The morphology and taxonomy of the red alga ... - SeaweedAfrica

Phycologia (1974) Volume 13 (1), 31-44
The morphology and taxonomy of the red alga Sarconema
(Gigartinales: Solieriaceae)
GEORGE F. PAPENFUSS* AND TrKVAH EOELSTEINt
* Department of Botany, University of California, Berkeley, California 94720, U.S.A.
t Atlantic Regional Laboratory, National Research Council of Canada, Halifax, Nova Scotia, Canada
A study of the structure and reproduction of two species of Sarcollema from East Africa has shown that the genus
is correctly placed in the Solieriaceae. The genus agrees with Solieria in that the auxiliary cell is indistinguishable
before diploidization. A sheath of peri car pic filaments is not produced about the gonimoblast as it is in CallophyclIs,
Solieria, Agardhiella and Meristotheca. Only two species of Sarcollema are recognized hy us, S. filiforme (Sander)
Kylin and S. scillaioides B0rgesen. S. Jurcellatum Zanardini, S. montagnei (Grunow) Kylin, S. indicum (J. Agardh)
Kylin, S. Jurcatum B0rgesen and S. filiforme f. gracillima Rayss are regarded as representative of S. filiforme.
Dicranema setaceum Sonder from Queensland, Australia, and D. setaceum var. "polensis Grunow from Samoa
were also found to be S. filiforme.
Introduction
The history of the family Solieriaceae, to which
Sarconema belongs, begins with J. Agardh, who in
1852 (pp. 718 and 721) erected the tribe Solierieae for
the genus Solieria J. Agardh (1842). Harvey (1853,
p. 115) elevated the Solierieae to the rank of subfamily
(except that he called it suborder) and included
in it the genus Eucheuma J. Agardh (1847) in addition
to Solieria. In 1872 J. Agardh included in the
Solierieae his new genus Meristotheca and Rhabdonia
Harvey (Hooker & Harvey, 1847), in addition to
Solieria and Eucheuma, and in 1876 he added:
Gelinaria Sonder (with a query; the genus is now
placed in the Cryptonemiaceae), Carpococcus
J. Agardh (currently treated as asynonym of Sarcodia
J. Agardh of the family Sarcodiaceae), and Catenella
Greville (1830) (with a query; the genus is now
referred to the Rhabdoniaceae).
Hauck in 1883 (pp. 17 and 186) elevated the sub­
family Solierieae to the rank of fami Iy and in 1886 he
placed Sarconema Zanardini (1858) in the family.
Schmitz in 1889 (p. 442) placed the following seven
genera in the Solieriaceae (J. Agard h) Hauck:
Rhabdonia Harvey, Erythroclonium Sonder (1852),
Areschougia Harvey (1855, nomen conservandum),
Solieria J. Agardh, Ellcheuma J. Agardh, Sarconema
Zanardini, and Thysanocladia (Endlicher) Lindley
Issued as NRCC No. 13699.
(now known as Callophycus Trevisan, 1848; Silva,
1957). The same seven genera were placed in the
Solieriaceae by Schmitz & Hauptfleisch (1896).
However, in both these accounts the family was
treated as a subfamily of the RhodophyJIidaceae.
De Toni (1897, 1924) adopted the classification of
Schmitz & HauptfJeisch.
.
Kylin in 1925 (p . 25) erected the family Rhabdoniaceae
for the reception of Rhabdonia, Agardhiella
Schmitz (Schmitz & Hauptfleisch, 1896), Anatheca
Schmitz (Schmitz & Hauptfleisch, 1896), Flahaliltia
Bornet (1892) and Solieria. (The inclusion of Solieria,
the type genus of the Solieriaceae, in the Rhabdoniaceae
by Kylin illegitimatizes the family.) In
1932 Kylin removed Solieria, Anatheca, Flahaultia
and Agardhiella to the Solieriaceae (the existence of
which he apparently had overlooked in 1925) and he
referred Catenella, Erythroclonium and Areschougia,
in addition to Rhabdonia, to the Rhabdoniaceae. In
his Gattungen Kylin (1956) added Catenellocolax
Weber-van Bosse (1928) to the Rhabdoniaceae and
dates the family as of 1932. Searles (1968, pp. 44 and
63) has provisionally also assigned Caulacanthus
Kiitzing (1843), Taylorophyclls Dawson (1961) and
Heringia J. Agardh (1844) to the Rhabdoniaceae.
