203.

On the basis of these features it is suggested that Aorothamnion is

developed from an Antithamnion-like ancestor, and because Antitharnnion

characteristically has pinnae in opposite pairs, Acrothamnion butleriae vdth

2 pinnae per whorl is ~sibly the most primitive species. The occasional

devel~poent of relic gland-cells also suggests its primitive nature.

Instability of pinna form occurs also in Acrothamnion preissii in Tihich 4

pinnae are formed (sometioes reduced to 3 in the lower parts of the plant by

loss of orie of the minor pinnae). Acrothamnion arcuatum with evenly-spaced

whorls each of 4 pinnae is the most stable form, and hence is probably the
most advanced. Details of its structure, however, need confirmation from

investigation of fresh material.

(Sand.)
TYPE species: Aorothamnion preissii/nov. comb.

Key to the Southern Australian species of Acrothamnion.

1. Uhorl-branchlets (pinnae) arranged dorsiventrally in each

whorl consisting of 2 opposite major pinnae between which 2

(or one) minor pinnae occur ••• ... A. preissii

1. Whorl-branchlets (pinnae) arranged in evenly-spaced whorls

each of 4 equal pinnae ••• ... ... ... ... .A. arcuatum

Note: Acrothamnion butleriae, although referred to in the above

discussion, does not occur in southern Australia.

 204.

1i.CROTIWOOON PREISSI!( 3or.d):. nov. comb.

Callithamnion preissii Sander 1845: 19; 1847: 166. Agardh, J. 1851:

33; 1876: 25. Kuetzing 1849: 651·

J.ntithamnion preissii De Toni 1897: 1414.

CaJ.lithamnion pulchellum Harvey 1854: 561; 1863J synop: No-. 692.

J. 1i.gardh 1876: 20. De Toni 1897: 1338. Lucas 1909: 49. Lucas and

Perrin 1947: 333.

Acrothamnion pulchellum (Harv.) J. l~ardh 1892: 25, f'igs. 6-10.

De Toni 1924: 451 . Reinbold 1898: 53; 1899: 50. Tokida and Inaba

1950: 124·

TYPE Locality: - Rottnest Island, Western 1~ust.

HOLOTY.~ : MEL, 10260, Preiss No. 2536.

DISTRIBilliON: , Western Australia to Port Campbell~

Victoria, oceurring epiphytically 0n a. variety o:£' algae (e.g. Ballia

oallitricha, B. mariana, Wrangelia orassa, Ace:ecarpia paniaulata and

Codium mamillosum),

(Fig. 24).

General Features: - The plant climsists of a creeping prostrate portion

attached to the host by digitate rhizoids and bearing free, erect branches

up to 1 .5 em long (Fig. 24A).

Each axial cell bears two opposite major whorl-branchlets (pinnae),

each with distichous, pinnately arranged, opposite pinnules and usually

terminated b,y a conspicuous gland10ell (Fig. 24 B,K). Two minor pinnae,

reduo~d to one in the oentre.:L and. ]..ower pal"ts. o~ the thallus, occur side

by side between the pairs of major pinnae. These are unilaterally placed

on the axes, all minor pinnae on the one branch being on the same side

so that the branch is distinctly dorsiventral (Fig. 24B).

Structure of the Thallus: -

(1) Form and Develo~ment of Main Axes:-- Elongation of the prostrate

axis occurs by transverse divisions of the dome-shaped terminal cell

(Fig. 24D). Often only two pinnae, reduced in form, are developed on each

cell of this axis; the basal cell of' one pinna of the pair bears an erect

branch while the other produces rhizoidal attachment organs (Fig. 24D a & b))

Eroot branches occur at irregular intervals, usually close together, along

the axis and soon become dominant over the pinna developed on the same cell ~

(Fig. 24Dp). Each erect branch may bear several lateral branches which also

arise from the basal cell of a pinna (Figs. 24A,Ba).

