTH);
2 mi W of Ravenswood, A. S. Weston 7404 (PERTH); 4 km S of Three Springs, f. & J. Whaite 4227
(PERTH); 25 km S of Perenjori, P. G. Wilson 8253 (PERTH),

The subspecific epithet refers to the fact that the stigmas frequently remain enclosed
within the perianth.

The species Halosarcia leptoclada is peculiar in having prominent lateral perianth
lobes which in fruit protrude as a short beak (in the flower, at least in subsp. inclusa,
they are flush with the perianth apex). In neither subspecies have fertile anthers been
observed although in subsp. leptoclada a well-formed (but sterile) anther ca. 0-6 mm long
is present. The staminode in subsp. inclusa is minute (ca. 0-2 mm long) and has no recog-
nisable anlherode; in this subspecies the stigma does not always emerge, in fact it normally
remains folded within the perianth. This is the situation as it has been observed in the
field in a population near Meckering and in herbarium material from near Lake King.
It is likely that in other localities the plant is normally hermaphrodite since a collection
from near Pingaring (A. S. George 9361) bears flowers with a fertile stamen and ovary; it
is more robust than the other collections of subsp. inclusa and the spikes frequently become
intercalary. From the material available it is not possible to say whether it is a hybrid,
or whether it should be recognised as a distinct ta.xon.

A collection made by Cecil Andrews at Canning Plains (near Perth) in October 1902
has the general appearance of H. leptoclada subsp. inclusa, but the material (which is
in bud) has been badly preserved and it is not possible to be certain of its identity; the
flowers have fertile anthers. Duplicates of this collection are present in the herbaria
BM, K and PERTH.

52

Nuylsia Vol, 3 (I) 1980

Halosarcia leptoclada siibsp. inclusa x H. pruinosa?

A plant which is intermediate in appearance between H. leptoclada ssp. inclusa and
H. pruinosa has been collected near Three Springs and at Ajana, Western Australia. A
description of the putative hybrid follows:

Small much-branched shrub to 50 cm high. Branchlets thin. Vegetative articles doleiform
to obovoid, 3-8 mm long, dull to glaucous, with shallow broadly rounded lobes; margin
broad and scarious, entire. Spikes terminal to branchlets, short, of 3-6 articles, the
bracts similar to but shorter than the vegetative articles, circular in cross-section and
somewhat swollen to give the spike an uneven appearance, ca. 3-5 mm diam. Flowers
obscured by bracts (the triads contiguous), free from each other and from bracts; abaxial
side steeply ascending; perianth beaker-shaped, lower half with fleshy lateral walls, upper
(distal) part membranous and somewhat dorsiventrally compressed, apex wide, open
and truncate, in bud consisting of a pair of lateral equitant lobes which are not apparent
at anthesis. Fruiting spike: bracts thin and shrivelled; perianth i horizontal with ascend-
ing strongly dorsiventrally flattened tongue-like apex, walls thin and soft, deeply honey-
combed outside, smooth and glossy within; pericarp membranous, absent tow'ards base.
Seed ovoid, ca. 1 mm long; testa thin, very pale brown, smooth; embryo slightly curved;
perisperm lateral.

Distributiun: Ajana and Three Springs. S.E. of Ajana, 6 Aug. 1976, R. Hnatiuk s.n. (PERTH); 5 km
S of Three Springs, by salt lake, 2 Oct. 1976, T. & J. Whaite 4225 and 4226 (PERTH).

12. Halosarcia flabelliformis P. G. Wilson, sp. nov. Fig. 8, 24F-H, 50, 51.

Fruiiculosiis ad 20 cm altus. Articiili anguste obovoidei vel doleiformi. ca. 5 x 2 5 mm, margine
fere truncalo inlegro. Spicae c> Imdraceae vel aliquantum attenuatae, 20-40 inm longae, ca. 5 mm diam.,
in ambilo laevi, apicc obtuso vel rotundato. Bracteae oppositae. librae, flabelliformes, facie exteriori
Irapezoidea. Flores ad axem spicae verticales, ad vicinas et bracteas liberi, apice truncattt, trapezoidea
Permiilhiiim succulentum. parietibus lateralibus et apice crasso, pariele adaxiali tenui: lobis laterali duo!
niagni; lobus abaxialis minulus, inlernus. Ovarium lenue. Spica fructifera demum in parte discreta
secedens; perianthium in dimidiis translucidis secedens; pericarpium membranaceum, librum. Semen
applanatum. late ellipticum ca. 1 -8 mm longum; testa tenuis, transluceda, super radiculam tuberculata
aliter laevis.

Type: Just behind Webb Beach (34 27'S, 138 16 E), South Australia; 30 Jan. 1977; R.
./. Cbinnock 3368 (holo: AD, iso: K, MEL, NSW, NY, PERTH).

Woody perennial to 20 cm high. Branches ascending: articles dull to glossy green,
narrowly obovoid to barrel-shaped, ca. 5 x 2 • 5 mm, margin almost truncate, entire. Spikes
terminal to main and lateral branchlets (not sessile) cylindrical or somewhat attenuate,
20-40 mm long, ca. 5 mm diameter, outline smooth, apex obtuse to rounded. Opposite
bracts free from each other, Uabelliform (from above), trapezoid on outer surface, not
obscuring the flowers. Flowers vertical to spike axis, free from each other and from tipper
and lower bracts; perianth succulent, tran.slucent; sides and apex thick; adaxial surface
very thin in medial position, apex truncate, trapezoid in shape, with two thick lateral
lobes which occupy most of the surface and a very small triangular inner abaxial medial
lobe, anther I mm long; ovary thin and weak, with a pair of short weak stigmas. Fruiting
spike eventually disarticulating into separate bracts, perianth halves, and pericarps w'ith
enclosed seed; perianth translucent, pericarp membranous. Seed flat, broadly elliptical
ca. 1 8 mm long, testa thin, translucent, with several rows of small tubercles over the
embryo, otherwise smooth.

Habitat: Coastal; Samphire association, forming a band below the Arthrocnemum-
Salicornia zone on wet blue-grey fine silty muds” (fide R. Chinnock in sched )■ “Centre
of gypsum swamp" (fide T.G.B. Osborn in sched.). ’ ’

Distribution: Only known from the coast north of Adelaide, and from Flinders Is off
the west coast of Eyre Peninsula, South Australia. Map 11.

Smnh Australia: Port Parham, R. J. Chinnock 2984, 3363, 3364, 3365, 3366 (AD)- Webb Beach R I Chin
nock 3371 (AD): Port Prime, R. J. Chinnock 3375 (AD); St, Kilda swampC 23 Mar 1923 / R rfjZu

SJSIT “£1’,AD," "■ S'- 'S'”" II j N tr Adilaidel:

Paul G. Wilson, Australian Salicornieae

53

Figure 8. Halosanki f/cihel/iformis: in samphire flat at Webb Bcaeh, South Australia. Photo by R. .1.
Chinnock.

This species is interesting in that it is intermediate in morphology between the genera
Halosarcia and Tccticornia. It could, in fact, equally well be placed in either genus.
The characters which suggest its afiinities to Tecticoniia are :( I ) its free flabelliform bracts;
(2) a perianth which divides readily into two lateral halves; (3) the membranous pericarp;
and (4) a similar seed to that found in T. austrulasica (Fig. 64, 65). None of these features
is restricted to the genus Tecticoniia, and other characters, such as the perennial habit
and the presence of an abaxial perianth lobe, are more indicative of a relationship with
the genus Halosarcia.

The specihc epithet refers to the fan-shaped appearance of the bracts.

13. Halosarcia peltata P. G. Wilson, sp. nov. Fig. 26. 52.

Fniticit/iis acl I m altus. sordido viridi. obovoidei, 2-5 mm longi, leviter lobati, plerumque

apiculati, marginem integri, granulares. Spicae late cylindraceae, 5-10 mm longae, 5 mm diam.. apicem
rotundatae; bracteae discretae, nec imbricatae, face! exteriore semicircular!, margine late scarioso. Flores
ad axem spicae verticales, ad vieinas et bracteam superam perl'ccte conjuncti. Periaiilliiiim succulentum;
lobi laterales grandes ab apice truncato protrudentes; lobus medius abaxialis parvus, internus. PerUmthiiim
in fructo succulentum demum medullosum, lobi.s lateralibus prominente protrudentibus; pericarpium
tenue medullosum, latcraliter compressum. ad periantliium conjunctum. Semen ovatum. ca. 1-3 mm
longum; testa tenuis, pallido brunnea, laevis. Spicci fnicrif'era demum a ramo secedens.

Type: 5-5 km S of Morawa, Western Australia; growing on salt pan; plant 0-5 m high,
dull green; 4 Sept. 1970; P. G. Wilson 9969 (holo; PERTFI, iso; CANS, K, MEL. NSW).

Much branched siihslinih to 1 m high. Arlicles dull pale green, papillose, obovoid,
3-5 mm long; apex somewhat lobed and frequently apiculate; margin granular (especially
when dry). Spikes terminal to short branchlets, broadly cylindrical and of up to 9 articles,
5-10 mm long, 5 mm diam. (when fresh), apex rounded; bracts free (or almost free) from

54

Nuytsia Vol. 3 (1) 1980

each other, not imbricate, sub-orbicular on surface, with broad scarious entire margin;
cymules well separated from each other. Flowers hermaphrodite, compact, aligned at
right angles to spike axis, fused along entire length to each other and to lateral and upper
bracts (but distinguishable by slight indentation in surface of spike), and along proximal
half to lower bract. Perianth succulent; apex truncate; lobes 3, lateral lobes large and
vertical, protruding from spike, overlapping the small inner hemispherical medial abaxial
lobe. Anther ca. 2-5 mm long. Ovary membranous; style arms delicate. Fruiting
perianth fleshy (pithy crustaccous when dry) with prominently exserted vertical (lateral)
lobes; pericarp thin, pithy, laterally compressed, fused to and scarcely distinguishable
from perianth. Seed ovate, ca. 1-3 mm long; testa thin, pale browm, smooth; embryo
curved: perisperm lateral. Release of seed by separation and eventual decay of perianths
in the spike which breaks away from the branch.

Habitat: Margin of salt lakes.

Distribution: Western Australia, from Lake Macleod south to Northam and east to
Kalgoorlie. Map 11.

Western Australia: Lake Annean. Aug. 1971, H. G. Baker (PERTH); S of Hamelin Pool. E. M. Bennett
3319 (PERTH); Nanga Stn., A. Devitt 20 (PERTH); Lake Macleod. A. S. George 10193 (PERTH); Dirk
Hartog Is.,/(.5. George 1 1430 (PERTH); Comet Vale,./. T.Jntson 195 (PERTH); .libberding, M. Koch 1373
(MEL); 4 km ENE Yelma H.S., 12 Sept. 1970,7. Loirn’ (PERTH); 16 km S of Widgiemooltha. R. A. Saffrey
1576 (PERTH): Mongers Lake, T. &J. H^/iatVe 4 1 29 (PERTH); 2.1 mi S of Three Springs, T. &J. Whaile 4221
(PERTH); Waeel, P. G. Wilson 6402 (PERTH); Lake Raeside, P. G. Wilson 7558 (PERTH).

The specific epithet refers to the peltate appearance of the bracts.

In H. peltata the perianths are intimately fused with the upper and lateral bracts,
and in their proximal half with the lower bracts. This condition continues to the fruiting
stage; when mature, the spikes eventually fall entire from the branches. The lateral
lobes of the perianth prominently protrude from the mature spike like a duck’s beak
(vertically positioned); this gives it a characteristic warty appearance.

The flowers and fruit of this species resemble those found in H. leptoclada to which
it is probably closely related.

14. Halosarcia pruinosa (Paulsen) P. G. Wilson, comb. nov. Fig. 27D-F, 53.
Arthrocnemum pruinosion Paulsen. Dansk Bot. Ark. 2 (8); 63, Fig. 25 and pl.6f.3 (1918).

Type: Carnarvon, 31 Oct. 1914, C. H. Ostenfeld 349 (iso: K).

Much branched suh-shruh to I m high. Articles obovoid, 3-8 mm long, often greyish
green; lobes rounded, entire. Spikes terminal to branches or to short lateral branchlets
(rarely sessile), narrowly cylindrical, 1 •5-3-5 (10) cm long, 3-5 mm diameter, occasionally
continuing growth vegetatively. Bracts shortly barrel-shaped, closely overlapping,
margin slightly undulate, entire. Flowers fertile or with either the stamen or ovary sterile,
coalescent to each other and fused to upper bract in proximal half, entirely concealed by
bracts; adaxial side rh horizontal, abaxial side steeply ascending, extreme apex dorsi-
ventrally compressed, acute. Perianth fleshy but rather thin with a pair of large con-
duplicate lateral lobes and a smaller oblong to deltoid inner abaxial lobe. Ovary thin,
free from perianth; style slender. Fruiting bracts spongy, remaining shortly cylindrical
or cup-shaped and imbricate. Fruitlets totally enclosed within the bracts, eventually
free from each other and from bracts; perianth brittle, areolate, muricate on the outside
but with a firm crustaccous inner layer; pericarp free from perianth, thinly crustaccous
at apex, membranous or absent below. Seed ovoid, ca. I mm long; testa membranous
pale brown, faintly concentrically granular over embryo, otherwise smooth; embryo
slightly curved; perisperm prominent and forming a lateral bulge. Bracts and fruitlets
eventually separating from each other and from the slender axis; perianth splitting in
medial sagittal plane to release seed.

Habitat: Moderately saline situations both coastal and inland.

Paul G. Wilson, Australian Saliconiieae

55

Distribution: Western Victoria, South Australia, Western Australia south of 24° latitude,
south-western portion of the Northern Territory. Map 10.

Victoria: Hattah Lakes Nat. Park, /t. C. Beauglehole 19817 (PERTH); L. Wahpool, Waitchie, N. Mac-
fartane 851, 854 (PERTH).

South Australia: Lake Meramangyne, N. Fortie 546 (CANB); 150 km E of Cheeseman Peak, R. B. Major
102 (AD): St. Francis Is., N. M. H'ace 103 (AD); Pt. Paterson to Red Cliff Point, D. J. E. Whihtev 5461
(AD); Pt. Pine, P. G. Wihon 10254 (PERTH).

Western Australia: Boulder, IT'. D. Campbell 185 (K); Lake Thros.sell, A. S. George 2994 (PERTH): I -5
km W of Windarra, 29 .luly 1973, H. W/A/;; (PERTH); Hamelin Pool, Sept. 1976, O'. & M. Pen r ( PERTH).

Northern Territory: SE corner of Lake Amadeus, G. Chippendale (NT 6376); Lake Neale, P. K. Latz 4256
(PERTH).

Most collections studied appear to have flowers with a fertile ovary and a vestigial
stamen, or a fertile stamen and an apparently non-functional ovary. However, some
collections had fully fertile flowers and in others it was not clear whether the ovary was
functional but matured considerably later than the stamen.

Hulosurcia pruinosa is somewhat variable in morphology as may be expected from a
plant with such a wide distribution. It may be recognised by its fruiting spikes which
retain their shape (the bracts do not shrivel) and by the fruitlets which separate from each
other and have a firm crustaceous perianth that is muricate on the outside. The relation-
ships of this species are to H. iindulaia, from which it may be distinguished by the firm
shortly cylindrical fruiting bracts and the steeply ascending florets.

The perianth in H. pruinosa is similar to that in H. auriculaia and in the flow'ering
condition the two are difficult to distinguish. In both the lateral lobes are conduplicate,
that is they arc dorsiventrally compressed; on the orifice (which is at the acute apex)
on the abaxial (lower) side they are separated by a medial abaxial lobe. In //. auriculatu
the abaxial lobe is large and broad and the lateral lobes overlap it only on its margins;
in H. pruinosa the abaxial lobe is much smaller and becomes inwardly folded, while the
lateral lobes are large and overlap each other so as to obscure the medial lobe. In H.
auriciilata the florets are fused to each other and the seed is released by the sagittal splitting
of the fruitlets.

Between Marree and Mt. Lyndhurst in South Australia has been collected a plant
which is very similar in fruiting structure to H. pruinosa and may, in fact, intergrade with
that species. It differs from U. pruinosa in having minutely papillose, acutely lobed,
ciliolate articles and in the fruiting spikes taking on a pale straw colour (rather than dark
browm). The following collections represent this taxon (all PERTH); 14 km NW of
Lyndhurst, M. Lazarides 8259; 27 km SE of Marree, M. Lazarides ?i319, 8780. Further
collections are required before it can be determined whether it should be recognized as
a distinct species.

15. Halosarcia auriculata P. G. Wilson, sp. nov. Fig. 27A-C, 54, 55.

Fruticuhts divaricalus ad 0-5 ni altus. Articiili doliiformes glaiiei. rnargine -Jt inincalo, integro.
Spicae terminales, erucaeformes, 1-2 cm longae, ca. 4 mm diam.; bractcac lirmae aliquantum tumidae,
interdum demum suberosae. Flores bractcis occLihi, vicinas et bracteas superas coalescentes. parti proxim-
ali cum bractea intera conjuncti; facies abaxialis valde adscendens. Perianthiinn apicem dorsi-ventraliter
applanatum. parietibus lateralibus crassis, aliter tenue; facie abaxialc cum lobo medio grandis, lobis lateralis
auriculatis superpositis. Perianthiinn in fructo crirstaceum, in piano sagitlali secedens. Pcricarpiunt
membranaceum vel ad apicem crustaceum. Semen applanato ellipsoideum, ca. 1 mm longum; testa
tenuis.