Joly & Alveal (1969) and Augier & Boudouresque
(1971) have provisionally also placed the new genera
Montemaria and Feldmannophycus, respectively, in
the Rhabdoniaceae. As an appendix to the present
paper, the family is proposed for conservation.
31

32 Phyc% gia, Vol. 13 (1), 1974
In 1932 Kylin placed the following ten genera in
the Solieriaceae: Thysanocladia ( = Callophycus),
Flahaultia, Sarcodiotheca Kylin, gen. nov., Agardhiella,
Solieria, Sarconema, Eucheuma, Anatheca,
Meristotheca, and Euryomma Schmitz (Schmitz &
Hauptfleisch, 1896). In 1934 Kylin included Opuntiella
Kylin (1925) and Turnerella Schmitz (Schmitz
& Hauptfleisch, 1896) in the family and in 1956 he
credited the family with two additional genera,
namely, Predaea G. DeToni (1936) and Gardneriella
Kylin (194 1).
Dawson (1949) added the monotypic new genus
Reticulobotrys to the family and B0rgesen (1953)
added the new genus Tenaciphyllum. Feldmann
(1942) removed Predaea to the Nemastomaceae, in
which family the genus had been placed by Setchell
& Gardner (1930) when they described it (as
Clarionea gen. nov.). Dawson (1961) also placed
Predaea in the Nemastomaceae and Kraft & Abbott
(1971) have also convincingly demonstrated that
this genus belongs in the Nemastomaceae rather than
in the Solieriaceae. Kraft & Abbott also reduced
Yadranella Ercegovic (1949) to asynonym of Predaea.
The features whereby the Solieriaceae may be
characterized are: (1) the thallus is multiaxial in
construction and has a filamentous medulla; (2)
procarps are lacking; (3) the auxiliary cell initiates
only one gonimoblast cell; (4) the gonimoblast at
first develops inwardly; (5) the cystocarps are
embedded in the thallus or project from the thallus
surface and are usually provided with a distinct
ostiole; (6) a pericarp is usually, but not always, prod­
uced; and (7) the tetrasporangia are zonately divided.
The family Rhabdoniaceae which has been
confused with the Solieriaceae differs from the latter
especially in that (1) the thallus is uniaxial in
construction, and (2) the cystocarp of all genera
lacks a special pericarp.
Information of a greater or lesser amount on the
ontogeny of the cystocarp in the Solieriaceae is
available for the following genera: Sarcodiotheca
(Kylin, 1925, as Anatheca), Turnerella (Kylin, 1934;
Tokida & Masaki, 1957),* Sarconema (Kylin, 1932;
Rayss, 1963), Opuntiella (Kylin, 1934), Agardhiella
(Osterhout, 1896; Kylin, 1928), Solieria (Bornet, in
Bornet & Thuret, 1880; Kylin, 1932), Meristotheca
(Kylin, 1932), Gardneriella (Kylin, 1941) and
Callophyclls (Hewitt, 1960).
* Tetrasporophytes have been unknown in Tlirnerella.
Recently South et al. (1972) showed that T. pennyi) of
the north Atlantic Ocean possesses a crustose, Crlloria-like
tetrasporic phase.
In view of our very meagre knowledge of the
development of the female reproductive apparatus
and the carposporophyte in the Solieriaceae and the
availability of fertile material of Sarconema from
East Africa, the present study was undertaken with
the hope of contributing to a better understanding
of this genus and the family. Two species (the only
two in the genus in our opinion, as will be pointed
out in later pages), S. filiforme (Sonder) Kylin and
S. scinaioides B0rgesen were studied. Both liquid
preserved and herbarium specimens were examined.
Structure of the thallus
Thalli of Sarconema (Figs. 1-6, 19, 20 and 23-25) are
terete, solid, 0,8-2 mm in diameter near the base, up
to 20 cm long, dichotomously to subdichotomously
branched, and attached by a branched, hapteroid
hold fast. The branches taper gradually toward the
tip. Regeneration occurs from branches whose tips
have been lost. One or several new branches may be
produced from such tips, which frequently results in
a disturbance of the dichotomous habit.