Mature axial cells in the erect branches are about 1~3 (~5) times as
~-
long as broad ((*50-60 x 120-160 (-190)p). The lower cells of lateral

branches mey be smaller than those above so that the branch tapers tow¥'ds

the base. Variation in proportions of axial cells results in variation in

overlap of the pinnae and proximity of the pinnules.

( 2) Development, Arr9ement and Branching of Pinnae: - Pinnae are

initiated one to three cells below the apex of branches and form initially

a chain of rachis cells, the basal cell of which alw~s remains smaller

than those next above it. Cells of the rachis are about as long as broad

(to 66p). The two major pinnae develop simple, distiohous, opposite
 206.

pinnules and, when mature, are up to 380 ( -580))-1 long with a rachis of

1 D-12 ( -14) cells. The longest pinnules are toward the base of the pinnae

and are about 9 cells 125 ( -190)}1 long, (Fig. 24B). Two (sometimes only

one) minor pinnae are initiated on one side of the axis betwden the two raajox·

pinnae; one is nearly always lost before long so that on most of the

thallus only one occurs. The minor pinnae vary in form (Fig. 24 E-I),

becoming more reduced from the upper to the lower parts of the plant. The

longer ones have a rachis of about 9 cells (150(-300)p). The major

pinnae may also be reduced towards the base of the plant so that a genera:

reduction in size of' all pinnae occurs, Those on the prostrate axis are

always reduced in both size end fo~ and. often bear unilaterally arranged

pinnules only.
C-
Gland-ciells: - Gland-cells occur terminally on the rachides of both

major and minor pinnae and hence do not form until the apical cell of the

rachis has ceased to dlivide transversely. The final pair of pinnules curv-8

upward and clasp the somewhat elongate and often angular gland-cell which is

about 25 x 18p (Fig. 24L). When a gland-cell does not develop several

small cells occur at the apex of the pinna (Fig. 24K).

Rhizoids and ~lttaohment Ol'ga.ns: - No axial cortioation occurs, but on the

pl'Ostrate axes l'hi.zo~dal at-t.ac.hment organs are borne on the basal cells of

the pinnae. Several multicellular rhi,90ids 1 soii1etimes sparingly branched,

mey arise from one basal cell.. When young the rhizoids are narrow and

elongate, but on attaohment to the host, the cells contract somewhat and

a d.igitate holdfa.st develops terminally (Fig. 24D, b & d).

 207.
'.

Tetrasporangia: -- Tetrasporangia are borne terminally on elongate pro-

~rusions from the upper side of the basal cells' of ·the·. p:i;m_ae. They are
~ :. ; ..
usually single -~n the ~or ~inn4e in the . upper p~s of the thallus (Fig .
. ;

24J), but occasionally more than one mey develop on a 'c~l.l; they may a1 so

ocoUl" on minor pinnae.

Each tetrasporangium is spherical when mature, about 4qu diam.

~ ~ ~-

..
• . •• ,._ ';. , • •· : ·I • • ' •. r •• U

within the gelatinous Sheath and qivision is cruciate although someti~es

., "-.
f! ; ; . . ..• "!' ... o#

forming sporangia apparently tetrahedral in form.

-~· ... . ·:... .·.
< col.
Spermatangia: Spermata.ngiYm- mother cells ( 2.5 - 4.5 .)l diameter) are borne

in terrrdnal ~lusters of 2-4 cells on branches of modified pinnules usually

toward the basal part of the pinnae. The axis of the fertile pinnule is

usually only 3-.6 cells long and with branches from each cell; occasiol"l.al.ly,

however, branching with development of spermatangial mother-cells takes

.·.
place from the. several uppermost cells of an elongate pinnule.
. ...\ . '

The branches which bear the spermatangi~ mother-cells consist of

1-4 successive whorls each of 2-4 cells (fig. 24C).