Type: 13 km N of Sandfire Roadhouse, approximately midway between Pt. Hedland and
Broome; blue colour evident, 25-30 cm high; 30 .luly 1973; R. A. Saff'rev 1707 (holo;
PERTH).

Much-branched spreading shrub to 0-5 m high. Articles doleiform, glaucous;
margin i truncate, entire. Spikes terminal to branches, cruciform, 1-2 cm long, ca.
4 mm diameter; bracts firm, somewhat sw'ollen, sometimes corky when mature. Flowers
in ± contiguous triads, completely obscured by bracts, steeply ascending on abaxial side,

56

Nuytsia Vol. 3 (I) 1980

coalescent to each other and to upper bract, fused to lower bract in proximal half. Perianth
strongly dorsiventrally flattened at apex, thin above and below in medial position but
with thick lateral walls, at first succulent but soon becoming crustaceous; abaxial face
of flattened apex with a large rounded medial lobe which is overlapped on either side by
smaller auriculate lobes, these being continuous with the flat adaxial margin; anther ca
I -2 mm long; style short and firm with two short stigmatic lobes. Fruit lets obscured by
bracts, firmly fused laterally to each other, adaxial margin vertical to spike axis, abaxial
margin steeply ascending, apex strongly dorsiventrally flattened; perianth crustaceous,
thin in medial position and readily splitting in sagittal plane, lateral walls firm; pericarp
laterally compressed, membranous and hyaline apart from the thickened apex, free from
perianth. Seed ovate, I mm long, adherent to the pericarp; testa thin, pale brown,
concentrically granular over embryo; embryo curved. Seed with pericarp released by
the decay of the spike and the sagittal splitting of the perianth.

Habitat: Saline clay pans.

Distribution: North-western Western Australia, predominantly in near coastal situations.
Map 13.

H'esieni Australia: b-S km from Beagle Bay road on Willie Creek road, A. C. Beauglehole 48198 (PERTH);
43 km S of Lagrange Mission turnolT, A. C. Beauglehole 48279 (PERTH); 16 mi S of Vlaming Head. A. S
George 6623 (PERTH); Lake Macleod, Mav 1977. M. Lewis (PERTH); 13 mi E of Broome, M. Lazaiides
6578 (AD. CANB. K, PERTH); 2 mi NW of Well 24, Canning Stock Route, B. R. Masliu 2252 (PERTH);
50 km SW of Onslow, A. A. Mitchell 515a (PERTH); Pt. Hedland, I3.V.1971. G. /. Parlerliel (PERTH).

Haiosarcia aiiriculata is closely related to H. syncarpa; they have similarly shaped
flowers, (which are fused together), and virtually identical seed. It differs most obviously
in having the spikes terminating the branches, and in the lobing of the perianth; the
fruiting and flowering spikes look very different as do the general habits of the two species
for in //. auriciilaia the articles are barrel-shaped and of a bluish glaucous colour whereas
in H. syncarpa they are obovoid and green.

Haiosarcia priiinosa, has a similar appearance to H. auriculata but differs in the shape
of its spikes, the lobing of the perianth (no abaxial lobe or only a very small one) and in
having free fruitlets with muricate lateral walls to the perianths.

All the collections studied of H. auriculata have a bluish glaucous appearance, a
character which appears to persist throughout the life of the plant.

In H. auriculata the flowers are fused to the lower bract in their proximal half; this
IS unusual in the genus where normally fusion only occurs with the upper bract (if at all).
Because of this fusion the fruitlets remain attached to the lower bract when the spike is
broken whereas in other species they preferentially adhere to the upper bract.

The specific epithet refers to the small ear-like perianth lobes which are positioned
on either side of the large rounded abaxial lobe.

16. Haiosarcia syncarpa P. G. Wilson, sp. nov. Fig. 29E-G.

Ft uticulosus decumbens vel erectus, 0-2-1 m altus. Articuli late obovoidei, ca. 5 mm long! et lati
apiculati, margme denticulato vel integro. Spicae primum terminales, jam intercalares. Flores ± obtecti’
abaxiale valde adscendente. Periatnhium praeter latos laterales tenue, apicem’
dorst ventraliter applanatum; lobi duo, laterales conduplicati. imbricati. Spica fructifera intercalaris-
bracteae demum tenues, siccae (vel mterdum medullosae), fructicull.s obtectis (vel interdum demum ex-
Mhpr nf ‘^'■“Staceum vel conaceum (in medio tenui) ad vicinas conjunctum; pericarpium

liberum, lateraliter compressum, spongiosum vel crustaceum, ad apiceiim durum. Semen ovoideum,
, l‘’n8us. ad pericarpium - adhaerens; testa membranacea. pallido brunnea, supra embryo
I’iberatilni’” fructirerum demum in piano sagitali fissum. scminibiiis et pericarpm

m 00 ^ 0 - 7 Grace, by main Tarin Rock— Lake Grace Road, 33°7'S,

u^t of May 1969, P. G.

H’dson 8277 (holo; PERTH, iso; CANB, R, NSW).

Paul G. Wilson, Australian Salicornieae

57

Decumbenl to erect siih-shnih 0-2-1 m high. Articles broadly obovoid, ca. 5 mm long
and wide, glossy or dull, bluntly to acutely lobed, apiculate, margin entire or denticulate.
Spikes sessile, or terminal to short lateral branchlets, soon becoming intercalary through
vegetative development of apex, of 1-10 articles, the bracts similar at anthesis to the
vegetative articles. Flowers in triads, r obscured by bracts but with apex protruding at
anthesis, coalescent to each other and in proximal half to upper bract, free from lower;
abaxial surface steeply ascending. Perianth thin at anthesis except for the thickened
lateral walls, deltoid in shape, apex strongly dorsiventrally flattened; lobes 2, lateral,
conduplicate and imbricate (with sometimes a small inner abaxial lobe); style short and
firm with a pair of ligulate stigmas which are thickened at their base. Fruiting spike:
bracts eventually becoming thin, dry or sometimes pithy (in one variant becoming firm)
i totally enclosing the fruitlets or shrivelling to expose them; perianths with leathery to
crustaceous lateral walls which are fu.sed to the neighbouring perianth, thin or torn in
medial position; pericarp laterally compressed, spongy to crustaceous (often hardened
at apex), free from perianth. Seed ovoid, ca. 1-2 mm long, i adherent to pericarp:
testa membranous, pale brown with distinct concentric granulation over embryo; embryo
curved; perisperm lateral. Seed released with pericarp through decay of bracts and
the splitting of perianth in sagittal plane.

Habitat: Principally on margin of salt lakes under strongly saline conditions but also
coastal.

Distribution: Western Victoria, southern South Australia, and southern Western Aus-
tralia. Map 13.

Victoria: Near Dimboola, Reader 20 (MEL); Lowan, 15 Aug. 1897, Reader (MEL); Wimmera, anon.
(MEL).

South Australia: Browns Beach. Yorke Peninsula, C. R. Atcuck 4967 (AD); 5 mi SE ol Wellington, F. M.
Hilton 1536 (ADW); Stamford Hill, P. (7. Wilson 2667 (AD).

Western Australia - Israelite Bay, 1885, 5. Brooke (MEL); 30 km NNE of Stokes Inlet, Hj. Fielder 20320
(AD, PERTH); 9 mi S of Pingrup. K. Newhey 3074 (PERTH); 13 mi N of Norseman, B. L. Turner 5278
(PERTH); 3 km E of Meckering, P. G. Wilson 8678 (PERTH).

Halosarcia syncarpa — Sclerosiegia arhuscula ?

South Australia: Browns Beach, Southern Yorke Peninsula, C. R. Alcock 4964, 4965 (AD); Lower end
of Coorong, D. E. Symon 10484 (ADW, PERTH).

Halosarcia syncarpa — H. undulata

Western Australia: Meckering, P. G. Wilson 8669, 11623 (PERTH).

The description provided by G. Bentham (1870) for Salicornia arbu.scula consisted
of a combination of characters derived chiefly from Sclerosiegia arbitscula and Halosarcia
halocnemoides. The material he cited, however, included specimens of additional species;
three of the specimens were of H. .syncarpa, these are as follows (all MEL); “Wimmera,
Dallachv" p.pte., "Sandflats N of Stirling range, F. Mueller", and “Swan river, Drummond,
I st coll.”

Halo.sarcia syncarpa may be readily distinguished from related species by the perianths
which are fused laterally to the neighbouring members of the cymule. The medial portion
of each perianth is membranous (or absent) allowing the fruitlets to split along the sagittal
plane to release the pericarp with its enclosed seed. The spikes vary considerably in size;
in the typical form they consist of only 1-3 articles but in other forms the spikes may
contain as many as 7; in all cases their apices continue growth vegetatively so that the
fruiting spike is always intercalary.

The great variability exhibited by this species suggests that more than one taxon is
involved. Hybridization with H. undulata and H. appears to occur in Western

Australia, and with Sclerosiegia arhuscula in South Australia. This obviously increases
the difficulty in distinguishing the constituent taxa within the complex. In South Australia
and Victoria the perianths have a small but obvious inner abaxial lobe while in Western

58

Nuytsia Vol. 3 (1) 1980

Australia this is absent (or is extremely minute). In Western Australia plants of //.
syncarpa with very different aspect but with the same fruit type can be toimd growing
together without any apparent sign of introgression, whereas plants gathered from the
total area of its distribution so vary as to cover the entire morphological spectrum. Much
critical field work is needed before the "syncarpa' complex can be understood.

The specific epithet refers to the lateral union of the fruitlets.

17. Halosarcia undulata P. G. Wilson, sp. nov. Figs. 29A-D, 56.

Fniticiilosiis ramosus. Ai ticidi opaci vel glauci, late doiiiformes ca. 7 mm longi; lobi rotundati (inter-
dtim aeuminati), integri. Spicae conicae vel cylindraceae, 10-30 (50) mm longae. 3-5 (9) mm diam.,
ambito undiilalo; bracteae breves, arete imbricatae. eonvexae, margine valde imdiilato, integro. Flores
oblecti parte proximal! ad vicina,s et braeleas coalescenti, alitcr liberi, facies abaxialis plana, leviter vel
valde adscendens, parte distali valde dorsiventraliter applanata: lobi duo, lateraic.s, conduplicali, imbricati.
I'riicticuli demum libri; perianthium spongiosum vel crustaeeum, nitidum; pericarpium membranaceum
(vel ad apice crustaeeum); stylus ad basim durus, acicularis. Semen ovoideum ca. I mm longum; testa
tenuis, brunnea, fere laevis. Bracteae in statu IVuctifcro demum ab axe secedentes; fructiculi in statu
integro cadenti.

Tvpe: 1 km W of Meckering, Western Australia, Mortlock River bed; plant 0-2 m high;
23 Nov. 1967, P. G. Wilson 6404 (holo; PERTH, iso; CANB, K).

Much-branched suh-shnih. Vegetative articles dull to slightly glaucous, broadly
barrel-shaped, ca. 7 mm long; lobes rounded (sometimes acuminate), entire. Spikes
terminal and lateral (when usually sessile by a broad base, the lowest bract pair being very
short), conical to cylindrical, 10-35 (50) mm long, 3-5 (9) mm diameter, outline very
undulate when fresh, the apex sometimes continuing growth as a short vegetative branchlet;
bract-pairs short, closely imbricate, fleshy, outer face prominently convex, margin markedly
undulate (face very thin in inter-bracteal position), entire. Flowers totally enclosed,
united in proximal half to each other and to upper bract; abaxial (lower) surface fiat,
slightly to steeply ascending; adaxial (upper) surface ascending to horizontal, distal
half .strongly dorsiventrally fiattened, apex acute; perianth at first fieshy at sides, thin at
apex, soon becoming brittle, ca. 2 mm long; lobes 2 lateral, conduplicate (each sharply
folded due to dorsivcntral compression of fiow'er), imbricate, extending - half way
along abaxial surface; anther 1 -5-2 mm long (frequently vestigial); style linear, thickened
towards base; stigmas slender. Fniillels separating from each other (sometimes tardily),
apex dorsiventrally fiattened; perianth thin, firm and spongy to crustaceous (especially
within), glossy; pericarp laterally fiattened, membranous (or crustaceous at apex), usually
absent towards axis or. if sterile, then often entirely crustaceous, style base firm and
acicular. Seed ovoid, ca. 1 mm long; testa thin, pale to dark brown, practically smooth
(minutely granular over embryo); embryo slightly curved; perisperm lateral. Bract-pairs
eventually separating from spike axis, the fruitlets falling entire.

Habitat: Saline or gypseous soil.

Distribution: Northern portion of South Australia, southern half of Western Australia,
and southern portion of the Northern Territory. Map 19.

South Australia: Dalhousie Springs, P. K. Latz 4195 (NT).

IVesrern Australia: Cundeeies, P. Boswell J2I (PERTH); 2-3 mi E of Carnegie HS., A. S. George 5545a
(PERTH); Lake Annean, R. Hnutiiik (PERTH); Haniersley estuary, K. Newhev 4900 (PERTH)- Lake
Hinds, P. G. HT/.vfw 1 1649 (PERTH).

Northern Territory: Erldiinda Sin.. T. S. Henshall 54 (NT).

Halosarcia undulata may be recognised by its sessile, broadly based, lateral spikes
and their short markedly undulate bracts. It is probably mo.st closely related to H.
syncarpa which may be distinguished by its predominantly intercalary fruiting spikes and
by the lateral fusion of its fruitlets (free in H. undulata). The fruitlets of H. undulata
are usually very strongly dorsiventrally fiattened towards the apex but in some variants
they are more obtuse, in which case they are similar in shape to those found in H. pruinosa.
In the latter species, however, the perianth is muricate on the outside. Another difference

Paul G. Wilson, Australian Salicornieae

59

between the two species, but one not easily observable, is that in H. pruinosa the flower
has a medial abaxial perianth lobe within the overlapping lateral lobes; in H. iindulata
there is no abaxial lobe; only the lateral lobes are present and these are prominently dorsi-
ventrally compressed.

Specimens of H. uiuliilata from the north-eastern region of its distribution in Western
Australia have acuminate articles and bracts whereas in the southern region the plants
have shortly apiculate articles. The material from the different regions is itself otherwise
far from uniform, but the variation observed could frequently be due to conditions of
growth, rather than a reflection of genetic diversity, since spikes on different parts of the
same plant, representing growth of different seasons, may vary considerably in both size
and shape.

There is only one record of the occurrence of H. undulata in the Northern Territory
and this specimen differs from those from Western Australia in not having sessile spikes
and in the fruitlets adhering to each other. Further work may indicate that the Northern
Territory plant should be recognised as a distinct species.

A number of collections appear to be either of a species closely related to H. undulata,
or of a hybrid between H. undulata and another species. These specimens have ovoid
to ellipsoidal pedunculate spikes, pithy perianths, and black carbonaceous pericarps.
Very few of the flowers set seed, and when they do the seed is adherent to the pericarp
and possesses a noticeably papillose testa (almost smooth in //. undulata). Examples of
this putative hybrid are as follows:

Western Aiisiraliti; Cue,J. H. Makleii (NSW 1.^6567); Lake Austin, P. G. Wilson 8,S69 (PERTH): Mongers
Lake, P. G. Wilson «(S02 (PERTH); Lake Moore, P. G. Wihon 8637 (PERTH): Luke Bailee, P. G. Wilson
8856 (PERTH): Lake Miranda, P. G. Wilson 8924 (PERTH).

18. Halosarcia bulbosa P. G. Wilson, sp. nov. Fig. 30.

Fniticnliis effaiusca. 1 maltus. ,‘)/7/c////grossi. doliil'onnes, ca. (5x 12 mm, truncati, pruinosi. Spicae
laterales, breviter pedunculalae, cylindrico-fusiformes, 15-20 mm longae, saepe demum intercalares;
bracteae undulatae, imbricatae. Fiores hermaphroditi. in parte proximali ad vicinas et bracteas coales-
centes, in parte di.stali actiminati, dorsi-ventraliter applanati. Perianthinnt ad apicem prol'unde bilobatum.
Spicae in statu rrucliferis: bracteae cupulatae, leviter undulatae, coriaceae; fructiculi libri, ascendentes;
perianthium crustaecum, nigrum, profunde areolatum, apice truncate prominente roslrato; pericarpium
ad apicem cruslaceum, after membranaceum. Semen ovatuni ca. 1-3 mm longum; testa membranacea,
laevis, pallido brunnea.

Type: Western Australia, 10 km E of Morawa, 29 April 1979, P. G. Wihon I 1702 (holo:
PERTH, iso: CANS, K).