The thallus is muItiaxial in construction and
consists of three clearly defined tissues (Figs. 19 and
20); an outer cortex of one or two layers of small,
tightly packed pigmented cells (Figs. 8 and 20); an
inner cortex of several layers of irregularly shaped,
pseudoparenchymatous cells, which in size decrease
toward the outer surface (Figs. 19 and 20); and a
central medulla of interwoven thick-walled filaments,
composed of elongated cells with lateral pit connections
between adjacent filaments (Figs. 19 and 20).
Development of the female reproductive system
As in other members of the Solieriaceae, procarps
are not formed in Sarconema. Carpogonial branches
are produced in large numbers in the tips of the
thallus branches. * The carpogonial branch is com-
* Young stages were best observed in material of S. scinaioides
that was collected by Papenfuss & Scagel (PR-YIl-\02) near
Dar-es-Salaam, Tanganyika, in October 1962. Rayss's (1963)
cystocarpic material of Sarconema from the eastern Mediterranean
was collected in July and October. Hauck's (1886)
material of this genus, collected by Hildebrandt in Somalia,
contained cystocarps in March, Newton's (1953) plant from
the Hanish Islands, Red Sea, contained tetrasporangia in
September, and Aleem (1948), who was the first to report the
genus from the Mediterranean, obtained plants with tetrasporangia
in October. B0rgesen (1932) had tetrasporic
specimens gathered (in Pakistan or India) in January.

,'.2
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2em
Papenfuss and Edelstein: Morphology and taxonomy of Sarconema 33
2 em
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FIGS. 1-3. Sarcol1ema /ili/orme. FIG. 1. Habit of a liquid preserved cystocarpic specimen collected by Papenfuss &
Scagel (PR·YUJ-48) near Dar·es·Salaam, Tanganyika, on 11-13 October 1962. FIG. 2. Habit of herbarium
specimen (showing the dichotomously branched nature of the species) collected by Papenfuss & Friedmann
(E62/20036) at Entedebir lslet, Dahlak Archipelago, Ethiopia, on 13 March 1962. FIG. 3. Voucher specimen of
Borgensen's 1934 record of S. jill'carul1I Borgesen from Pakistan (NY).

34 Phyc% gia, Vol. 13 (1), 1974
FIGS. 4-6. Sarconema scillaioides. FIG. 4. Habit of a cystocarpic
specimen collected by Papenfuss & Scagel (PR-VII [-
102) near Dar-es-Salaam, Tanganyika, on 11-13 October
1962. FIG. 5. Habit of a sterile specimen from the same collection
as F!g. 4. FIG. 6. Type of the species from Pakistan
(NY).
posed of three or four cells (Figs. 7-9) borne on an
undifferentiated, intercalary cell of the inner cortex.
This cell, the supporting cell, is larger than the cells
of the carpogonial branch (Figs. 7 and 8). As in
other Solieriaceae, it produces a single carpogonial
branch on the side facing the medulla (Fig. 8). The
trichogyne bends as it emerges from the carpogonium
and grows toward the surface of the thallus (Fig. 8).
An intercalary cell of the inner cortex functions as
auxiliary cell (Fig. 10). This cell is spatially removed
from the carpogonial branch and is not distinguish­
able from neighbouring vegetative cells before
diploidization and the initiation of the gonimoblast.
In this latter feature, Sarconema resembles Solieria
(Kylin, 1932), whose auxiliary cells also are indistin­
guishable before diploidization. In the other investi­
gated genera of the Solieriaceae, for example,
Sarcodiotheca (Kylin, 1925, as Anatheca) and
Callophycus (Hewitt, 1960), the auxiliary cell is
recognizable at an early stage by its dense contents
which stain deeply with dye.
After fertilization of thecarpogonium, a connecting
filament is produced from the region between the
carpogonium and the trichogyne, which structure is
often pushed to one side (Fig. 9). Long, branched
connecting filaments are common in fertile branch
tips, but their union with an auxiliary cell was not
observed. The abundance of carpogonial branches
and paucity of gonimoblasts in fertile tips suggest
that as in other Solieriaceae (Kylin, 1956, p. 274) the
number of auxiliary cells is small in comparison with
the number of carpogonial branches.