Development of Carpogonial Branch and Carposporopgyte: -- Carpogonial

I , ) '

branches, of the form found in Antitha:ID:ri:i:.cin~· develop on the basal cells

(supporting cells) of young ~innae near branch apices (Fig. 24 P,M) •

•
Four to eight m~ develop suocessively on one branch apex, and two of

these may initiate early stages in development of the carposporophyte •

• f I •:

After fertilization an elongation from the base of the carpogonium develops

...
t\ij~ds the r oundei auxiliary cell cut off from the upper side of the
.. . .
supporting cell (Fig. 24t'i), Presumably f'usion occurs between this
t \ • .; : ·;
 208.

protrusion of thG o.arpogonium e.nd. the auxiliary c-ell.. The auxiliary cell

then divides transversely to form a. lower foot cell and an upper rounded

central cell (Fig. 24 0). The first gonimolobe develops terminally from

the central cell and outs off several sterile cells which in turn give rise

to the carposporangia (Fig. 24 P,Q,R). The second gonimolobe is lateral

and develops ca.rposporangia qy a sioilar series of cell divisions (Fig, .

24. p ,.Q,R.).
After initiation of carpogonial branches the branch apex ceases

further elongation, and the carposporophyte is protected by the upwardly

directed surrounding pinnae. Pinnae on the upper cells of the branch apex

remain immature often with several smaller cells at the base (Fig.24 P) o

Discussion: -

Individual plants var,y in form due to variation in cell size and

proportion. The type material, for example, has comparatively short ceJ l .•

and, hence, close proximity of adjacent pinnae and pinnules. One collect ~ a ~ ~

from Flinders Bay, W.A. (f~• A 29,287) represents another extreme in fo~uJ

~nd approximate limits for dimensions of plants from this collection have

been indicated in brackets in appropriate places~. Many intermediate forrr..

are recorded, and cell size is not a satisfactory feature for the separat:;. ·'

of species. A Japanese species has also been described (Yendo 1916: 262;

Yamada 1928: 528; Yamada ~~Jnagaki 1935: 37) as Acrothamnion pulche,hllF'

/I se n £~ :L ? tC
/\.Yamada (not J, 1'..gardh) but~s probably more closely r el at ed to the New
Zealand Antitha.mnion a:p;llici,Plm J. Ag. The species Jl.crothamnion

pulchellum J. Ag. is desoribed by !okida and Inaba (1950) as having

 209.

sessile. tetra..spol'Qilgia on the "ba..sal. oo.l).& '0£ piDna.e but without the

elongate protrusion which occurs in the Australian species. This Japanese

plant m~ therefore also represent a different species. Acrothamnion

pulchellum (Harv.) J. Ag. is identical with the earlier described species

Callithamnion preissii Sander. On the basis of priority, the latter epiphet

must be retained. The type of Harvey' s C. pulchellum is in TCD 1 Herbarium

Horvey, No. 230.

li.CROTHQMNION ARCUATUM n. sp.

Thallus prostratus ramis erectis 1 aut non nullis em altis. Vortico-

raouli 4 e quoque cellula, usque ad 200).1 longi, cellula infima parva

pnaediti, curvato-adscententes. Vortico-ramuli ramis 1-jugatis, 2-6 cellulae

longis, exteriore obliquis, saepe ab latere superiore 1-pluribus ramis

brevi. bus instructis. Cellulae glandulosae in rachidibus vortico-ramulorum

terminales. Tetrasporangia, spermatangia et carposporophyti ignoti.

TYPE LocaJ.ity: Middle River, Kangaroo Is., South Aus.

~LOTYPE: -

DISTRIBUI'ION : - Known only from the type locality. Growing on

coralline algae.
(Fig. 23M - 0).
General Features: The plant consists of long unattached axes, one to

several om. high, arising from prostrate axes which adhere to the host by

means of branched rhizoid.al filaments bearing digitate terminal attachoent

organs.