Sprawling shruh ca. I m high and 2-3 m diameter. Branches spreading: articles
barrel-shaped, ca. 15 x 12 mm, truncate with only a very narrow scarious rim, bluish green
and pruinose when fresh. Spikes lateral, sub-sessile, fusiform-cylindrical, 15-20 mm
long, often becoming intercalary w'ith age; bract-pairs closely imbricate, undulate; triads
completely enclosed. Flowers hermaphrodite, united in proximal half to bracts and to
each other, a.scending, acuminate and dorsiventrally flattened in distal half. Perianth
with lateral walls succulent, otherwise thin, the distal (acuminate) half deeply divided
into tw'o lateral lobes, a small abaxial lobe sometimes present; anther ca. 1-7 mm long;
ovary ellipsoidal; style arms ligulate. Fruiting spike persistent, dark-brown (when dry);
bracts imbricate, cup-shaped, leathery, with an abaxial flange which partially covers
apex of flowers. Fruitlets ascending, free from each other and from bracts; perianth
crustaceous, black, deeply areolate where previously united to lateral fruitlets and bracts,
apex oblong and smooth with a beak-like apiculum; pericarp thinly crustaceous at apex,
membranous below, free from perianth. Seed ovate, ca. 1-3 mm long; testa membranous,
smooth, pale brown.

Habitat: Open Melaleuca shrubland on slightly saline reddish-brown loam (Soil pH
8-7, NaCI 3-4%).

Distribution: Known only from one locality between Morawa and Koolanooka Hills
ca. 150 km SE of Geraldton, Western Australia. Map 8.

Western Australia: 10 km E of Morawa, 1977, H. Demarz iPEKlH).

60

Nuytsia Vol, 3 (1) 1980

This species is noticeable because of its large prtiinose articles which exceed those of
any other species of Halosarcia. It is evidently most closely related to H. pruinosa and
H. iinclulata as is suggested by the similar fruits.

The old spikes appear to remain attached to the plant for several years and presumably
fall only with the decay of the dead branches. The fruitlets are tightly enclosed within
the cup-shaped bracts and are securely held in place by a flange on the underside of the
bracts. Evidently, therefore, the seed is released only after the decay of the bracts and
perianth.

The size of the articles in H. hulhosa may be partly a response to the particular soil
in which the plant grows for when transplanted to non-saline sandy loam it develops
narrower branchlets. However, the other Halosarcia species found with H. bulbosa
had articles of normal size; these species were H. halocnemoides subsp. halocnenwides,
H. pergranidata subsp. pergraniilata, and H. pruinosa.

19. Halosarcia entrichoma P. G. Wilson, sp. nov. Fig. 31A-D.

fn/nVH/o.m.T decumbens. ArtkuU crpaa vcl glauci. late obovoidei, lobis rolundalis, margine cilialato.
Spicae terminates, ovoideae, ambito undulato; bracteae tumidae, profunde undulatae, margine ciliolato.
Flores bracteis occulti, liberi; facies abaxialis valde adscendens. Periantluiim tenuiter coriaceum, lateraliter
compressum, apicem dorsi-ventraliter applanatum, orificio terminali; lobi laterales super iacieni abaxialem
inserti, imbricati, denticulaii; lobus mediiis abaxialis parvus, semi-eircularis, internus. FructicuU liberi,
ab axim facile secedentes, integri; perianthiiim tenuiter cartilagineum; pericarpium ad apicem crustaceum,
basim versus membranaceum. Semen ovoideum ca, 2 mm longum; testa membranacea, laevis, straminea.

Type: Frank Hann National Park, 79-2 km E of Fake King township, 32''52'S, 120 25 'E,
Western Australia, 15 Oct. 1978, D. Monk 482 (holo; PERTH, iso; CANB, K, MEL,
NSW).

Dwarf semi-decumbent shrub. Articles dull green to glaucous, broadly obovoid,
ca. 10 mm long, 5 mm broad; lobes rounded, margin cililate. Spikes terminal to branches,
ovoid, ca. 20 mm long, 8 mm broad, outline uneven; bract pairs deeply undulate, swollen,
margin ciliolate. Flowers hermaphrodite, obscured by bracts, adaxial face horizontal,
abaxial face steeply ascending. Perianth thinly coriaceous, laterally compressed; apex
dorsiventrally flattened with orifice at terminal edge; lateral lobes imbricate on abaxial
face, denticulate; medial abaxial lobe semi-circular, small, overlapped by lateral lobes.
Fruitlets free from each other and from bracts, readily separating from axis, entire (not
tearing at base); perianth cartilaginous; pericarp crustaceous at apex, membranous towards
base. Seed laterally flattened, ellipsoidal, ca. 2 mm long; testa membranous, smooth,
pale fawn in colour.

Habitat: Margin of slightly brackish lakes in clay soil.

Distribution: Only known from the type locality near Lake King township. Western Aus-
tralia. Map 10.

Western AtistmUa: yvank Hann National Park, 32 53 'S, 120 26 'E, around lake (fresh in winter) D N
Butcher (PERTH). ’

Halosarcia entrichoma is readily recognisable on account of its large obovoid articles
which are ciliolate on their margin; these, with the free fruitlets and cartilaginous perianths
clearly demarcate it from other species.

The lake around which the collections have been made is relatively fresh in winter
and only becomes brackish in summer; this situation is unusual in inland Western Australia
and may account for the apparent rarity of the species. An analysis of the soil after the
lake had dried up indicated an NaCI content of 0-24% in the top 10 cm and I 05'/ at
a depth of 30-40 cm.

The specific epithet is a Greek word for eyelash and alludes to the ciliolate margin
of the bracts and leaf-lobes.

Paul G. Wilson, Australian Salicornieae

61

20. Halosarcia ctiartacea P. G. Wilson, sp. nov. Fig. 20 D F, 57.

Fniticiilns ad I m altus. Ariiciili angusle obovoidei, ca. 10 mm longi, margine truncate, integro.
Spicae terminales cylindraceae, 20-40 mm longae, 4-6 mm diam.; bracteae confertim imbricatae truncatae,
margine integro. Flores I'ere occulti. ad vicinas et bracteam superum coalescentes. Perianihiiim tenue,
apicem dorsiventraliter applanatum; lobi 2, laterales, conduplicati. Ovarium membranaceum, ad basim
perianthii affixuin; stylus tenuis. Periamhium in friicto librum integrum, aliquantum urceolalum, ad 4
mm longum, chartaceum, lateribus reticulatis, semine incluso; pericarpium diaphanum. Semen late
ellipsoideum ca. 2 mm longum; testa diaphana, laevis.

Type: 5 km S of Warren Bore, ca. 115 km ENE of Bandya HS, Western Australia; flooded
clay pan, plant I m high; 28 Aug. 1968, P. G. Wilson 7446 (holo: PERTH, iso: AD
96947008, NSW).

Erect dwarf s hr uh to I m high. Vegetative tirlicles narrowly obovoid, ca. 10 mm long,
apices with a scarious truncate margin, entire. Spikes terminal to short lateral branchlets
(often of one internode) but apparently not to main branches, cylindrical, 20-40 mm
long, 4-6 mm diam.; bracts closely imbricate (becoming separated on drying), margin
dr truncate, entire. Flowers with the opposite triads contiguous, steeply ascending,
almost entirely included with bracts, adherent to each other and at first to upper bract;
perianth thin (not succulent), the apex obtuse, dorsiventrally llattened with terminal
orifice; lobes 2, lateral and conduplicate anther ca. 2 mm long; ovary membranous,
attached to base of perianth, style and stigmas tenuous. Fniiilels eventually free, somewhat
urceolate, to 4 mm long; perianth chartaceous, reticulate where formerly adherent to
bract, completely enclo.sing seed; pericarp diaphanous (somewhat thicker at apex) closely
enfolding seed. Seed broadly ellipsoidal; perisperm copious along length of embryo.

Habitat: On only slightly saline clay pans.

Distribution: Central Western Australia. Map 12.

Western Australia: Lake Yindarlgooda 70 km E of Kalgoorlie, Jane Elkinglon 628 (PERTH); llgarari
Mine, 120 km S of Mt. Newman, A. Mitchell 318 (PERTH).

This species is distinctive in having fruitlets which readily fall away from the dry
spike, a papery perianth, and a diaphanous pericarp and testa. It evidently occurs only
on clay pans and in these situations it may form a mono-specific community.

The specific epithet refers to the papery consistency of the perianth in fruit.

21. Halosarcia cupuliforniis F. G. Wilson, sp. nov. Fig. 32.

Friiticulosiis ca. 25 cm altus. Rannili tenues; articuli anguste cylindracei ca. 10 mm longi, lobis
rotundatis breviter acuminatis, Spicae terminales anguste cylindraceae, ad 8 cm longae, 4 mm diam,;
bracteae cupulil'ormes ca. 2-5 mm altae, breviter acuminatae, in sicco laxe imbricatae. Flores omnino
occulti, ad vicina et bracteam superam coalescentes, facie abaxiali valde ad.scendente. Perianthium suc-
culentum. 2 mm longum in plana sagittal! fere divisum in dimidiis succulentis, apicem dorsi-ventraliter
applanatum; stylus tenuis, Perianthium in fructo liberum, integrum, leviter medullosum, in dimidiis
lateralibus s'ecedens; pericarpium membranaceum (in parte proximal! abscens). Semen ovatum ca. 1-2
mm longum; testa pallido fusca, super radiculam minute tuberculata.

Tvpc: Pulchera Waterhole 30 mi WNW of Sandringham Stn., Queensland; rare on edge
of waterhole in white day soil; 21 Jan. 1970: P. K. Latz 518 (holo: PERTH, iso: CANB,
NT 26711 (n.v.) ).

Sub-shrub ca. 25 cm high. Branches slender; articles dull or glaucous, narrowly
cylindrical, ca. 10 mm long, apex with rounded shortly acuminate lobes. Spikes terminal,
narrowly cylindrical, to 8 cm long, 4 mm diameter; bracts cupuliform, ca. 2-5 mm high,
loosely imbricate in dried state, obtuse and .shortly acuminate, with a prominent scarious
margin; triads completely hidden, contiguous. Flowers adherent to each other and to
upper bract throughout their length, abaxial surface steeply ascending; perianth succulent,
2 mm long, divided on abaxial surface in sagittal plain to form two succulent lateral halves,
apex obtuse and dorsiventrally compressed; anther ca. I mm long; style and stigmas

62

Nuytsia Vol. 3 (I) 1980

long and slender, Fruiiiiiii spike not seen in mature state; fruitlets (perianth with enclosed
pericarp and seed) eventually free from each other and from bracts; perianth weakly
pithy, separating into two lateral halves; pericarp membranous (absent in proximal half).
Seed ovate ca. 1-2 mm long; testa pale brown, minutely tuberculate over embryo; embryo
almost straight; perisperm lateral.

Habitat: Only known from the description accompanying the type specimen.

Distribution: South-west Queensland and north-east South Australia. Map 2.

Queenskuut: Pulchera Waterhole. 30 mi WNW of Sandringham, P. K. Lutz 518 (CANB, NT, PERTH).

South Australia: Eyre Creek, 26 S, 1889, A. Henry (MEL). Note: 26 latitude delineates the Queensland-
South Australia border.

Halosarcia cupuliformis may be recognised by its distinctive spikes in which the
abruptly acuminately lobed bracts form cup-like rings around the axis. In flower structure
it is similar to H. pruinosa but the perianth is much thinner and does not harden with
age.

The specific epithet refers to the cup-shaped appearance of the bracts when dry.

22. Halosarcia calyptrata P. G. Wilson, sp. nov. Fig. 25A~C, 58, 59.

Fruticiilosus ad 0-8 m altus. Ramii/i tenues: articuli doleiformes vel obovoidei ca. 6 mm longi, 3 mm
diam., margine leviter lobato; integro. Spicae angustc cylindraceae 10-20 (40) mm longae, 3 mm diam.;
bracteae breyiier doleiformes, ca. 3 mm altae, -t truncatae, confertim imbricatae, integrae. Flores bracteis
occulti ad vicinas et bracteas superas parti proximali conjuncti. Perianthium apicem dorsi-ventraliter
applanatum, parietibus latcralibus crassis, lobis lateralis iiiibricatis (lobo abaxali absent! ?). Spica fruetifera
persistens; braetea et perianthium crustaceum; pericarpium ad apicem crassum, crustaceum, operculatum,
ad basim membranaceum; stylus ad basim durus, persistens. Semen ovoideum, ca. I mm longum; testa
membranacea, super radiculam granulata.

Type: 18 km S of Wiluna, Western Australia; claypan; sub-shrub 0-3 m high, articles
dull pale green; 28 Aug. 1970; P. G. Wilson 8939 (holo; PERTH, iso; CANB, K).

Divaricately branched sub-shrub to 0-8 m high; branches slender. Articles doleiform
to obovoid, ca. 6 mm long, 3 mm diam., dull to slightly glaucous; margin broad and
scarious, lobes very short and rounded. Spikes narrowly cylindrical, 10-20 (40) mm
long. 3 mm diam., terminal (frequently becoming intercalary) or sessile and lateral; axis
thick and woody; bracts shortly doleiform. ca. 3 mm high, almost truncate, closely imbricate
thick and woody; bracts shortly doleiform, ca. 3 mm high, almost truncate, closely im-
bricate, entire; triads ± contiguous. Flowers completely obscured by bracts, fused
laterally to each other and (sometimes?) in proximal half to upper bract; perianth succulent
on lateral margins, otherwise thin, deltoid-acuminate in lateral view, -F vertical to spike
axis (abaxial side ascending and adaxial descending), orifice at the acute apex; lateral
lobes imbricate and conduplicate (no abaxial lobe observed), the perianth 1 splitting
at anthesis into two lateral halves; anther ca. 1-3 mm long; ovary deltoid, fixed along
vertical length of flower to spike axis; style and .stigmas slender. Fruiting spike persisting
on dead branchlets or, if intercalary, around living branch; bracts and perianth usually
crustaceous; pericarp thick and crustaceous at apex forming an ellipsoidal cymbiform
cap to seed, membranous towards axis; style base hard and persistent frequently forming
a small conical projection in centre of fruitlet. Sm/ ovoid to narrowly ellipsoidal, I mni
long; testa thin, pale brown, granulate in longitudinal lines over radicle; embryo straight.

Habitat: Margin of gypseous salt lakes (Northern Territory) and on slightly saline clay
pans (Western Australia).

Distribution: Central Australia and north central Western Australia. Map 22.

Northern Territory: Lake Amadeus. 26 June 1959. G. Chippendale (NT 6377, AD); Lake Mackav 17 June
(N^) ^ iuppendaie (NT 3401. AD, BRI, CANB, NSW, PERTH); Lake Amadeus, T. S. Henshaii 748

Western Australia: Mininer to Mt. Vernon Stn„ A. S. Mitchell 76/189 (PERTH)- ligalonu R D Rovre
1557 (PERTH): 49 km N of Mt. Magnet, P. G. Wilson 8563 (PERTH); 34 km W of Wiluna P G When
8963 (PERTH): 25 km NE ol Balfour Downs H.S„ P. G. Wilson 10371 (PERTH)- Rtidall River P G
Wilson 10578 (PERTH).

Paul G. Wilson, Australian Salicornieae

63

The plant as found in the Northern Terrtiory has terminal inflorescences while in
Western Australia these are usually lateral and sessile. Further collecting may indicate
that two species are involved which would then account for the apparent differences in
habitat.

A distinguishing feature of this species is its thick cap-shaped pericarp; the plant
is also interesting in having a hard fruiting spike due to the indurated bracts, perianth,
and pericarp, and to (he thick lignified axis. In those cases where the spike-axis dies the
spike may persist for some time (a year or more) as a short lateral branchlet; these fruiting
spikes eventually break up into separate segments composed of the opposite triads with
the surrounding pair of bracts. In other cases the spike axis continues growth vegetatively
and the fruitlets persist on the branch for some years until they are eventually sloughed off
in a similar manner to that of the bark.

Collections from near Wiluna (including the type) frequently have two spikes arising
from a leaf axil (that is, four spikes to a node), or one of the spikes may be replaced by
a branchlet. No anatomical investigation has been made of this phenomenon.

The specific epithet refers to the cap-like covering to the seed formed from the hardened
apical portion of the pericarp.

23. Halosarcia indica (Willd.) P. G. Wilson, comb. nov.

Saliconiia indica Willd.. Ges. Naturf. Fr. Neue Schr. 2:111 t.4 f.2 (1799 ). — Arihrocnemiim indicum (Willd.)
Moq., Chenop. Enum. 113 (1840), p.p.; Moq. in DC., Prod. 13 (2): 151 (1849), p.p.; Ungern-Sternberg,
Versuch Syst. Salicornieen 69 (1866); Ungern-Sternberg, D. Aui del Cong. Int. Bot. Firenze 1874: 282
( 1876); J. D. Hooker. FI. Brit. India 5:12 ( 1886): Brenan in Turrill et Milne-Redhead, FI. Trop. E. Africa —
Chenopodiaceae 18 (1954); Tolken, Bothalia 9:276 f3/3 (1967 ). — Sarcathria indica (Willd.) Rafinesque
ex D. Jackson, Ind. Kew. 2:803 (1895).

Type: Tranquebar, India, J. G. Klein (B).