One of the first signs that an auxiliary cell has
been diploidized (presumably) is the development
toward the surface of the thallus of a small-celled,
deeply staining nutritive tissue from the auxiliary
cell (Fig. 10). No trace of the nutritive cells was
recognized in half-grown cystocarps, which fact
leads us to conclude that their contents are used up
during the development of the gonimoblast or they
become incorporated in the fusion cell. The gonimoblast
is initiated inwardly (Figs. 11 and 14-18) as in
Callophyclls (Hewitt, 1960), Solieria (Kylin, 1932)
and other genera of the Solieriaceae. At an early
stage, the auxiliary cell, some of the neighbouring
vegetative cells, and probably some of the first­
formed gonimoblast cells fuse and form a large
irregularly shaped, deeply staining fusion cell
(Fig. 15). In older stages, arms of the fusion cell
extend into the cortex in the region where the ostiole
will be formed (Figs. 12, 16- 19 and 21). In the
mature condition, gonimoblast filaments radiate
from the fusion cell (Fig. 22). Carposporangia are
cut off in chains from the tips of the gonimoblast
filaments (Figs. 17-19, 21 and 22). [(According to
Kylin (1932) the carposporangia are formed singly,
an erroneous impression based on the fact, no
doubt, that he had only herbarium material at his

-I
Papenfuss and Edelstein: Morphology and taxonomy of Sarconema 35
[ 20fLm
FIGS. 7-12. Sarconema scinaioides (PR-VlII-I02). FIG. 7. Carpogonial branch and supporting cell. FIG. 8. Part of
cross section through fertile tip of thalllls, showing a carpogonial branch and its supporting cell, which is an intercalary
cell of the inner cortex. FIG. 9. Two carpogonial branches, the carpogonium of one of which has produced a
connecting filament. FIG. 10. Part of cross section through fertile branch of thallus, showing an auxiliary cell (an
intercalary cell of the inner cortex) and a group of small nutritive cells which have been produced by it toward
the surface of the thallus. FIG. II. Auxiliary cell and young gonimoblast produced by it toward the interior of the
thallus. FIG. 12. Older gonimoblast with a large fusion cell and carposporangia.
F,G. 13. S. filiforme (PR-VIlT-48). Part of cross section of tetrasporic plant, showing tetrasporangia embedded
in the outer part of the inner cortex (aux, auxiliary cell; ca, carposporangia; cf, connecting filament; cp,carpogonium;
cpb, carpogonial branch; fc, fusion cell; go, gonimoblast; ic, inner cortex; nc, nutritive cells; oc, outer
cortex; sc, supporting cell; tr, trichogyne).
disposal.] Some of the gonimoblast filaments remain
sterile and seem to have a nutritive function. Early
in the development of the carposporophyte the
region where the ostiole will develop becomes
recognizable by the formation of files of small
cells (Fig. 14). However, the pore is formed only
when the gonimoblast approaches maturity (Fig. 22).
The carposporangia are round to ovoid, measuring
12-15 x 15-25 ",m. Mature gonimoblasts are 500-750
13
",m in diameter, more or less embedded in the
thallus (Figs. 17, 18 and 21), although the cystocarps
are readily recognized with the unaided eye as
bulges produced on the thallus (Figs. 1, 18 and 21).
In our material cystocarpic plants wer¢ rare-of
over 100 specimens examined only four bore cysto­
carps.
No male plants were observed in the material
examined by us.

E
E
10
o
16
'"
O.lmm
FIGs. 14-18. Sarconema scinaioides (PR-VIII-102). Parts of cross sections through fertile thallus, showing stages
in the development of the gonimoblast (aux, auxiliary cell; ca, carposporangia; fe, fusion cell; go, gonimoblast).
17
,
.\
]
t8

E
E
10
d
E
E
10
d
FIG. 19. Sarconema scinaioides (PR-VIlI-102). Part of cross section through fertile thallus, showing three embedded
cystocarps, the core of medullary filaments, and the pseudoparenchymatous inner cortex.
FIGs. 20-22. S. fili/orme (PR-VllI-48). FIG. 20. Longitudinal section of a vegetative branch. FIG. 21. Cross section
through thallus, showing a cystocarp. FIG. 22. Part of cross section through thallus, showing a mature cystocarp
(ca, carposporangia; fc, fusion cell; go, gonimoblast; ic, inner cortex; mf, medullary filaments; oc, outer cortex;
os, ostiole).
E
E
10
o
·22

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Development of tetrasporangia
The tetrasporic plants are morphologically similar
to the female plants. The sporangia are scattered
over the surface of the thallus, except that they are
not present in the lowermost parts of the branches.
They develop as lateral outgrowths from cortical
cells. When mature, they are zonately divided, appear
as dark, round spots, as seen in surface view, are
covered by one or two layers of cortical cells (Fig. 13),
and are relatively large, measuring 20-25 ",m in
width and 35-50 ",m in length.
Discussion
In spite of the then scanty knowledge of Sarconema,
Hauck (1886) and Kylin (1932) were correct in
placing the genus in the Solieriaceae.
In some members of this family, for example,
Callophycus (Hewitt, 1960, pI. 29), Solieria (Bornet &
Thuret, 1880, pI. 50, fig. 5; Kylin, 1932, fig. 3B;
Thein, personal communication), Agardhiella (Kylin,
1928, fig. 45A; 1956, fig. 206G), and Meristotheca
(Kylin, 1932, fig. 6A; 1956, fig. 208B), a sheath of
pericarpic filaments is formed about the gonimoblast.
Such a sheath is not produced in Sarconema (Figs.
18 and 22; see also Kylin, 1932, fig. 4A; 1956, fig.
208D; Rayss, 1963, fig. 6). A somewhat weakly
developed cellular peri carp is formed, however, by
proliferation of the cortex (Figs. 16-18, 21 and 22).
In Agardhiella fenera * (Osterhout, 1896, p. 416,
fig. 1) and Ellchellma procrusteanllm (Kraft, 1970,
fig. 4), sterile gonimoblast filaments establish
secondary pit connections with the pericarpic tissue.
In CallophycliS africanus the fusion cell is connected,
by means of filaments, to the pericarpic cortex
(Hewitt, 1960, fig. 21) but in this taxon the filaments
issue from the inner cortex. Comparable filaments
were not observed in Sarconema. It is seen, therefore,
that the Solieriaceae contains some genera whose
cystocarps are provided with a filamentous pericarpic
tissue about the gonimoblast and others (Sarconema,
Turnerella and Opllntiella) in which a filamentous
pericarpic tissue is lacking.
Se:!rles (1968, p. 75) was inclined to doubt that the
• This species is now known as Agardhie/la bailey; (Harvey
ex Ktitzing) Taylor (Taylor & Rhyne, J 970, p. J 3).
Papenfuss and Edelstein: Morphology and taxonomy of Sarconema 39
Sarcodiaceae Kylin (1932) should be retained as a
family distinct from the Solieriaceae. As far as
Sarconema and other better-known genera of the
Solieriaceae (see introduction) are concerned, there
are several features, both anatomical and reproduc­
tive, whereby the members of this family differ from
the type of the Sarcodiaceae, Sarcodia montagneana
(J.D. Hooker & Harvey) J. Agardh, as revealed by
Rasmussen's (1964) study of this species. It would
seem, therefore, that the Sarcodiaceae should be
retained for the genus Sarcodia (and perhaps the
little-known Chondrymenia Zanardini, see Kylin,
1956) and another home sought for the other
genus currently placed in the family, namely,
Trematocarpus Kiitzing [see Searles (1969), who has
merged Dicllrella Harvey ex J. Agardh in Trematocarpus).
Taxonomy
When Zanardini (1858) erected the genus Sarconema
he credited it with only one species, S. fllrcellatllm,
from the Red Sea. Kylin (1932) placed four species
in the genus: S. fllrcellatllm, S. monfagnei (Grunow)
Kylin (=Dicranema monfagnei Grunow, 1873-74=
Plocaria furcellata Montagne, 1850) from the Red
Sea, S. indicum (J. A-gardh) Kylin ( = Solieria indica
J. Agardh, 1852) from India, and S. /i/iforme
(Sonder) Kylin ( = Dicranema /ili/orme Sonder,
1845, 1846) from Western Australia.
B0rgesen (1932) reported Sarconema furcellatum
from Pakistan and India, and in 1934 added two new
species from Karachi, Pakistan, to the genus: S.
furcatum and S. scinaioides. In 1937 he reported
S. indicllm from Cape Comorin in southern India,
and in ] 939 he reduced S. montagnei and S. indicllm
to the synonymy of S. fllrcellafllm. Newton (1953)
agreed with B0rgesen that S. montagnei and S.
indicllm were merely variants of S. fllrcellatum. She
thought that S./iliforme may well belong in the genus
Dicranema, where Sonder (1845, 1846) had originally
placed it, rather than in Sarconema. On the other
hand, she found that No. 316a of Harvey's Australian
Algae (named D. /i/iforme by Harvey), a duplicate
of which Kylin (1932) had seen in the Agardh
Herbarium, was indeed a species of Sarconema, viz.