Decumbent to erect perennial, frequently glaucous. Articles of branchlets slender
or thick, cylindrical and truncate to somewhat flattened, obovoid and deeply lobed, margin
entire or ciliolate. Spikes terminal to main and lateral branchlets, smooth and cylindrical
and up to 4 cm long or ovoid and up to 2 cm long. Opposite bracts united, the bract-
pair truncate to deeply lobed, entire or ciliolate. the cymules completely disjunct. Flowers
coalescent to each other and to upper bract or free except near axis, completely enclosed
or the apex exposed: upper face horizontal, lower (abaxial) face ascending; apex rounded
or obtuse. Perianth soon becoming fleshy (especially the lateral walls); lobes 3, the
two lateral thin, large, • valvate to deeply imbricate, (the outer one frequently fimbriate);
lower (abaxial) medial lobe within, triangular and small to semi-circular and then equal
in size to lateral lobes (sometimes apparently absent). Fruiting spike broadly ovoid
(SLibsp. hiclens) to cylindrical; bracts leathery, corky, or spongy. Fruitlets -± obscured
by bracts (except in stibsp. /;//6'CC«). laterally compressed, triangular in side view, I -2-2 mm
high, apex rounded; perianth soft and spongy or pithy and firm, sometimes crustaceous
within, pericarp hard and horny all over, often adherent to perianth, eventually splitting
in medial sagittal plane. Seed broadly ovate to almost circular; testa membranous,
very pale brown, smooth and glossy (faintly reticulate and dull in subsp. /u/wccfl); embryo
curved, perisperm lateral. Spike disarticulating with age and releasing the bract-pairs
and fruitlets, the axis often persisting; bracts and perianth in semi-mature spikes frequently
swelling when wet (at least in subspp. hidens and Iciostachya).

Halosarcia indica, as here circumscribed, exhibits considerable variability and an
attempt has been made to provide infra-specific names for the more prominent of these
variants. The typical sub-species is almost prostrate in habit and inhabits the tropical
coasts of east Africa and of India where it is very uniform in appearance. A plant
apparently identical to the type is also found on the north coast of Australia; here, however,
it grades imperceptibly into subsp. Iciostachya.

64

Nuytsia Vol. 3 (I) 1980

In Malaysia is found a plant which differs sharply from that of India and Africa;
it has an erect habit, slender branches, and narrow cylindrical spikes. The articles (both
vegetative and fertile) are noticeably ciliolate. This plant was referred by Backer in
Flora Malesiana 4: 104 (1949) to Arilirnau’imim imiicum. Brenan, however, in Flora of
Tropical East Africa— Chenopodiaceae 20 (1954), considered the Malaysian plant to
represent a distinct species. The plant found in Malaysia was described by Ungern-
Sternberg as A. ciliolatiini, which name was based on material collected in Timor and
Sumbawa (Indonesia). On the north coast of Australia may be found plants which
closely match those of Indonesia and which again appear to grade into the common
Eremaean taxon which is here referred to Hatosarcia indica subsp. kiostachya.

In southern temperate Australia is found a variant with markedly lobed articles
and a robust habit. This plant was described as A. hidens, but again there is no clear
separation between it and the Eremaean taxon.

Halosarcia indica (as Salicornia indica) was first recorded for Australia by Robert
Brown (1810). Brown's description was largely based on material of Sarcocornia quin-
queflora ( Ung.-Sternb.) A. J. Scott but his species concept was broad and in this case
encompassed a number of taxa including H. indica sensu lato.

Halosarcia indica (as Arthrocnennim indicnm) was again recorded for Australia (with-
out validation) by Moquin (1840). This record was probably based on Robert Brown's
treatment for in a later revision Moquin (1849) omitted any reference to Australia but
tentatively (and incorrectly) referred Brown's misapplication of the name to Halocnennim
aiistralasicuni. C. E. Moss (1954) also recorded //. indica for Australia and from his
text it would appear that he had seen material of plants here referred to the sub-species
hidens and leiostacliya.

In related species important characters can be found in the lobes of the perianth,
but these are usually difficult to distinguish in Halosarcia indica. They consist basically
of a pair of large semi-circular lateral lobes extending downwards from the upper margin
of the apex of the flower. These lobes extensively overlap one another and are usually
somewhat erose or ciliolate; the inner of the lateral lobes may (at least initially) be infolded.
Similarly the lower (abaxial and innermost) lobe may not be observable, but although
it varies considerably in size its apparent absence is frequently due to it also being infolded.
In the sub-species jidacea the abaxial side of the perianth is e.Ktremely thin and soon
ruptures with the development of the anther: the lateral lobes scarcely (or do not) overlap
and the abaxial lobe is either minute or absent. In some variants of subsp. kiostachya
the abaxial margin is also extremely thin and the lobe very small.

The variation in perianth .structure would probably form the basis of a sound infra-
specific classification, however, this structure is only decipherable in young flowers in a
good state of preservation (preferably either fresh or in a fluid preservative) and since
only a few collections are suitable for detailed analysis it is not possible to make much
use of this character.

The texture of the perianth is also extremely variable. In many collections the
flowers when young have a thin soft perianth and may persist in this state even when
seed formation is well advanced. Normally the perianth rapidly becomes succulent
and changes to a dry and spongy te.xture as the fruitlets mature. The factors which
influence the various conditions are not clear but generally speaking the fruiting spikes
do not fragment until a year after anthesis and in desert situations they may remain
attached to the branches for several years.

As has been mentioned earlier, Halosarcia indica differs markedly in its anatomy
from all other species of the genus and, in fact, from all other members of the Salicornkae.
The palisade layer consists of a single row of cells which is made up of groups of chloren-
chyma cells (situated beneath the stomata) and groups of colourless cells (that may form
a reticulate pattern between the chlorenchyma clusters). Beneath the palisade layer is
a single layer of small close-fitting thick-walled isodiametric chlorenchymous cells and
this forms a continuous sheath between the aqueous and palisade tissues. The walls of
these isodiametric cells often become lignified with age. The nature of the chlorenchyma

Paul G. Wilson, Australian Salicornieae

65

and associated tissues provides a means of identifying H. indica when dealing with vege-
tative material, for the distinctive anatomy may be readily observed in fresh specimens
with a hand lens, or in dry specimens with a microscope (Fig. I).

Key to subspecies

1. Pericarp horny all over; testa smooth and glossy; fruitlets l obscured by bracts 2
1*. Pericarp crustaceous at apex, membranous below; testa faintly reticulate, dull;

fruitlets not obscured by bracts 23d. subsp. julacea

2. Plant decumbent or prostrate; articles of branches corky with age, d: truncate

entire (tropical coastal) ,,,, . 23a. subsp. indica

2*. Plants • erect ; cortical tissue of articles shrivelling with age, often lobed, d: ciliolate 3

3. Spike cylindrical or ellipsoidal, lowest ring of bracts small; vegetative articles

truncate or shallowly lobed 23b. subsp. leiostachya

3*. Spike ovoid, lowest ring of bracts large and deeply lobed; vegetative articles
deeply lobed 23c. subsp. bidens

23a. subsp. indica. Fig. 61.

Decumbent perennial forming mats ca. 1 m diameter. Artkies cylindrical to barrel-
shaped, to 10 mm long, 4-6 mm diameter, becoming corky with age. Spikes cylindrical
to 2 cm long, grey with age; bracts closely overlapping, eciliate, the lowest pair di equal
in size to (or smaller than) those above it. Flowers fused to upper bract and to each other,
predominantly (always?) female. Fruitlets: perianth spongy; pericarp hard, horny all
over. Seed ovate; testa smooth and glossy.

Habitat: Tidal mudflats.

Distribution: North coast of Australia; the tropical coasts of the Indian Ocean; according
to Tolken, l.c., also occasionally on the west coast of Africa. Map 9.

QiieeiislancI: Curtis Is. 21 Apr. 1962, tC. Maciiae (BRI); Mouth of Settlement Ck., R. A. Perry 1250 (BRI,
CANB, NSW); Karumba, G. tC. Trapnetl 197 (BRI); Townsville Common, 24 Feb. 1977, P. G. Wilson
(PERTH).

Northern Territory: Roper River near mouth, N. Byrnes 1616 (NT); Bing Bong Stn., E. Paine (NT 8513).
India: Peninsula In. Orienlalis, ex Herb. Wight (NSW 136656).

Tanzania: Kunduchi, A. McCusker 242 (NSW).

This is the most widespread of the sub-species of H. indica and shows very little
variability throughout its range, although in Australia it grades into the local variants of
subsp. leiostachya. In its typical form it is found in Australia only along the north and
north-east coasts of Queensland and the east coast of the Northern Territory. Rather
similar plants, however, have been collected elsewhere on the Northern Territory coast
and on the north coast of Western Australia. Brenan, l.c., assumed the species to be
dioecious since he failed to find any stamens in the flowers he examined. Similar comments
have appeared on some herbarium sheets, but in all cases I have examined the supposedly
male plants have turned out to belong to a quite different species. Tolken, l.c., observed
stamens in fresh material from Mozambique and found the flowers to be normally herm-
aphrodite. 1 have been unable to find stamens in any plants from Africa or in plants of
the typical variant in Australia. At Townsville subsp. indica was observed in the field,
and again only female plants were seen although seed-set was occurring. It would appear
probable that this sub-species is normally apomictic. (See also comments on partheno-
genesis in H. pergraniilata subsp. queenskindica).

The mature spikes in subsp. indica break down through the separation of the bract-
pairs which fall away as distinct articles from the slender axis. The triads of fruitlets
at first remain attached to a bract-pair (usually the upper) which are buoyant in water.

This sub-species diflers from subsp. leiostachya in habit and in gross morphology;
it may also be distinguished by the eciliate margin of the leaves and bracts. Some Aus-
tralian material, how'ever, probably as a result of introgression with subsp. leiostachya,
is minutely and irregularly ciliolate.

91670 — ( 5 )

66

Nuytsia Vol. 3 (1) 1980

23b. subsp. leiostachya (Benth.) P. G. Wilson, stat. et comb. iiov. Fig. 9.

Salkornia leioslachya Benih., FI. Austral. 5: 203 (1870 ). — Arthrocnemum leiosnichya (Benth.) Paulsen,
Dansk Bot. Ark.2(8):6l (I9I8);J. M. Black, Trans. Roy. Soc. S. Austral. 43 : 360 (1919).

Ledotype: Kyejeron Creek, J. P. Murray (holo: K, iso: MEL 70595 and 70596, “A. W.
Howitt”), lecto. nov.

? A. ciliolalKm Bunge ex Ung.-Sternb., Versuch Syst. Salicornieen 69 (1866); Ung.-Sternb., D. Atti del
Cong. Int. Bot. Firenze 1874; 283 (1876). Type: From Timor and Stimbawa, n.v.

S. ciUolutum Bunge, Linnaca 28: 573 (1856), nomen.

A. henlhcimii Paulsen. Dansk Bot. Ark. 2 (8): 62, pi. 6 f.2 (1918). Lectotype: Point Samson (Cossacks),
C. H. Ostenfeld 1 143 (iso: NSW), lecto. nov.

A. hrachvstachvum Paulsen, op. cit. 64, pi. 6 f.4. Type: Carnarvon, 31 Oct, 1914. C, H. Ostenfeld 352
(iso: K, MEL, NSW).

Figure 9. Hcilosareia iiidica subsp. leioslachya: from Derby, Western Australia.

Decumbenl to erect suh-shruh. Articlex thick, cylindrical to obovoid, to 10 mm long
slightly lobed, ciliolate. Spikes ellipsoidal to cylindrical 5 40 mm long, with smooth
outline when mature, bracts closely overlapping, the lowest pair smaller than, or equal
in size to, those above it Flowers initially fused to upper bract and to each other’, normally
hermaphrodite. Fiuitiiig spikes leathery to corky: perianth soft and spongy* pericarp
hard and horny all over. Seed ovate; testa smooth and glossy.

Paul G. Wilson, Australian Salicornieae

67

Habitat: Tropical coast and inland around sail lakes.

Distribution: Widespread in Australia; Malaysia. Map 14.

Queensland: Lake Galilee, Adams 1233 (BRl. CANB); 15 mi N of Brisbane, L. A. S. Johnson (NSW 16079)'
Waroo, K. I. Morris 5 (BRl).

New South H'ales: Onepar Stn., E. G. Cuthberlson (CANB); Broken Hill, Haviland (NSW).

Victoria: 25 km NW of Kerang, A. C. Beaugtehole 55720 (PERTH); 12 km N of Manangatang, A. C
55989 (PERTH).

South Australia: Andamooka, 4 Nov. 1929, y. «. Cleiand (AD): Lake Eyre North, G. C. Cornwall 5 (AD)'
Leigh Creek, T. R. N. Lothian 3028 (AD); Oakden Hills. B. J. Murray 512 (AD).

We-aern Australia: Lake Macleod, A. S. George 10192 (PERTH); West Is., Lacepede Islands, R. Johnson
(PERTH); 13 mi E of Broome, M. Lararides 6579 (BRl, CANB).

Northern Territory: Wessel Islands, P. K. Lutz 3503 (CANB); Mulga Park Stn., P. K. Latz 5036 (NT)'
5'6 mi S of Mt. Wedge HS., D. J. Nelson 104 (AD, NSW, NT).

Java: Bungil, April 1934, Altmann (K); Sumarang, Herb. Otto Kuntze no. 5476 (K); Japore, leg. Teysmann
(K).

Sumba Is.; Waingapu, tboet 23 (B, K).

Madura Is.; Sampang. C. A. Backer 19607 (K).

Timor: Bena-Panite. Sauveneur C!) L50(K).

Sub-species ieiostachya exhibits a multiplicity of forms and this explains to some
extent the several names which have been applied to it. Part of this plasticity is obviously
due to environmental factors since in different seasons the same plant may produce both
branches and spikes which are very different in morphology. Part of the variation is
certainly due to genetic differences since in cultivation some variants retain a distinctive
appearance. Noripally only one variant is present in any locality but occasionally, due
to introgression with subsp. bitiens or subsp. iiulica, a range of forms may be found.

The Malaysian plant, which was named A. ciliolatuni by Ungern-Sternberg, has
slender branches and slender, evenly cylindrical spikes; the leaf-lobes and bracts are
distinctly ciliolalc. When dry it usually assumes a brown colour, unlike the grey or
fawn colour of typical H. iiulica ssp. ieiostachya. The bracts when mature are more
leathery than corky and do not enlarge with age. In Malaysia this plant is invariable
and. in fact, is the only member of the genus present. A plant very similar to the Malaysian
has been collected on the north coast of Australia but here it obviously introgresses with
subsp. ieiostachya. I have therefore referred all material of this complex to this sub-species
but realise that from the point of view of Malaysian botanists it would be more inforimative
to recognise this plant as a species distinct from H. iiulica.

The lectotype of A. Ieiostachya (from Kyejeron Creek) was stated by Bentham (1870)
to have been collected on M'Douall Stuart's Expedition; it was, however, evidently collected
by J. P. Murray (whose name is on the herbariuin label), probably in 1862 on a Victoria
Exploring Expedition relief parly led by A. W. Howitt (see J. H. Willis, 1962). I have
chosen this collection as the lectotype since it is typical of the eremaean variant of the
sub-species; of the other collections cited by Bentham, those from Providence Hill and
McAdam Range represent the north coast variant (approaching A. ciliolatuni) while the
Drummond collection cited is a vegetative specimen of H. cloleiforniis.

23c. subsp. bidens (Nees) P. G. Wilson, stat. el comb. nov. Fig. 10, 31E-F.

Arlhrocnemiim bidens Nee.s In Lehm.. PI. Preiss. 1: 632 (1845); Moq. in DC., Prod, 13 (2); 151 (1849);
Ungern-Sternberg, Versuch Sy.st. Salicornieen 71 (1866). Salicornia bidens (Nees) Benth., FI. Austral.
5; 204(1870).

Type: Swan River, Preiss 1261 (iso: K, MEL).

Robust shrub to 2 m high. Articles thick, obovoid, circular in cross-section or
somewhat (lattencd, 5-10 mm long, deeply and often acutely lobed, keeled towards apex,
usually ciliolate. Spikes ovoid, pale brown to grey with age, the lowest article at first
larger and jnore deeply lobed than those above. Flowers fused to upper bract and to
eacTi other. Fruitlets: perianth soft and spongy; pericarp hard and horny all over. Seed
ovate; testa smooth and glossy.

68

Nuytsia Vol. 3 (1) 1980

Figure 10. Halosarcia iiidica subsp. hidens; near Forest Beach. South West ol Capel, Western Australia.
Photo by B. R. Maslin.

Habitat: Moderately saline areas both coastal and inland.

Distribution: North western Victoria, southern South Australia, Western Australia south
of 26 latitude and west of 123° longitude. Map 14.

Victoria: 10 mi NNW of Underbool, A. C. Beaitgiehok 40385 (PERTH).

South Australia: Grange, 19 Nov. 1926, J. B. Clclaud (AD); Goolwa Barrage. Hi. Eichler 17122 (AD);
N of Cowell, C. R. Atcock 895 (AD).