S. fllrcellatllm.
FIGS. 23-25. Sarconema fili/orme. FIG. 23. Type of DicI"Gnema filiforme Sonder from Western Australia (MEL).
FIG. 24. Lectotype of Dicranema selacellm Sonder from Queensland, Australia (MEL). FIG. 25. Lectotype of
D. selacellm var. lIpolensis Grunow from Samoa (W).

40 Phyeologia, Vol. 13 (1), 1974
An important point as regards the nomenclature
of Sareonema /ureellatum Zanardini (1858) that
needed to be settled was whether Dieranema filt/orme
Sonder (1845, 1846) is representative of Sareonema
[as Kylin (1932) and BrJrgesen (1934, 1938, 1943,
1950, 1952) believed] or of Dieranema [as Newton
(1953) thought], and if it should prove to be a species
of Sareonema, whether it is an autonomous species
or the same as S. /ureellatum, a species which
Newton (1953) had shown to occur in Western
Australia. The second author was privileged to see
Sonder's type specimen (Fig. 23) while she worked
in Dr Womersley's laboratory at the University of
Adelaide, South Australia, during part of 1972.
Examination of this specimen (MEL) showed not
only that Sonder's species is a species of Sareonema
but that it indeed is the same as S. /ureel/atum. The
Australian plants of this taxon are somewhat more
slender and elongate than those from the Red Sea
and seldom exceed 0·8 mm in diameter. B0rgesen
(1934) thought that the cells of the parenchymatous
tissue were larger in S. fili/orme than in S. /ureel/atum
but we have seen no evidence of this. In writing to the
first author about Sonder's specimen, Dr Womersley
commented as follows: 'The type specimen of
[Dieranema] fili/orme fits the habit description of
Sonder. The plant is tetrasporangial and many of
the axes are truncated, probably by grazing. New
branches have arisen from many of the truncations,
and thus appear "articulated" and somewhat
swollen, as referred to by Sonder. They bear tetra­
sporangia and might give the impression that the
sporangia were confined to them if no other parts
were sectioned. Sporangia are general throughout
most of the plant.' It is the misconception of Sonder
that the tetrasporangia were confined to the some­
what swollen tips of the branches that caused Newton
(1953) to conclude that Dieranema /ili/orme Sonder
probably is a species of Dieranema rather than of
Sareonema.
Since the binomial Dieranema fili/orme Sonder
(1845) has priority over Sareonema /ureel/atum
Zanardini (1858) the correct name of the species that
for more than 100 years has been known as S.
/ureellatum is S. fili/orme (Sonder) Kylin.
While in Adelaide, the second author also had the
opportunity of examining the lectotypes of Dieranema
setaeeum Sonder (1871) (Fig. 24) and D. setaeeum
var. upolensis Grunow (1873-74) (Fig. 25),
which specimens Dr Womersley had borrowed
from Melbourne and Vienna, respectively, for
Mr G. Kraft's study of Dieranema. Both these taxa
proved to be representative of Sareonema fili/orme.
A comparison of one of the Kotwal specimens
(in Herb. N.Y. Bot. Gard., Fig. 3) of Sareonema
/ureatum from Pakistan, cited by B0rgesen (1934)
when he erected this species, with material from East
Africa and elsewhere has convinced us that this
species also should be merged in S. fili/orme.
Rayss in 1963 described two new formae under
Sareonema fili/orme, forma curta (from Bombay,
Karachi, Yemen, Mauritius and the eastern Mediter­
ranean) and forma graeillima (from the Mediter­
ranean coast of Israel). Papenfuss (1968) concluded
that forma curta probably is representative of S.
fureel/atum. However, Rayss failed to indicate a
type for the taxon, which renders it invalid in
accordance with Article 37 of the Seattle Code. In
the light of present knowledge of the variability of
S. fili/orme it seems reasonable to conclude that
S. fili/orme forma gracil/ima is the same as S.
fili/orme.
It would seem, therefore, that as far as known,
Sareonema includes only two species, S. fili/orme
(Sonder) Kylin and S. scinaioides B0rgesen. S.
scinaioides (Figs. 4-6) is similar in general appearance
to S. fili/orme (Figs. 1-3 and 23-25), but
plants of the former are of greater diameter (up to
2 mm in diameter), of a much softer texture and
adhere more firmly to paper. The two species were
obtained at the same locality near Dar-es-Salaam,
Tanganyika, by Papenfuss & Scagel. [Schmitz
(1895, p. 139) had previously reported S.fili/orme [as
S. /ureel/atum] from Dar-es-Salaam and Zanzi­
bar.]