Western Australia: Wyola, T. E. H. Aplin 694 (PERTH); E of Mullewa, A. M. Ashhv 2591a (ADI; Phillips
River, E. M. Bennett 3123 (PERTH).

This sub-species is typically a sub-tropical plant and is relatively common in the
southern portion of Australia south of 26° latitude. It occasionally occurs in association
with subsp. leiostachya in more northern localities but here often appears to be a recent
arrival since it may be only represented by isolated bushes. Towards the north it grades
into subsp. leiostachya and with many collections it is not practicable to designate them
as belonging to either one sub-species or the other.

Sub-species hidens exhibits considerable plasticity of form; in particular the spike
may vary greatly in size, shape and texture. The size is much dependent on the environ-
mental conditions which when satisfactory cause the spike to become much longer than
normal. The corkiness of the spike increases with age but the texture is obviously also
dependent on other faetors.

The fully mature fruiting spikes break up in a manner similar to that described under
subsp. indica, but immature fruiting spikes (those which have not assumed a dull pale
fawn appearance) of both subsp. hidens and subsp. leiostachya will often swell up and
disarticulate when placed in water. This reaction is due to the gelatinous nature of the
cell contents of the bract tissue and the perianth. A similar reaction has not been observed
in any other species.

Paul Cl. Wilson, Australian Salicornieae

69

23d. siibsp. julacea P. G. Wilson, siibsp. nov. Fig. 62, 63.

Perennis decumbens; raniuli graciles. Ariiciili anguste cylindraeei, 4-10 mm longi, 2-3 mm diam.,
margine obtuse lobato, eciliato. Spicae cylindraceae; bracteae truncatae, integrae. Spicae fructificantes
eruciformes; perianthium medullosum, in piano sagittal! secedens; pericarpium ad apicem crustaceum
basin versus membranaceum. Semen sub-circulare, ca. 1 -5 mm diam: testa straminea, leviter reticulata.

Type: Near Derby (17 I7'S, 123 37 'E), Western Australia, July 1975, V. Semeuiuk 174005
(liolo: PERTH, iso: CANB, K).

Decumbent perennial. Braiichlets slender; articles narrowly cylindrical 4-10 mm
long, 2-3 mm diameter, margins shortly and bluntly lobed, eciliate. Spikes cylindrical
2-5 cm long. 4-5 mm diameter; bract-pairs truncate, entire. Flowers free from each
other but adherent to upper bract in pro.\imal half; perianth lobes 2, collateral, not over-
lapping. Fruiting spike cruciform with bulging fruitlets: perianth firm and pithy, eventually
separating into two lateral halves; pericarp crustaceous at ape.\, membranous below.
Seed sub-circular, ca. 1-5 mm diameter; testa stramineous, irregularly wrinkled, loosely
surrounding embryo and perisperm, with faint reticulate ornamentation.

Habitat: Tidal flats in tropics.

Distribution: Coastal tropical Australia. Map 9.

Queensland: Woody Is., H. Flecker 8962 (NSW); St. Lawrence, W. Macnae (BRI); Townsville Common,
24 Feb. 1977, P. G. Wilson (PERTH).

Western Australia: Pt. Wvndham, A. C. Beaiiglehole 47043 (PERTH); near Derby, V. Semenhik 174005-6
(PERTH).

Northern Territory: Darwin Harbour. BaHooy 1419 (PERTH); Peron Is., T. S. Henshall 855 PERTH);
Melville Is., G. Stocker 611 (NT).

Around the tropical coast of Australia in regions where either H. indiea subsp. indica
or the tropical coastal variant of subsp. leiostachya occur, is frequently also found a plant
which is similar to each of these sub-species but which differs from both in having the
flow'ers and fruitlets exposed, in having a thin pericarp, and in having a sub-circular seed
with faint porcate sculpturing. This plant varies considerably in habit and morphology;
it is predominantly associated in the field with one or other of these //. indica sub-species
and usually occupies an intertidal situation. Three of the collections studied are known
to have been growing in association w'ith one of the tropical coastal variants ol H. per-
granulata (as well as with H. indica), i.e. near Derby, on Sir Graham Moore island, and
at Townsville. At Derby and at Townsville, where it has been observed by the author,
it has the appearance of being a distinct taxon, but in both situations it grades imperceptibly
into the local variant of H^indica. Some of its organs are intermediate in morphology
between H. indica ssp. indica and H. pergramdata (e.g. in texture and ornamentation of
testa, texture of pericarp, and shape of articles). This suggests that the plant is a hybrid
between the two species. Flowever, in vegetative anatomy (i.e. in the single row ot palisade
cells and the underlying sheath of small thickened cells) it agrees with H. indica. At
Derby it occupies a distinct zone in the llalosarcia association but at Tow nsville it is found
growing intermixed with ssp. indica and H. pergranulaia, a difference that is due evidently
to a noticeable inclination of the mudfiat at Derby compared with an almost horizontal
tidal flat at Townsville. The mature seed appears to be normal except that the testa is
wrinkled, that is, it does not fit closely around the embryo and perisperm as is usual in
the genus. I have identified specimens of this complex as H. indica s,ubsp. Julacea but
further field w'ork over its whole range is required to understand its true relationship with
the other sub-species of H. indica and possibly with H. pergranulaia.

Hcdosarcia indica subsp. julacea differs from subsp. indica and leiostachya m having
slender eciliolate articles, in the texture and lobing of the perianth, the texture of the
pericarp (not horny) and in the shape of the seed and ornamentation of the testa. It
is probable that eventually a complete range of variability will be found connecting these
taxa but where subsp. julacea grows adjacent to subsp. leiostachya the two do not appear
to hybridise.

The specific epithet refers to the resemblance of the spikes to catkins (Julus, a catkin).

70

Nuytsia Vol. 3 (1) 1980

5. Sarcocornia A. J. Scott

A. J. Scott. Bot. J, Linn, Soc. 75: 366 (1978).

Saliconna subgenus Arthrocnemoides Ung.-Sternb., Versuch einer Systematik der Salicornieen 54 (1866).
Lectotype: Salicuniia frmicosa (L.) L,. fide A. .1. Scott, l.c.

Saliconim sect. Pevennes Duval-Jouve, Bull. Soc. Bot. France 170 (1868), n.v.. Moss, J. Bot. British &
Foreign 49: 178 (191 1). Lectotype: Saliconiia peremus Miller, fide A. J. Scott, l.c.

Arthrocnemum subgen. Gymnanthemum Moss, J.S. Afr. Bot, 20: 8 (1954).

[Salicontia auct. non L. s.str.: .1. M. Black, Trans. Roy. Soc. S. Austral. 43: 365 (1919); J. M. Black, FI.
S, Austral, ed. 2, 321 (1948); .1. H. Willis, ttandb. PI. Victoria 2: 110 (1972); Beadle. Evans and Carolin,
FI. Sydney Reg. 188 (1972).]

{Arthvocnenmm auct. non. Moq. s.str.; C. E. Moss. J.S. Afr. Bot. 20: I (1954); tt. R. Tolken, Bothalia
9; 255 (1967).]

Decumbent to erect perennial herbs or suh-shruhs often rooting at the nodes. In-
florescence a terminal spike-like thyrse consisting of 3-12-fiowered cymes sessile in the
axils of the bracts. Bracts united in opposite pairs, 1 shield shaped on outer surface.
Flowers apparently embedded in the spongy mesophyll of the succulent axis, hermaphrodite
or unisexual by abortion (rarely plants dioecious), arranged vertical to spike axis, free
distally from bracts and from each other, fused towards base. Perianth succulent,
gamotepalous; apex (outer surface) truncate, exposed, the orifice a short central vertical
slit; lobes 3 or 4, two long lateral lobes and a small semi-circular outer adaxial lobe (some-
times also with a small semi-circular abaxial lobe). Stamens 2, one on both the abaxial
and adaxial side of pistil. Ovary semi-circular to ovoid, succulent: style short; stigmas
2 (3), linear, papillose. Spike enlarging in fruit, the axis and bracts not disarticulating
at maturity. Fruiting perianth becoming dry and spongy: pericarp membranous, !
fused to perianth and both eventually .separating from axis just above their base. Seed
vertical, broadly ovate to orbicular; seed coat membranous of two separate layers, the
testa papillose or with slender (sometimes hooked) hairs: embryo hippocrepiform, radicle
incumbent, inferior; perisperm absent.

Type species: Sarcocornia perennis (Miller) A. J. Scott (Salicornia perennis Miller).

About 16 species throughout the world; three are native to Australia, two of which
are endemic.

Origin of name: From the Greek word sar.x, flesh, and the Latin word cornu, horn, referring
to the succulent nature of the plants, the shape of the leaves, and with allusion to the
genus Salicornia (cf. the name Pachycornia).

The genus Sarcocornia includes species which were referred to Salicornia by Ungern-
Sternberg ( 1 866, 1876), Volkens ( 1892), J. M. Black(19l9), Ulbrich ( 1934), and by authors
of recent Australian State floras. However, Moss (1954), Tolken (1967), and Ball (1964)
treated these species as belonging to the genus Arthrocnemum, a move that was accompanied
by a misinterpretation of both Standley’s (1916) and Ulbrich's (1934) lectotypification
of that genus. This lectotypification relates back to Ungern-Sternberg's critical assess-
ments of the genera within the Salicornieae. He recognised that Moquin's original
description of Arthrocnemum encompassed species belonging to disparate genera and he
effectively excluded all but three taxa. i.e. A. fruticosum var. macrostachvum (Moric.)
Moq. ( - A. glaucum (Delile) Ungern-Sternb.), A. indicum (Willd.) Moq.. and A. arhuscula
(R.Br.) Moq (the last species with a query). P. Standley (1914) pointed out the confusion
which existed in European floras over the use of the names Salicornia and Arthrocnemum
and loiter (1916) lectotypified the latter genus on A. fruticosum [non (L.) Moq.j Moq,
Is'-’totypification was accepted by Ulbrich ( 1934). Unfortunately
both Moss and Tolken apparently misunderstood the intention of the earlier authors
and (jonsidered the type to be A. fruticosum s.str., a move which ran counter to the intentions
ot Ungern-Sternberg, Standley, and Ulbrich (see also A. .1. Scott, 1978). The genera
Sahcornia jand Arthrocnemum as previously understood together contained five distinct
groups which are here recognised as separate genera, the additional ones being Sarcocornia,
Sclerostegia and Halosarcia,

Paul G. Wilson, Australian Salicornieae

71

The Australian members of the genus Sarcocornia can be readily distinguished from
those of the related genera by the truncate apex of the perianth in which the orifice is
surrounded by a pair of vertical (lateral) lobes and a small semi-circular outer adaxial
lobe (there may also be a small abaxial lobe); in no other Australian genus of the Sali-
cornieae is this adaxial lobe present. The llow'ers are also distinctive in having two
stamens (one in other Australian genera), and in having a seed in which the embryo is
horseshoe-shaped and albumen absent. In their vegetative morphology members of the
genus Sarcocornia are distinctive in ha\ing either plain or spirally thickened sclereids
in the palisade tissue of the stem, a character not present in the other Australian members
of the Salicornieae (fig. 3E, F). Additional characters which readily distinguish the two
common species of Sarcocornia from all other Australian members of the Salicornieae
are the number of llowers in a cymule (5 or more) and the decumbent habit with adventitious
roots arising at the nodes.

The genus Salicornia is similar in its floral morphology to Sarcocornia and differs
principally in: (1) containing only annual species; (2) all of the branches of the plant
terminating in an inllorescence; and (3) the shape of the (normally 3-nowered) axillary
cymes in which the lateral llowers are below the central and contiguous, their truncate
apices forming the shape of a triangle. This genus is not represented in Australia.

Key to the species of Sarcocornia

1. Decumbent or caespitose woody perennial often rooting at nodes; flow'ers 5-12

in each axillary cyme , .. .. .... .. 2

I*. Erect sub-shrub, not rooting at nodes; flowers in 3-flowered cymes; seed minutely
tuberculate over radicle, otherwise smooth 3. S. globosa

2. Flowers of cyme in single row; seed with slender acuminate hairs sometimes hooked

at their tips I- S. quinqueflora

2*. Central flowers of cyme often in two rows; seed papillose with short rounded
hairs 2. S. blackiana

1 . Sarcocornia quinqueflora (Bunge ex Ung.-Sternb.) A. J. Scott, Bot. J. Linn. Soc. 75:
368 (1978). Fig. 2A.

SuUconiia cjniiujiiefiora Bunge ex Ung.-Sternb., Vers, Syst. Salic. 59 (1866); Ung.-Sternb,, Atti Cong. Int,
Bot. Firenze 1874, 302(1876).

Syntvpes: “Herb. Steetz: Melbourne (gesam. von Hildebrand)! Port-Adelaide (Blon-
dowsky, Ferd. Muller) ! Port-Jackson (Rieder)!)". Lcctolypc: Port Adelaide, F. Mueller
(K, MEL 71432) lecto nov.

5. cjiixlralii- Solander ex Forst.f., Flor. Instil. Austral. Prod. 88 (1786) nomen.

5. australis Solander ex Bcnth., FI. Austral. 5: 205 (1870): H. H. Allan, FI. N. Zealand I: 232 (1961).
Lectotype: New Zealand. Forster (BM, n.v.), lecto nov.

Arthroenenmm heptiflorum Moss. ,I.S. African Bot. 20: 18 (1954), nom. illeg. non Moss ex Fourc. (1941).
Type: New Zealand, leg. Forster (BM, n.v.).

[5 iiiclica auct. non Willd.; R.Br., Prod. 411 (1810) p.pte.; Richard. Essai FI. Nouv.-Zel., 162 (1832) (in
Dumont d'Urville, Voy. Decouv. I’Astrolabe); Hook.f., FI. Tasm. I: 317 (I860); Hook.f., FI. Nov.-Zel.
216 (1853).]

[A. ausirulasiciini auct. non (Moq.) Moss: Moss, .I.S. African Bot. 20: 19 (1954), as to the New Zealand
and New- Caledonian specimens cited.]

Decumbent to erect perennial, frequently caespitose or, when decumbent, rooting
at the nodes, up to 0- 5 m high. Branches slender; articles cylindrical to narrowly obovoid,
5-15 mm long, minutely to prominently and acutely lobed, sometimes glaucous when
young, older articles often retaining shape due to the thick firm spongy mesophyll; uni-
formly thickened (or sometimes spirally thickened) sclereids frequently present in palisade

72

Nuytsia Vol, 3(1) 1980

tissue. Spikes (!) 2-5 cm long, cylindrical or narrowing towards the apex when in Mower,
4-5 mm diam. when in fruit; axillary cymules 5-9 Mowered in a single row, central Moret
(quadrate to) oblong or cuneate on outer face. Perianth (in fruit) ca. 2 mm high on outer
face; adaxial lobe small, semi-circular, outside the lateral pair; abaxial lobe minute,
triangular to semi-circular, usually within the lateral pair. Seed circular, covered with
short or slender acute hairs, appressed or raised (sometimes uncinate), prominent around
the periphery, frequently absent from the sides.

Nomenclatiiral Notes: The name Arthrocnemuni heptifiorum Moss (1954) was published
posthumously which could account for the confusion with its earlier homonym published
by Moss ex Fourcade in 1941, also after Moss’s death. Moss (1954) stated that the
name A. heptiflonim was a ‘nom. nov.'; in his synonymy he cited three names, viz. “S.
australis Sol. ex Forst. Prodr. 88. 1786 (nomcn nudum); S. imiica Hook. Handb. N.Z.
FI. 233. 1864 non. Willd. 5". pachystachya Black FI. S. Austral. 2. 208 f.l. 1924. non Bunge
ex Ung.-Sternb.". Moss* died in 1930 and the editors of his manuscript were evidently
unaware that Ulbrich (1934) had already provided a new name for S. pachystachya Black
and that Bentham (1870) had validly published the name S. australis (based on Solander’s
manuscript name). Moss, in calling his name a 'nom. nov.’, and in citing the type as
being the Forster specimen from New Zealand, was evidently intending a nomen novum
for 5. australis Sol. ex Forst. f., though it is unclear why he felt this required a new name.
There is the possibility that he was aware of Bentham's 'S. australis' and assumed it to
be based on an Australian plant. He was, therefore, providing a new name for the New
Zealand entity. In any case, the name A. heptifiorum (1954) is illegitimate being both
superMuous and a later homonym. Salicornia pachystachya Black is distinct from the
New Zealand species and is a nomenclatiiral synonym of Sarcocornia hlackiana. In
his paper Moss also cited specimens from South and East Africa; these are referred by
Tolken (1967) to A. decumhens Tolken ( Sarcocornia decumhens (Tolken) A. J. Scott).

The name A. australasicum was misapplied by Moss (1954) to at least two dilferent
species. The New Zealand and New Caledonian specimens cited belong to S. cpdnqueflora
while the African specimens belong to Sarcocornia decumhens (Tolken) A. J. Scott (see
Tolken, 1967). The type of A. australasicum was stated by Moss to be a Robert Brown
specimen from the East Indies in herb. BM; the type is, however, a specimen of unknown
collector in herb. P; the name has been transferred to Tecticornia as T. australasica (Moq.)
P. G. Wilson, a species quite different from those considered by Moss (see Wilson, 1972).

Robert Brown applied the name Salicornia indica Willd. to the species considered
here and his application was followed by both Richard and J. D. Hooker. Moquin
(1849) recognised a misidentification by Brown and considered that the latter’s specimens
probably belonged to Halocnemum australasicum Moq. However, .1. D. Hooker (1853,
I860), who evidently discussed the matter with Brown, adhered to a broad (and obviously
incorrect) concept ot the species. Brown (1810) appears to have based his description
of 5. indica on material of Sarcocornia ? quincpieflora from Fowlers Bay, South Australia
(cf. his Flora manuscript at the British Museum), but his cited distribution of the plant
encompassed specimens or observations of several diM'erent species.

Key to subspecies

Plant decumbent (or shrubby); spikes slender; seed with slender often uncinate
trichomes subsp. quinqueflora

Plant caespitose, spikes shortly cylindrical; seed with short acute trichomes

lb. subsp. tasmanica

l.rt hv M c prepared by his wife and b> Prof. R. S, Adamson from notes

lett by Moss after his death m 1930. These editors added to and altered the original manuscript (including
the synonymy citation); many of the contradictory statements evident in this paper could have emanated
irom these later additions.

Paul G. Wilson, Australian Salicornieae

73

la. subsp. quinqueflora. Fig. 33A-B, 68, 69.

S. australis Solander ex Benth., I.c.

S. heptiflorum Moss, I.c,

Decumbent to erect perennial rooting at the nodes, to 0-5 m high. Spikes 2-5 cm
long, narrowly cylindrical; central floret oblong to cuneate on outer face. Seed with
slender acute (often uncinate) hairs.

Habitat: Moderately saline situations which are frequently inundated, e.g. estuarine
localities.

Distribution: Coastal or near coastal situations around the east and south coasts of Aus-
tralia, and on the west coast as far north as Carnarvon; inland in south-west Western
Australia: also found in New Zealand and New Caledonia. Map 20.

Queensland: Sandgate, S. T. Blake 2384 (BRl);. Hannibal Is., Gt. Barrier Reef, T. Done (BRl); Stradroke
Is., C, E. Hubbard nif,

New South Wales: Narrabeen, D. O. Cross (NSW); Bermagui, W. Dunn (NSW); Nowra, 7 Feb. 1956,
E. Gauha (BRl, NSW. PERTH).

Victoria: Port Phillip Bay, H. I. Aston 269 (MEL); Wingan Pt., A. C. Beauglehole 32662 (PERTH); Williams-
town, R. J. King 4626 (MEL).

Tasmania: Sorell, Dec. 1908, R. A. Black (MEL); Eaglehawk Neck, N. T. Burbidge 3221 (HO); King Is.,
Nov. 1887, C French (MEL).

South Australia: Lake Robe, J. R. Dodson 139 (AD); Ethelton, 9 Feb. 1918, E. H. Ising (AD); 30 km N
of Ardrossan, A. E. Orchard 2790 (AD),

Western Australia: Swan River, Perth, Mav 1902, W. V. Fitzgerald Lake Coyrecup, K. Newbey

3156 (PERTH); Busselton, A. & E. Pries (MEL).

Morphological Notes. The presence of sclereids in the palisade tissue of the stem has
been commented on by a number of workers. Cooke (1912) recorded ‘spiral tracheides’
in material which presumably came from New Zealand. Moss (1954) commented on
them but owing to a taxonomic confusion it is not clear to which species he was referring.
Tolken (1967) recorded spiral sclereids in S. c/uimpieflora but in this case it is not clear
whether from his own observations or whether he was relying on those of Cooke. I
have examined specimens from all Australian States and from New- Zealand and have
often been able to find sclereids. When present, however, they have been uniformly-
thickened except in some specimens from Western Australia and from South Australia
where spirally thickened sclereids were found (see Fig. 3E-F). In no plant examined
were both types of sclereids present (although when using dried herbarium material they
may have been overlooked) and Tolken (1967) found that in the South African species
of Sarcocornia (which was treated under Arthrocnemum) one or other type was typical of
a particular species. If this situation is paralleled in Australia it would suggest that more
than one species should be recognised within the complex now referred to S. c/uinqueflora.
It has not, however, been possible to consistently correlate the presence or absence of
spiral sclereids with other characters, nor has it been possible to demonstrate the presence
of sclereids in the majority of dried herbarium specimens, therefore at present no attempt
has been made to erect additional taxa on this anatomical data. Specimens of 5. quin-
cpieflora in which spiral sclereids were observed are as follows;

South Australia: S. Neptune Is., Jan. 1923, T. G. B. O.v/irt/v/ (NSW); Flinders Is., 7 and 8 Jan. 1924, T. G. B.
Osborn (NSW); N. Pearson Is., 10 Jan. 1923, T. G. B. Osborn (NSW).

Western Australia: Lake Hope, G. J. Keigherv 1520 (PERTH); W of Moora, 27 Apr. 1969, J. & D. C.
Lowrv (PERTH); Shark Bay, 9 Apr. 1972, C. Stone (PERTH); Carnarvon, P. G. Wil.son 8352 (PERTH);
New Beach, ca. 30 km S of Carnarvon, T. & J. Whaite 4188 (PERTH).

74

Nuytsia Vol. 3 (I) 1980

Tolken (1967) noted that the abundance of sclereids in the palisade tissue was often
related to the salinity of the soil; the greater the salinity the more numerous the tracheids.
I have not observed this correlation in the Australian species where relative abundance
appears to be determined by geography rather than ecology.

Tolken (1967) recognised that S. t/uinqueffora did not occur in southern Africa, as
had been stated by Moss (under S. australis) and he described the African segment of
the 'australis' complex as a new species, Arthrocnemum clecumhcns. Tolken compared
A. decumhens with both S. quinqueflora and S. hlacklana and stated that the latter two
species had spiral sclereids whereas in A. decumhens they were uniformly thickened. As
noted above, spiral sclereids do occur in a variant of 5. quinqueflora, but this variant has
not been recorded from near Adelaide, Melbourne, or Sydney and since plants from these
areas tend to be more slender than those in which spiral sclereids occur it is unlikely that
such will be found. S', quinqueflora was described from material that was collected near
these cities and it is therefore probable that none of the syntype specimens have palisade
tissue with spirally thickened sclereids.

Collectors have sometimes indicated that specimens were dioecious and Black (1919)
found S. quinqueflora to be functionally so, the male plants having ovaries which were
sterile and smaller than those of the female plant. A study of herbarium material suggests
that the plants of some localities are dioecious with the male flowers as described by
Black and the female with small staminodes, however, in general specimens bear what
appear to be hermaphrodite flowers. It is not clear how this sexuality is related to the
plant's distribution or whether it is dependent on environmental factors. (For a general
discussion on this phenomenon see J. Heslop-Harrison, 1957, 1972.)

Ecology. The ecological conditions which generally operate in segregating 5. quinqueflora
and 5. hiackiana are not clear. Both species will grow in situations which are periodically
inundated, with S. quinqueflora generally occurring where the water has a salt content
roughly equivalent to or less than that of the sea; Sareocornia hiackiana, when allopatric
with the former species, often grow's in areas which are drier for much of the year or are
likely to have a higher salt content; it will also grow in saline loam which is very dry in
the summer and here it adopts a bushy rather than decumbent habit. Where the two
species grow together they tend to be virtually identical in gross morphology. It is
interesting to note that in Western Australia the variant of 5. quinqueflora which has spiral
sclereids is found growing in conditions where water stress is likely to be at times extreme;
this anatomical character may therefore be an advantageous adaptation to these situations.

lb. sLibsp. tasmanica P. G. Wilson, subsp. nov. Fig. 33C, 72.

caespitosa ad 20 cm alta; caules graciles dense aggregati; articuli prominente lobati, Spicae
breviter cylindraceae ca. 1 5 mm longae 4 mm lalae. Periamhinni in fructo firnie spongiosum, apice quadrato
ca. 2 mm lato. Semen cum Irichomatibus brevibus acutis ornata.

Type: Bond Bay. Port Davey, Tasmania; rocks on sea edge subject to heavy spray at
times, 14 March 1954, M. Davis 1306 (holo; MEL 71455, iso: HO).

Plant caespitose or with a dense cluster of slender stems to 20 cm high arising from
short horizontal stolons. Branches slender, the aqueous tissue collapsing with age and
soon shed along with the outer tissues. Articles prominently lobed. 5-10 mm long.
Spikes shortly cylindrical ca 15 mm long, 4 mm wide. Fruiting perianth firmly spongy;
apex quadrate, ca. 2 mm wide. Seed orbicular, ca. I -3 mm diameter, covered with short
acute trichomes.

Habitat: Rocky coasts, usually in situations where the plants are subject to sea-spray
or to periodic inundation by the sea.

Paul G. Wilson, Australian Salicornieae

75

Distribution: Tasmania and probably the south coast of Victoria. Map 21.

Victoria: Mt. Martha, L. A. S. Johnson (NSW 136621).

Tasmania: Pt. Davey, 9 Feb. 1937, C. Davies (NSW); Blythe River, 3 Jan. 1937, G. & C. Davies (NSW);
Cape Portland, 1884, Baiulinel (MEL); Breaksea Is., M. Davis 1341 (CANB, HO); Cape Wickham, King
Is.. March 1885, Dobson (MEL); Southport, S. Jacobs 2027 (NSW); Bicheno, V. Jacobs 7 (MEL); Cape
Barren Is,, J. S. Whinray 422 (MEL).

It is not clear whether the characteristic caespitose form adopted by this subspecies
is due to the influence of its preferred habitat (rocky coasts) or whether it would in other
situations adopt this habit; however, in Tasmania plants with this habit and with slender
stems (from which the soft tissues have been shed) have, if in fruit, always been found
to have seeds with short acute trichomes. The subspecie.s tcisnuinica has often been
collected along with a larger decumbent plant which is here recognised as subsp. quin-
queflora and specimens of this latter subspecies, if in fruit, have usually been found to
possess seeds with slender hooked trichomes, although in some cases a variety of seed
types has been found in the one inflorescence. It is not clear whether such plants, having
seeds of different types, are hybrids or w'hcther the seed indumentum (and perianth shape)
is influenced by the environment, although the latter possibility seems unlikely.

Specimens have been collected which appear to be intermediate in gross morphology
between the two subspecies but these do not possess fruit so unequivocal identification
is not possible. Because of the difficulty in assigning non-fruiting material to one or
other taxon I have used the rank of subspecies rather than species, although, in fact,
typical members of the two plants appear to differ sufficiently from each other to deserve
the higher rank.

Sclereids in the palisade tissue have been observed in one specimen ( K Jacobs 7,
MEL 71420); they were evenly thickened as is usual for the S. quinquefiora complex.

The solitary collection cited of subsp. tcisnuinica from Victoria is not in fruit so the
record requires confirmation.

2. Sarcocornia blackiana (Ulbrich) A. J. Scott, Bot. .1. Linn. Soc. 75; 369 (1978). Fig.
33D-E, 70, 71.

Salicornia blackiana Ulbrich in Engler et Prantl, Nat, Pllanzenfam. ed. 2. 16c: 552 (1934), based on succeed-
ing.

Salicornia pachystachya J. M. Black, Trans. Roy. Soc. S. Austral. 45: 8, t.2 (1921) nom. illeg. non Ung.-
Sternb. ( 1 866).

Lectotype: Patawalonga River, Glenelg, 20 Jan. 1920,7. M. Black (holo: AD, ?iso: NSW)
lecto. nov.

Arlhi'ocnemnin heptifiormn Moss ex Fourcade, Bot. Surv. S. Air. Mem. no. 20: 19, 1 15 (1941) nom. illeg.
(superfluous name), based on above.

[A. heptifionim auct. non Moss (1954) sensu lectotypica: Moss, J. S. Afr. Bot. 20: 18 (1954) p.pte., as to
S. Australian specimen cited.]

Erect or decumbent perennial, to 0-8 m high, frequently rooting at nodes. Articles
to 10 mm long, narrowly obovoid to cylindrical, apex very shortly lobed; leaf-sheath
persistent and retaining shape due to the dry spongy mesophyll; palisade tissue with
numerous spirally thickened sclereids. Spikes 2-5 cm long, cylindrical, ca. 8 mm diam.
in fruit; axillary cymules 5-13 (lowered, frequently with the central florets in two tiers,
central lloret wedge-shaped on outer surface (apex). Perianth (in fruit) ca. 3 mm high
on outer surface; adaxial lobe small, semi-circular, overlapping the lateral lobes; abaxial
lobe minute, usually within the lateral lobes. Seed circular, testa covered with spreading
papilliform hairs (rounded at their tips) which are prominent around the periphery but
small on the faces.

76

Nuytsia Vol. 3 (1) 1980

Hahilat: Swampy or periodically waterlogged moderately saline soils, frequently in
estuarine situations. For notes on ecology see under S. quinqucflora.

Dislribulion: Coastal areas of Tasmania, Victoria and South Australia, in Western Aus-
tralia coastal as far north as Shark Bay, and inland south-western region. Map 22.

Viclorici: Allona. Port Phillip Bay, H. I. Aston 302 (MEL); The Lakes Nat, Park, A. C. Beauglehote 37748,
Corner Inlet, Sandy Is., Sept. 1952, C. I. Skcwes (MEL).

Tasmania: Ralphs Bay. F. H. Long 136 (HO); Muddy Plains. L. Railway 660 (HO),

South Australia: 9 km S of Cape Banks, 25 Feb. 1945, J. B. ClelamI {\D)\ 3 km N of Elliston, Hj. Eicliler
19448 (AD); Goolwa. 25 Aug. 1970, E. H. Ising (AD),

IVeslern Ansiralia: Middle Doubtful Is., Apr. 1977, /. Abbot (PERTH); Garden Is., T. E. H. Aplin 646
(PERTH); King George Sound, R. Brown (3080) (K); N of Stirling Range. Oct, 1867, F. Mueller (MEL);
2 km NW of Pt. Charles, Fitzgerald River, K. Ncwbey 4030 (PERTH).

Nomenclaiural Nates. C. E. Moss (1954) cited the collection 'Black 8’ from Gleneig,
South Australia, as being the type of Salicornia pachystachya and he listed this name as
a synonym of Artliracneniiim hcptifiontm. A specimen in herb. AD, collected by J. M.
Black on 20 January 1920 at Gleneig probably corresponds to this and has been selected
as lectotype. It bears a note in Black's handwriting to say that it was “sent to Paris as
no. 3, 30 1 20”; the no. 8 referred to in Moss’s paper may be a copying error for a “3”
on a specimen in herb. P. For additional notes on A. heptifiontm Fourcade (1941) and
A. hepliflonim Moss (1954) .see under S. quinqiteflora.

Morphological Notes. Surcoeonha blackiaita is rather uniform in appearance throughout
its geographical range (but it becomes slightly more robust towards the north-w'estern
limit). The papillae on the testa of the seeds provide the clearest character for distinguish-
ing it from S. qiiinque/lora and, in fact, from other members of the genus. In Victoria,
Tasmania, and South Australia the papillae are short and rounded, whereas in Western
Australia, in addition to plants which exhibit this typical form, may be found those in
which the papillae are slender and tend towards the shape of the irichomes found on
the seed of S. quinqueflora.

In all specimens e.xamined the palisade tissue had abundant spirally thickened sclereids
(fig. 3F) which, in material from much of its range, are useful in distinguishing this species
from S. quinqueflora (but see comment under the latter). The central florets of the
cymules in S. blackiana are frequently in two tiers although the flowers of the lower tier
may be vestigial. Old spikes from which the flowers have fallen still bear an indication
of the two tiers in the concavities of their axes. In the extreme western limit of its range
the single-tier condition is normal.

3. Sarcocornia globosa P. G. Wilson, sp. nov. Fig. 33F-K.

Fritticulus eiectu.s ad 20 cm altus. Articiili obovoidei ca. 5 mm long!, apice aeuminato, margjne
integro; textura vallo .sclereidis spiralibus numerosis. Iiiflorescentlae terminales, articulis sphaericis;
cymulae contiguae, 3-florae. Perlanthlam 3-lobum, lobo adaxiale parvo, hemisphaerico (lobo abaxiale
absent! ). Semen siib-orbiculare, ca. 2 mm diam,; testa super radicula minute luberculata aliter laevis.

Type: One Mile Rocks Reserve, 33 12'S. II9"46'E, 12 Nov. 1970, A. S. George 10502
(liolo: PERTH).