Sareonema scinaioides has no synonyms and the
present report of the presence of this species at
Mambrui, Kenya, and Dar-es-Salaam, Tanganyika,
constitutes the first record of its occurrence outside
of Pakistan, whence it was described by B0rgesen
(1934).
The synonymy of Sareonema fili/orme follows:
Sareonema fili/orme (Sonder) Kylin
Kylin, 1932, p. 22.
Dieranema fili/orme Sonder, 1845, col. 56; 1846,
p. 173.
Cystoclonium fili/orme (Sonder) Kiitzing, 1849,
p. 757; 1868, p. 6, PI. 18, Figs. a and b.
Sareonema fili/orme f. graeillima Rayss, 1963,
p. 101, Fig. 7.
Sareonema fili/orme f. curta Rayss, 1963, p. 98,
Figs. 1-6 (nomen nudum).

Sarconema furcellatum Zanardini, 1858, p. 264,
PI. 10, Fig. 1.
Plocaria furcellata Montagne, 1850, p. 243.
Graci/aria furcellata (Montagne) Zanardini, 1858,
p.266.
Trematocarpus furcellatus (Montagne) Kiitzing,
1869, p. 27, PI. 73, Figs. c and d.
Dicranema furcellatum (Montagne) J. Agardh,
1876, p. 436 (non J. D. Hooker et Harvey).
Dicranema montagnei Grunow, 1873-74, p. 43
(=Plocaria furcellata Montagne non D. furcellatum
J. D. Hooker et Harvey).
Sarconema montagnei (Grunow) Kylin, 1932,
p. 21, PI. 8, Fig. 16.
Solieria indica J. Agardh, 1852, p. 723.
Sarconema indicum (J. Agardh) Kylin, 1932,
p. 22, PI. 8, Fig. 17.
Dicranema setaceum Sonder, 1871, p. 58.
Dicranema setaceum var. upolensis Grunow, 1873-
74, p. 43.
Sarconema furcatum B0rgesen, 1934, p. 12, Fig. 8,
PI. 2, upper figure.
Material examined
Sarconema filiforme (Sonder) Kylin
Red Sea-Ethiopia-Dahlak Archipelago: Ente­
debir Islet (Landing Bay), 11. iii .1962, Papenfuss &
Friedmann E62/2003 (tetrasporic); 13 . iii .1962,
Papenfuss & Friedmann E62/20036 (Fig. 2) (tetrasporic)
and E62/20037; (Goliath Bay), 23.iii.1962,
Papenfuss E62/20167; Harmil Islet, 28 . iii .1962,
Papenfuss E62/20287 (tetrasporic). Massawa, iv.
1870, Issei (UC 90890; voucher material of Piccone's
1884 p. 320 Massawa record of Dicranema furcellatum).
French Somaliland: Djibouti (Le Plage), 15 and
16.ix.1962, Papenfuss & Scagel PR-I-65.
Somalia: Las Khoreh, iii .1873, Hildebrandt
(UC 408988; voucher material of Hauck's 1886
p. 167 record of S. furcellatum).
Kenya: Mambrui (north of Malindi), xii.1967,
Isaac (ADU 40349, 40418); Malindi (Outer Malindi
Bank), 5.x.1962, Papenfuss & ScageIPR-VC-55.
Tanganyika: Kingombe (35 km south of Tanga
Railway Station), 7.x.1962, Papenfuss & Scagel
PR-VII-34; Dar-es-Salaam (point north of Oyster
Bay), 11-13 .x.1962, Papenfuss & Scagel PR-VIII-48
(female, Fig. 1, and tetrasporic).
Zanzibar: Bwenju Reef, 18 .x .1962, Papenfuss &
Scagel PR-XV-62.
Papenfuss and Edelstein: Morphology and taxonomy of Sarconema 41
Mozambique: Praia Chokas (north of Lumbo),
15-17. xi .1962, Papenfuss & Scagel PR-XXVIII-92;
Zavora Reef, 4 .xi .1962, Papenfuss & Scagel
PR-XX-54.
Pakistan: near Karachi, no date, Kotwal (NY:
voucher specimen of B0rgesen's 1934, p. 12 record
of S. furcatum [our Fig. 3)).