Small erect woody perennial to 20 cm high. Articles obovoid, ca. 5 mm long, apex
acuminate, margin entire, palisade li.ssue with abundant slender spirally thickened sclereids.
Spikes terminal to branches, of one or two (rarely three) spherical articles ca. 6 mm diam-
eter; bracts with outer face deltoid; axis slender, extending as prominent vertical partitions
between the flowers; cymule 3-flowered, contiguous with that of opposite bract. Flowers
hermaphrodite (stamens not always maturing), vertical to spike axis, free from each other
and from bracts but united in proximal half to radial partitions of axis; apex oblong in
outline (laterally compressed), surface curved, ca. 5 mm high, 3 mm wide. Perianth

Paul G. Wilson. Australian Salicornieae

77

succulent, 3-lobed; lateral lobes straight, the outer one extending along two-thirds
of perianth apex, the inner one for one-quarter length of apex; upper (adaxial) lobe small,
semi-circular, overlapping the lateral lobes; abaxial lobe absent; orifice small, in centre
of apex. Ovary thin walled, translucent; stigmas 2, delicate, sub-plumose. Fruiting
perianth dry and spongy, readily separating (with seed enclosed in persistent pericarp)
from spike axis; pericarp somewhat firmer than, and loosely adherent to, perianth. Seed
loosely adherent to pericarp, vertical, sub-circular, ca. 2 mm long; testa thinly chartaceous,
minutely tuberculate over radicle otherwise smooth, very pale brown; embryo accumbent,
cotyledons broad.

Distribution: Inland south-western Western Australia from Meckering east to Lake King
district. Map 21.

Western Australia: Ml. Caroline, 1891, O'. Sewell (MEL 71532); One Mile Rocks Reserve (33 12'S, 119”
46'E), A. Weston 7075 (PERTH); 3 km E of Meckering, P. G. Wilson 10965 (PERTH).

Sarcocornia glohosa may be readily distinguished from all other members of the
genus by its spherical (and usually solitary) fertile articles. From the other two species
found in Australia it differs in its erect habit and in its seed which is only sparsely tuber-
culate. The perianth lobes are similar in form to those found in S. hlackiana and S.
quincjnefiora except that the abaxial lobe (which is small in these two species) is in S. glohosa
totally absent, and the inner of the two lateral lobes is much shorter than the outer. The
palisade tissue is of .several cells thickness (as is normal for the genus) and contains
abundant slender sclereids which are spirally thickened; these sclereids extend from the
epidermis to the water tissue.

Whereas both 5". hlackiana and S. quinciueflora are predominantly species of saline
waterlogged situations (they even flourish in water), S. glohosa grows in drier situations
in company with such species as Halosarcia pergranulata and H. lylei. Its preferred
habitat is reflected in its habit, for whereas the other two species have the main branches
decumbent and rooting at the nodes, S', glohosa is an erect dwarf shrub.

Sarcocornia glohosa is almost certainly more widespread than is apparent from the
few collections seen, but it is a small plant and easily overlooked in a samphire shrubland.
However, it is not common, and at the .salt-lakes near Meckering (one of the three collection
sites) only one plant has been found even though the area has been extensively investigated
on several occasions.

Acknowledgements

The scanning electron microscope photographs were prepared or supervised by
Stuart Craig of the Division of Plant Industry, CSIRO, Canberra. I am grateful to him
for putting so much careful work into their preparation and to the CSIRO for permitting
them to be done.

The illustrations have been prepared by Margaret A. Menadue and Bryony V. Wilson,
to both of whom I extend my thanks for the long and detailed study they made of the
plants.

References

Agardh, J. G. 1858. Theoria Sysleniatis Plantarum. Lund.

Bentham, G. 1870. Salicornia. Flora Australiensis 5; 201 -205. London.

Black, J. M. 1919. A revision of the Australian Salicornieae. Trans. Roy. Soc. S. Austral. 43 ; 355-367.
Brown, R. 1810. Prodromus Florae Novae Hollandiae et Insulae Van Dieman. London.

Carolin, R. C., Jacobs, S. W. L., and Vesk, M. 1975. Leaf structure in Chenopodiaceae. Bot. Jahrb.
Syst. 95: 226-266.’

Corner, E. J. H. 1976. The Seeds of Dicotyledons. Cambridge.

De Fraine, E. 1912. The anatomy of the genus Salicornia. J. Linn. Soc. Botany 41 : 317-346.

78

Nuytsia Vol. 3 (I) 1980

Dumortier, B. C. 3. 1827. Florula Belgica. Tournai.

Falin, A. 1963. The fleshy corte.x of articulated Chenopodiaceae. J. Indian Bot. Soc. 42a: 39-45.

Fahn, A. and Arzee, T. 1959. Vascularization of articulated Chenopodiaceae and the nature of their
fleshy cortex. Amer. J. Bot. 46: 330-338.

Heslop-Harrison, J. 1957. The experimental modification of sex expression in flowering plants. Biol.
Rev. 32: 38-90.

Hindmarsh, G. .1. 1966. An embryological study of five species of Bassia Ail. (Chenopodiaceae).

Proc. Linn. Soc. N.S. Wales 90: 274-289 (1966).

Hooker, J. D. 1880. Chenopodiaceae. In G. Bentham and .1. D. Hooker, Genera Plantarum 3: 63-66.
London.

Linnaeus, C. 1753. Species Plantarum. Stockholm.

Mohabale, T. S. and Solanky, I. N. 1954. Studies in the Chenopodiaceae. II. Embryology of Arthroc-
nemum indicum Moq. Proc. Ind. Acad. Sci. 39 sect. B: 212-222.

Moss, C. E. 1954. The species of Arthrocnemum and Salicornia in Southern Africa. J.S. Afr. Bot.
20 : 1 - 22 .

Scott. A. J. 1978. Reinstatement and revision of Salicorniaceae J. Agardh (Caryophyllales). Bot.
J. Linn. Soc. 75: 357-374.

Standley, P. C. 1914. The genus Arthrocnemum in North America. J. Washington Acad. Sci. 4:
398-399.

1916. Chenopodiaceae, in North American Flora 21 : 1-93. New York.

Tolken, H. R. 1967. The species of Arthrocnemum and Salicornia in Southern Africa. Bothalia 9:
255-307.

Ulbrich, E. 1934. Chenopodiaceae. In A. Engler and K. PrantI, Die Natiirlichen Pfianzenfamilien ed.
2. 16c: 379-584. Berlin.

Ungern-Sternberg, F. 1866. Versuch einer Systematik der Salicornieen. Dorpat.

1876. Salicorniearum Synopsis. Atti Congr. Int. Bot. Firenze. 1874: 259-343.

Volkens, G. 1892. Chenopodiaceae. In A. Engler and K. PrantI, Die Natiirlichen Pfianzenfamilien
III. la: 36-91. Berlin.

Willis, J. H. 1962. The botany of the Victorian Exploring Expedition (September 1860-June 1861)
and of relief contingents from Victoria (July 1861-November 1862). Proc. Roy. Soc. Victoria 75:
247-268.

Wilson, P. G. 1972. A taxonomic revision of the genus Tecticornia. Nuytsia 1 : 277-288.

Figure 11. Pacln’coriiia triandra: A— Inflorescence; B— L.S. central hermaphrodite flower- C— L S
Iruit; D— Tnad of young flowers, lateral male, central hermaphrodite; E^T.S. perianth apex’ (a— stele
b -bract, c — perianth, d — stamen, e — style, f — ovule, g — seed). From J. Verschuer, 27 Aug. 1971.

Paul G, Wilson, Australian Saiicornieae

79

2mm

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Nuytsia Vol. 3 (I) 1980

Figure 12. Sderostegia iHsarticidata: A — Inflorescence; B — Triad of flowers (a — central hermaphrodite,
b-— lateral male); C — Seed; D — L.S. (radial) of fruitlet (c — stele, d — seed, c — thick pericarp, f — upper
bract, g — perianth, h — lower bract); E — L.S. of central flower (i — stele, j — fleshy ovary wall, k — ovule,
i — upper bract, m — perienth, n — style, o — anther, p — lower bract). A — D from P. G. Wilson 8603;
E from P. G. Wilson 8850.

Paul G. Wilson, Australian Salicornieae

81

1mm 2mm

Figure 13. Sckrostegia urhtisciilu: A— Inflorescence; B— Triad of flowers fused to bract (abaxial view),
lateral flowers male, central hermaphrodite: C— I..S. fruit; D— T.S. fruiting article. Sderostegia mamli-
formis: E— Fertile portion of branch; F— L.S. fruiting article; G— T.S. fertile article with young flowers,
lateral flowers male, central hermaphrodite; H— T.S. fruiting article (a— seed, b — pericarp, c— perianth,
d— bract, e- -stamen, f stele, g — ovary, h — ovule, i — male flowers, j— hermaphrodite flowers).

A to D from M. Brown. Lauderdale, Tasmania; E to H from A. S. George 9385.

91670 — ( 6 )

82

Nuytsia Vol. 3 (I) 1980

1mm

1mm

Figure 14. Sc/eroslegia tenuis: A — T.S, fruiting spike (a — bract tissue, b — secondary thickening, c — seed,
d — remains of perianth, e — pericarp, f — secondary thickening of pericarp; B — Triad showing the enclosed
anthers and stigmas emerging from central flower (abaxial view); C — ^Terminal inflorescence; D — Apex of
perianth with emerging stigmas (below, T.S. of apex); E — L.S. through fruiting spike (g — stele, h — staminal
filament, i — seed, j — thickened pericarp); F — T.S. immature fruiting spike (k — chlorenchyma, I — aqueous
tissue, m — stele, n — staminal filament, o — perianth, p — ovary wall, q — ovule).

A — F from M. A. Wilson 34; G from T. E. H. Aplin 2236.

Paul G. Wilson, Australian Salicornieae

83

Figure 15. Sclerostegia medullosa: A — Flowering branch; B — Portion of branch showing triads of flowers;
C — Triad of flowers (abaxial view), central hermaphrodite and lateral male flowers; D — Apex of article
showing denticulate margin ; E — Stamens; F — Seed; G — L.S. fruit (e — pericarp); H — L.S. flower (a perianth,
b — ovary wall, c — stamen, d — ovule).

From D. Symon 11313 (dried material).

84

Nuytsia Vol. 3 (1) 1980

Figure 16. Tegicornia imiflora, male plant: A — Flowering branch; B — Flower (abaxial view) with enlarged
view of perianth lobes; C — Apex of flower with exserted anther; D — Anther; E — Flower, lateral view;
F and G — Anther in bud; H — Stamen with pistillode (adaxial view); I - Pistillode; J. — T.S. flower (a —
pistillode, b — staminal filament, c — perianth),

From P. G. Wilson 1 1625.

Figure 17. Tegicornia imiflora, female plant: A — Flowering branch (a — flowers); B — Flower (abaxial
view) with enlarged view of perianth lobes; C — Flower dissected to show ovule and vascular trace; D —
Ovary with vestigial staminode ‘a’; E — Ovule; F and G — Fruiting perianth showing tangential position
of seed; H — T.S. fruiting perianth showing normal (radial) position of seed; 1 to 1C — ^Seed.

From P. G. Wilson 11626.

Paul G. Wilson, Australian Salicornieae

85

1mm

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Nuytsia Vol. 3 (1) 1980

Figure 18. Halosarcia sp. x Tegiconiia uniflora: A — Terminal inflorescences (a — united bracts, b — free
bracts, c — terminal solitary flower); B — Triad with a central fertile flower and a pair of vestigial flowers;
C — Anther; D — Terminal bract adaxial view); E — Triad with central hermaphrodite flower and lateral
female flowers (abaxial view); (d — stigmas, e — staminode, f — abaxial perianth lobe); F — Central flower
with seed ‘g’; G — Seed.

From G. Keighery, 18 Jan. 1978.

Paul G. Wilson, Australian Salicornieae

87

Figure 19. Hahsarda halocnemoides subsp. halocnemoides: A — Inflorescence; B — Central flower, abaxial
view (showing position of bract and inner medial lobe). Hahsarda halocnemoides subsp. catenulata:
C— Inflorescence; D — L.S. flower (a — bracts, b— perianth, c — ovary, d — medial lobe, e — staminal filament).
Hahsarda halocnemoides subsp. lennis: E— Inflorescence; F— Central flower, apex.

A and B from P. G. Wilson, 22 Feb. 1979, Alfred Cove; C and D from P. G. Wilson 8814; E and F from
M. Trudgen, Aug. 1978, Dampier.

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Nuytsia Vol, 3 (I) 1980

Figure 20. Hahsarcia fimhiiata: A— Terminal inflorescence (a— flower); B— Central fruitlet showing
seed exposed at torn base, abaxial view; C — The same, lateral view, Hahsarcia charlacea: D — Terminal
inflorescence; E — Fruitlet, abaxial view; F — Fruitlet, medial longitudinal section (a — seed, b — base of
staminal filament, c— perianth, d — pericarp).

A— C from D. and J. Lowry, June 1969: D— F from P. G. Wilson 7446.

Figure 21. Hahsarcia pergranulata subsp. pergraniilata: A — Branch bearing inflorescences; B — L.S.
through fertile articles; C — Fertile article with triad of flowers exposed; D — Seed.

From M. A. Menadue, October, 1974.

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Nuytsia Vol. 3 (I) 1980

Figure 22. Hdosam'a pcrgranulata subsp. pergranulata: A — Terminal inflorescence; B — Fruitlet, lateral
view, showing seed exposed at torn base of perianth; C — Fruitlet, abaxial view. Hahsarcia pergranulata
subsp. elongata: D — Terminal inflorescence; E — Fruitlet, abaxial view. Hahsarcia doleiformis: F —
Portion of spike; G — Fruitlet, abaxial view.

A— C from P. G. Wilson, Nov. 1978; D — E from J. Maconochie 1032; F — G from P. G. Wilson 8310.

Paul G. Wilson, Australian Salicornieae

91

1mm

Figure 23. Ha/osurcia lylci: A — Terminal inflorescence; B — Fruitlet, lateral view (a — hard protruding
style); C — Fruitlet, front view. Halosarcia lepidosperma: D — Terminal inflorescence; E — Fruitlet, front
view; F — Fruitlet, lateral view, showing seed exposed at torn base of perianth.

A — C from P. G. Wilson 6448; D— F from K. Newbey 3103.

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Figure 24. Halosarcia pteiygosperma subsp, denticulata: A— Terminal inflorescence; B— Central flower,
front view. Halosarcia fontinalis: C — Terminal inflorescence; D — Fruitlet, lateral view; E — Fruitlet,
front view. Halosarcia flabelliformis: F — Terminal inflorescence; G — T.S. spike to show fan-shaped
bracts and opposite triads of flowers (a — bract, b — perianth, c — pericarp, d — seed). H — Central floret,
apex.

A and B from T. E. H. Aplin, Bellefin Prong; C to E from D. E. Symon 9302; F — H from R. Chinnock
3364.

Paul G. Wilson, Australian Salicornieae

93

Figure 25. Hahsarcia calypiralu: A--Fruiting branchlet; B — L.S. through whorl of friiillets (f-
pericarp); C — L.S. flowering spike (a — style, b — anther). Halosan-ia leptudada subsp. lepioclada:
E Separate fruitlets. Hahsarcia leptoclada subsp. inclusa: F — L.S. flower showing included style (c-
d — staminode, e — bracts); G— Terminal inflorescence; H — Seed; 1— Triad of fruitlets (abaxial view)

—hard
D and
-style.

A — C from P. G, Wilson 10578; D — E from D. Wilcox 53; F — I from P. G. Wilson 1 1645.

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Nuytsia Vol, 3 (1) 1980

Figure 26. Halosarcia peltata: A — Terminal inflorescence; B — Central flowers, apex; C — L.S, through
young fruit (a — bracts, b — perianth, c — style, d — staminal filament).

From H. G. Baker, Lake Annean.

Paul G. Wilson, Australian Salicornieae

95

Figure 27. Halosaicia aitriciilcila: A — Inflorescence; B — Central Floret, abaxial view (a — lateral perianth
lobes); C — L.S. of fruitlet (b — upper bract, c — lower bract, d — style, e— seed with embryo and perisperm,

f pericarp.) Halosarckt pruinosa: D — Terminal inflorescence; E — Central fruitlet, abaxial view; F —

Lateral view of fruitlet within bracts (g— upper bract, h--staminal filament, i— lower bract, j— spike axis,
k — lateral wall of fruitlet),

A— C from G. I. Parlerliet, 13 May 1971; D— F from D. Lowry. 14 Aug. 1971.

Nuytsia Vol. 3 (1) 1980

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: -O (u
3 P 00

Paul G. Wilson, Australian Salicornieae

97

3mm

O-Smm

Figure 29. Hahsurcia iiiichilala: A — L.S. fruit; B — Fertile article from below to show the opposite triads
of flowers; C — Fertile branch with flowering spikes; D — Apex of perianth to show imbrication of lobes.
Halosarcia syncwpa: E — Spike. F — Central flower with portion of lateral flowers attached, G — L.S. flower
(a — perianth, b — stamen, c — pericarp, d - bract, c — ovule, f — stele).

A to D from P. G. Wilson I 1649; E to G from K. F. Kenneally, 13 Aug. 1970.

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Figure 30. Hatosarcia btilhosa: A— Vegetative branch; B — Flowering branch; C — Spike; D — Triad of
flowers; E — L.S. flower; F — Stamen and ovary; G — Ovary, H — Stigmata; I — Fruiting spike; J — L.S.
Fruiting spike; K— Fruitlet from above; L — Fruitlet from side; M — Fruitlet apex; N — Utricle; O — Seed;
P — L.S. fruitlet within bracts (a -pericarp, b — staminal filament, c — perianth, d — lower bract.)

A — H from M. Demarz s.n. (cult.); J— P from P. G. Wilson 1 1702.

Paul G. Wilson, Australian Salicornieae

99

4mm Sisj

Figure 31. Halosarcia enlrichoma: A— L.S. through flower (a— stamen, b— bracts, c— perianth, d— ovary);

Terminal inflorescence; C — Seed; D — Triad of flowers abaxial view. Halosarcia inJica subsp. bidens;

—Terminal inflorescence; F- L.S. fruitlet (e— spongy perianth, f- hard pericarp, g— staminal filament,
—seed).

A to D from R. Crantield 804; E and F from P. G. Wilson, Feb. 1978 (Swan River).

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Nuytsia Vol. 3 (1) 1980

02 mm

0-5mm

Figure 32. Halosarcia cupidiformis: A — Terminal inflorescence; B — Spike; C — Triad of flowers, abaxial
view; D — Seed and pericarp; E — L.S. through spike to show developing seed.

From P. K. Latz 518 (resuscitated specimen).

Paul G. Wilson, Australian Salicornieae

101

0*5mm

2mm

'TS

Figure 33. Sarcocornia qiiinqueflura: A— Portion of inflorescenre; B— Apex of flower, Sarmcornia
quinqiieflora subsp. tasmanica: C— Seed. Sarcocornia blackiana: D— Portion of inflorescence, E— L.S.
through fruiting spike (a— perianth, b— stamens, c— pericarp, d— seed).

Sarcocornia globosa: F-lnflorescence; G--Apex of flower; H-Side view of flower ; I-Seed;J- Embryo;
K— Section through embryo.

C from W. M. Curtis, March 1957; D and E from E. M. Bennett 2913; F to J from A. S. George 10502.

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Nuytsia Vol. 3 (1) 1980

Figure 34. Scleroslegia disarliciilala: SEM[photo of seed x 108. V. Setneniuk, 19 Aug. 1976.

Paul G. Wilson. Australian Salicornieae

103

Figure 35. Sclerostegia arbuscula: SEM photo of seed x 102. J. Parham, 28 Feb. 1976.

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Nuytsia Vol. 3 (1) 1980

Figure 36. Tegicornia iiiiiflora: SEM photo of seed x 76.

Paul G. Wilson, Australian Salicornieae

105

Figure 37. Tegicornia iiitiflora: Detail of Fig, 36 x 410.

106

Nuytsia Vol. 3(1) 1980

Figure 38. Halosarcia halocnemoides: SEM photo of seed .x 102. P. G. Wilson 8698, Alfred Cove.

Paul G. Wilson, Australian Salicornieae

107

3 ,# ,i^,w ^

Figure 39. Halnsania hahcnemoides: Detail of Fig. 3l< x 410.

108

Nuytsia Vol. 3 (I) 1980

Figure 40. Hahsarcia halocnemoides: SEM photo of seed x 128. P. G. Wilson 8650, Jibberding.

110

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Figure 42. Hcihsairia peruniiinlaia: SEM photo of seed x 115. P. G. Wilson 10106

Paul G. Wilson. Australian Salicornieae

111

Figure 43. Halosarcia pergranidata: Detail of Fig. 42 x 410.

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Nuytsia Vol. 3(1) 1980

Figure 44. Hahsarcia lepidospenna: SEM photo of seed x 102. R. A. Saffrey 6216.

Paul O. Wilson, Australian Salicornieae

113

Figure 45. Hahsarcia tepidospernui: Detail of Fig. 44 .\ 275.

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1 14

Figure 4fi. Hu/n^arcia ptc'ry!r(i.tperni(if,ub^p. SEM photo of seed x 104- N. Macfarlane 852.

115

Paul G. Wilson, Australian Salicornieae

Figure 47. Halnsarcia ptervgospcnna sulisp. plerygospcrnia: Detail of Fig. 46 \ 475.

116

Nuytsia Vol, 3 (I) 1980

Figure 48. Halosarcia lylei: SEM photo of seed x 129. A. S. George 8650.

Paul G. Wilson, Australian Salicornieae

Figure 49. Halosarcia lylei: Detail of Fig. 48 x 324.

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Nuytsia Vol. 3 (1) 1980

Figure 50. Hahsarcia flabelliformis: SEM photo of seed x 80. R. Chinnock 2984,

Paul G. Wilson, Australian Salicornieae

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Paul G. Wilson, Australian Salicornieae

121

Figure 53. Hahsarcia pruinosa: SEM photo of seed x 108. T. E, H. Aplin 3258.

122

Nuytsia Vol. 3 (1) 1980

Paul G. Wilson, Australian Salicornieac

123

Figure 55. Ha/oMiria oiirictilaia: Detail of Fig. 54 x 378.

124

Nuytsia Vol. 3 (Ij 1980

Figure 56. Halasarciu nndiilala. SEM photo of seed x 128. P. G. Wil.son 88.30.

125

Paul G. Wilson, Australian Salicoi’nieac

Figure 57. Halosarcia chartacea: ShM photo of seed x 108. ,1. HIkington 62«.

126

Nuytsia Vol. 3 (I) 1980

Figure 58. HalMairiti calvplnilci. SFM photo of.seed \ 128. P. G. Wilson 8939.

Paul Ci. Wilson, Australian Salicornieae

127

128

Nuylsia Vol. 3 (I) 1980

Figure 60. Halosarcia lepinclada: SEM photo of seed x 108. B. R. Maslin 182.

Paul G. Wilson, Australian Salicornieae

129

Figure 61. Hahsan iu indica stibsp. iiidiiu: SEM photo of seed .x 76. P. G, Wilson, 24 Feb. 1977, Towns-
ville.

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Nuytsia Vol. 3 (1) 1980

Paul G. Wilson, Australian Salicornieae

131

Figure 63. Halosarcia ituHca subsp. jiilacea: Detail of Fig. 62 x 405.

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Figure 64. Tecticoniia australasica: SEM photo of seed x 76. Kalkman, BW6226, Merauke', New Guinea.

Paul G. Wilson, Australian Salicornieae

133

Figure 65. Tecticornia australasica: Detail of Fig. 64 x 275.

134

Nuytsia Vol. 3 (1) 1980

Paul G. Wilson, Australian Salicornieae

135

Figure 67. Haloswcia pluriflora: SEM photo of seed \ 118. C. W. Bonython, May, 1951, Lake Eyre,
ADW7194.

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Nuytsia Vol. 3 (I) 1980

Figure 68. Sarcocornia quinqueflom subsp. quinqueflora; SEM photo of seed x 108. K. Newbey 3156.

Paul G. Wilson, Australian Salicornieae

137

Figure 69. Sarcucurnia quinqueflora subsp. qiiinqiiejlora: Detail of Fig. 68 324.

138

Nuytsia Vol, 3 (1) 1980

Figure 70. Sarcocornia blackiana: SEM phoio of seed x 108. K. Newbey 4030.

Paul G. Wilson, Australian Salicornieae

139

Figure 71. Sarcocornia b/ackiana: Detail of Fig. 70 x 324.

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Nuytsia Vol, 3 (I) 1980

Figure 72. Sarcocornia quinqueftora subsp. tasmanica: SEM photo of seed x 102. J. S. Whinray 422.

Paul G. Wilson, Australian Salicornieae 141

+ +

142

Nuytsia Vol. 3 (1) 1980

143

Paul G. Wilson, Australian Salicornieae

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Nuytsia Vol. 3 (1) 1980

Paul G. Wilson, Australian Salicornieae 145

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Note added in proof

The type species of Arthrocnemum is stated earlier in this article to be A. glaucum
(Delile) Ung.-Sternb. (1876); however, the epithet glaucum dates from 1876 since the
combination is based on Salicornia glauca Delile (1813), an illegitimate name being a
later homonym of S. glauca Stokes (1812). Some recent authors (see synonymy) have
therefore used for the same species the name Arthrocnemum macrostachyum (Moric.)
C. Koch (1853) which is based on S. macrostachya Moric. (1820); however, according to
Index Kewensis (1895) an earlier synonym exists, viz. Salicornia mucronata Lag. (1817),
a name which has never been accepted by botanists nor has it been transferred to Arthroc-
nemum. Ungern-Sternberg (1876) appears to have been the first person to suggest the
synonymy of 5. mucronata with A. glaucum although he did so with a query, which was
understandable since the description of the former species was based on vegetative material.
Index Kewensis (1895) provided the same synonymy and Moss (1911) stated that he was
following that work. It is likely that the later authors did likewise since, in each of those
papers cited above in which the name Salicornia mucronata is mentioned (except for that
of Ungern-Sternberg), the page reference for the original publication is incorrectly given
as 58 instead of 53.

1 am not transferring Salicornia mucronata to Arthrocnemum in view of the lack of
evidence that any worker since Ungern-Sternberg has looked into its correct application.
Until such a study is undertaken it would appear desirable that the name applied to the
species in question should be Arthrocnemum maerostachyum. The relevant synonymy
is as follows:

Arthrocnemum macrostachyum (Moric). C. Koch, Hortus dendrologicus 96 (1853); Zohary,
FI. Palaest. 1:156 (1966); Jafri et Rateeb, FI. Libya 58 : 51 (1978). — Salicornia macrostachya
Moric., Flora Veneta I: 2 (1820) n.v. — A. fruticosum (L.) Moq. var. macrostachyum
(Moric.) Moq., Chen. Mon. Enum. 112 (1840).

? Salicornia glauca J. Stokes, Botanical Materia Medica 1: 8 (1812), non Arthrocnemum
glaucum Ung-Sternb..

Salicornia virginica Forsskal, Flora aegyptiaco-arabica 2 (1775) nom. illeg. non L. (1753). —
S. glauca Delile, Florae aegyptiacae illustratio 49 (1813) based on preceding, nom. illeg.
non Stokes (1812); Moss, J. Bot. Brit. & For. 49: 177 (191 1). — A. glaucum Ung.-Sternb.,
Atti Congr. Bot. Firenze 1874: 283 (1876); Ball in Tutin et al., FI. Europ. 1: 101 (1964);
A. J. Scott, Bot. J. Linn. Soc. 75: 370 (1977).

? Salicornia mucronata Lag., Mem. PI. Barrilleras 53 (1817); Lag. in G. A. de Herrera,
Agricultura general 1 : 280 (1818) n.v.

Paul G. Wilson, Australian Salicornieae

153

Index to Names

Entries in bold type are of new taxa; entries in roman type are accepted names; entries in italics are of
synonyms. Page numbers in bold type refer to the main taxonomic treatment.

Arthrocnemum

subgen. Angianthemum Moss
subgen. Cymnaiilhenuon Moss
sect. Leiospenna Black
sect. Tracbysperma Black
arhusciila (R.Br.) Moq.
austraiasicum (Moq.) Moss
belangerianum Moq.
benthamii Paulsen
hidens Nees

brachystachyum Paulsen

ciliolatiim Ung.-Sternb.

donaldsonii (Ewart et White) C. A. Gardner

fniticosum (L.) Moq.

glaiicum (Del.) Ung.-Sternb.

holocnemoides Nees

var. pergramdaliim Black
var. ptergyospernmm Black
heptiflomm Fourc.
heptiflorum Moss.
indicunt (Willd.) Moq.
leiostachyum (Benth.) Paulsen
lylei (Ewart et White) Black
macrostachyum (Moric.) C. Koch
pndnosiim Paulsen
subterminale (Parish) Standley
triandrum F. Muell.

Halocnemum

arbiiscida (R.Br.) F. M. Bailey
Halosarcia

auriculata P.G. Wils.
bulbosa P.G. Wils.
calyptrata P.G. Wils.
chartacea P.G. Wils.
cupuliformis P.G. Wils.
doleiformis P.G. Wils.
entrichoma P.G. Wils.
hmbriata P.G. Wils.
flabelliformis P.G. Wils.
fontinalis P.G. Wils.
halocnemoides (Nees) P.G. Wils.
sub.sp. caudata P.G. Wils.
subsp. catcnulata P.G. Wils.
subsp. longispicata P.G. Wils.
subsp. tenuis P.G. Wils.
indica (Willd.) P.G. Wils.

subsp. bidens (Nees) P. G. Wils.
subsp. julaeca P.G. Wils.
subsp. leiostachya (Benth.) P. G. Wils.
lepidospcrma P.G. Wils.
leptoclada P.G. Wils.

subsp. inclusa P.G. Wils.
lylei (Ewart et White) P.G. Wils.
nitida P.G. Wils.
peltata P.G. Wils.
pergranulata (Black) P.G. Wils.
subsp. divaricata P.G. Wils.
subsp. elongata P.G. Wils.
subsp. queenslandica P.G. Wils.
pluriflora P.(j. Wils.
pruinosa P.G. Wils.
pterygosperma (Black) P.G. Wils.

subsp. denticulata P.G. Wils.
syncarpa P.G. Wils.
undulata P.G. Wils.

91670— (1 1)

12 , 28 , 70
28

70
28
28

14

71
13
66
67
66

66 , 67
22

13 , 14

8 , 13 , 14 , 152
29 , 31
40

48
75
71

13 , 63
66

49

4 , 152
54

8 , 13

15

12

20

28

55 , 122 . 123

59 , 95 , 98 , 144
62 , 93 , 126 , 127
61 , 88 , 125 , 146
61 , 100 , 141
46 , 142

60 , 145

37 , 88 , 99 , 143

15 , 52 , 92 , 118 , 119 , 146

39 , 92 , 134 , 145

31 , 87 , 96 , 106 ^ 109 , 143

34 , 144

35 , 87 , 144
34 , 87 , 144
36

4 , 63 , 129 , 145

67 , 99 , 145 , 147
69 , 129 , 131
66 , 147

6 , 47 , 91 , 112 , 141

50 , 93 , 128 , 149

51 , 93 , 149

49 , 91 , 116 , 146

38 , 150

53 , 94 , 120 , 146

40 , 89 , 90 , 110 , 148
45 , 148

43 , 148

44 , 148

39 , 135 , 145

54 , 95 , 145

15 , 48 , 114 , 149
49 ,

56 , 97 , 147

58 , 97 , 124 , 150

154

Nuytsia Vol. 3 (1) 1980

Halostachys

13

Pachycornia

arbusciila (R.Br.) A. J. Scott
robusta (F. Muell.) Baill.
tennis (Benth.) Black
triandra (F. Muell.) Black

Salicornia

subgen. Arthrocnemoides Ung.-Sternb.

sect. Perennes Duval-Jouve.

arbiiscula R.Br.

australis Benth.

australis Forst.f.

hidens (Nees) Benth.

hlackiana Ulbrich

dliolaluin Bunge

donaldsonii Ewart et White

europaea L.

fruticosa (L.) L.

glauca Delile

glauca Stokes

indica Willd.

leiostachya Benth.

lylei Ewart et White

macroslachya Moric.

mucrnnata Lag.

pachystachya Ung.-Sternb.

pachystachya Black.

perennis Mill

quinqueflora Ung.-Sternb.

robusta F. Muell.

tenuis Benth.

triandra (F. Muell.) Druce
virginica Forskal

Sarcathria

indica (Willd.) D. Jackson

Sarcocornia

blackiana (Ulbrich) A. J. Scott
globosa P.G. Wils.
perennis (Mill.) A. J. Scott
quinqueflora (Ung.-Sternb.) A. J. Scott
subsp. tasmanica P.G. Wils.

Sclerostegia

arbuscula (R.Br.) P. G. Wils.
disarticulata P. G. Wils.
mcdullosa P. G. Wils.
moniliformis P. G. Wils.
tenuis (Benth.) P. G. Wils.

Tecticornia

australasica (Moq.) P.G. Wils.

12, 13, 16
20
16
22

16 , 79, 141
12, 28, 70
70

70
20

71
71
67
75
66
22

15
70
152
152
63, 71
66

49

152

152

75

75

70

71

16
22
16
152

12

63

13

5, 75 , 101, 138, 151
76 , 101, 150
70

5, 10, 71 , 101, 136, 150
74 , 140, 150

17

20, 81, 103, 142
19 , 80, 102, 141
24, 83, 142, 150
22, 81
22, 82, 142

53, 132, 133

Tegicornia

unillora P.G. Wils.

25,

7, 25, 84, 104, 143

Publication date of Volume 2 Number 6

The publication date of Volume 2 Number 6 was 21 December 1979.

91670/l/aO— IM

WILLIAM C. BROWN, Government Printer, Western Australia

Notes for Authors

Nuytsia publishes papers relating to the flora of Western Australia. All papers are refereed
outside the Western Australian Herbarium. The Herbarium reserves the right to reject
papers.

Manuscripts must be submitted in duplicate. They should be typewritten, double spaced,
with 2-5 cm margins and no underlining. Pages should be numbered. The desired
position of text figures may be pencilled in the margin. An abstract should be provided.
For style and layout, recent numbers of Nuytsia should be followed. Galley proofs will
be forwarded to authors for checking. Reprints may be ordered at the author’s expense.

CONTENTS

A revision of the Australian species of Salicornieae (Chenopodiaceae). By Paul G.

Wilson 3

Publication date of Volume 2 , Number 6 154