India: Bombay, xii. 1927 and i. 1928, Borgesen
(UC 581368 and 581375, respectively; voucher
material of B0rgesen's 1934 p. 11 record of S.
fi/iforme from Bombay).
Australia-Western Australia: no date, Preiss
[2557] (MEL, Type of Dicranema filiforme Sonder
1845 col. 56; 1846 p. 173 [our Fig. 23)); no date,
Harvey 316a (MEL, voucher specimen of Harvey's
Australian Algae No. 316a of D. filiforme); Perth
(North Beach area), iii .1959, Norris (ADU 22268);
Fremantle, iii.1951, Royce (ADU 14143); Cockburn
Sound, iv.1966, Kraft (G. T. Kraft's personal
herbarium); Safety Bay, vii .1966, Kraft 1913 and
1919 (Kraft's herbarium).
South Australia: Elliston (inner reef), i.1951,
Womersley (ADU 13695a).
New South Wales: Port Stephens, no date (MEL,
Algae Muellerianae curante J. G. Agardh distributae,
as D. filiforme).
Queensland: Port Denison, no date [Fitzalan]
(MEL, Lectotype of D. setaceum Sonder 1871 p. 58
[our Fig. 24]).
Samoa: Upolu, no date, Graeffe fYI, Collectio
Grunow No. 33896, Lectotype of D. setaceum vaL
upolensis Grunow 1873-74 p. 43 [our Fig. 25]).
Sarconema scinaioides B0rgesen
Kenya: Mambrui (north of Malindi), xii. 1967,
Isaac (ADU 40418).
Tanganyika: Dar-es-Salaam (point north of
Oyster Bay), 11-13. x. 1962, Papenfuss & Scagel
PR-VIII-102 (female, tetrasporic and sterile [Figs. 4
and 5]).
Pakistan: near Karachi, no date, Kotwal (NY,
'Type'; right half of the specimen illustrated by
B0rgesen 1934 PI. 2; our Fig. 6).
Acknowledgments
This work was done while the second author was on
leave of absence from NRCC at the Department of
Botany of the University of California, Berkeley,
and the Department of Botany of the University of

42 Phyc% gia, Vol. 13 (1), 1974
Adelaide, South Australia. The material from the
Red Sea and East Africa was collected under the
auspices of the United States Programme in Biology
for the International Indian Ocean Expedition, which
Programme received its funds from the National
Science Foundation. As a participant in the United
States Programme the first author had two opportunities
in 1962 of collecting marine algae in East
African waters. During March and April he was a
participant in the Israel South Red Sea Expedition
and from September to December Dr R. F. Scagel
of the University of British Columbia and he had
their own expedition to East Africa. He should like
to express his appreciation to the Israel South Red
Sea Expedition and especially to its leader, the late
Professor H. Steinitz of the Hebrew University of
Jerusalem, for inviting him to join the expedition.
Dr E. Imre Friedmann helped with the collection of
some of the Red Sea material. Dr H. B. S. Womersley
and Mr G. T. Kraft kindly allowed us to incorporate
in our study the material of putative taxa of Dicranema
which they had borrowed from Melbourne
(MEL) and Vienna (W) for Mr Kraft's study of this
genus and which proved to be representative of
Sarconema. Mr Kraft generously also gave us photographs
of these taxa (reproduced as Figs. 23-25).
We are indebted to Miss Charlotte Mentges for help
with the illustrations. The processing of the Papen­
fuss-Scagel collections was done with the aid of a
grant from the National Science Foundation
(GB-1656).
Appendix
Rhabdol1iaceae Kylin (1932) proposed for conservation.
Rhabdoniaceae Kylin, Lunds Univ. Arsskr., N.F.,
Avd. 2, 28 (8): 32, J932. Nomen /amiliarum conservandum
propositum. Genus typicus: Rhabdonia
W. H. Harvey. Nomen /amiliarum rejiciendum
propositum: Rhabdoniaceae Kylin, Lunds Univ.
Arsskr., N.F., Avd. 2, 21 (9): 38, 1925 [fam. illeg.
pro Solieriaceae (J. Agardh) Hauck, 1883].
The Rhabdoniaceae as currently circumscribed
includes five, and possibly ten, genera. The family
has no synonyms; it would be advantageous, there­
fore, to conserve the name rather than propose a new
family name.
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