r" ...

• I

Filho, Coll: TIte Genus Porphyrrz in the American South Atlantic

/ 191

Botanica Marina M.e Juan, Manue I lopez Vivas

Vol. XVIII, p. 191-197, 1975

110 - l'AeC

KM 107 Can. TIj-Fns.

174-4601 Ext 173 Ens.B e

The Genus Porphyra C. Ag. (Rhodophyta-Bangiales) in the American South Atlantic. 1. Brazilian
Species

E. C. de Oliveira Filho and J. con·

Dep. Bottinica, Unillersidade de Sáo Paulo, C. Postal 11461, S. Paulo, Brasil

(Rec. 2. 8. 74)

A revision of the literature and ~he study of abundant material showed that P. vulgaris C. Ag. and P. lacinia!a
(Lightt) C. Ag. (= P. umbilica/is (L.) Kuetz.), are not present on the Brazilian coast, the same being tIlle for the
more recent references of P. atropurpurea (Olivi) De Toni. P. TOseana Howe, though known only from the original
collection, is considered to be a good species. Two species, with two varieties each, namely P. acanthophora varo
acanthophora and varo brasiliensis, and P. spira/is varo spira/is and varo amplifolia are proposed as new taxa. A thorough
discussion to warrant the new view presented is also given.

Introduetion (3) P. roseana was described by Howe (1928) based on

Four species of Porphyra C. Ag. were referred to the
Brazilian coast up to now: (1) P. lIulgaris C. Ag., (2)
P. /aciniata (Lightf.) C. Ag. varo umbilicata C. Ag., (3)
P. roseana Howe, and (4) P. atropurpurea (Olivi) De
Toni.
(1) P. vulgaris was first referred by Greville (1833,
p. 449) based on material collected by Saínt Hilaire at
Macaé (R.J.). The same reference was later quoted by
Montagne (I839, p. 42). Today this species is eonsidered
under P. umbilicalis (L.) Kuetz. (ef. De Toni, 1897,
p. 20, as Wildemannia umbilicalis (L.) De Toni). These
were the only references of this species (or of P. umbili­
calis) to Brazil. The descriptions given by Greville and
Montl!grte (Le.) are not clear enough in order to ascer­
tain the present name that should be applied to the
material they studied.
(2) P. laciniata varo umbilicata was first referred to the
State ofMaranhao by Martius (1833, p.20). The same
citation was repeated by Martens (1870), and later
referred several times to Brazil (Zeller, 1876; Piccone,
1886,1889; Moebius, 1890). J. Agardh (1882) consí·
dered this taxon as a form of P. umbilicalis. Kylin (1944)
following J. Agardh considered it also as a forro of
P. umbilicalis, but later (1945) as an independent species,
P. laciniata (Lightf.) Kylin; however this same author
in 1949 united it again with P. umbilicalis (last two
references from Drew, 1954).
* Under an University oC SAo Paulo student-fellowship at the
time this paper was prepared.
material col1ected by J. N. Rose and P. G. Russen
(August 8,1915) at Cabo Frio (R.J.), the type locality.
This still remains the only reference to the species.
(4) P. atropurpurea (Olivi) De Toni was first referred
to Brazil, though with sorne doubt, by Joly (1957, .
p. 92). Later it was reported from several points along
the brazilian coast (Joly and Yamaguishi, 1963; Joly,
1965; Ferreira·Correia and Pinheiro-Yieira, 1969;
Yoneshigue-Braga, 1971 and Ugadim, 1974).
A close examination of abundant material from several
points of the Brazilian Httoral has convinced us that
none of these could be identified with any of the four
previously mentioned species and that it was necessary
to propose two new species, with two varieties each, to
accomodate the studied material.
Descriptions
P. roseana Howe
In arder to complement the descriptions of the Brazilian
species we transcribed here the original description of
Howe (1928, p. 190): "Densely cespitose, tawny violet
or olivaceous; thallus very shortly stipitate, usually
with several succesive convolute-spreading or cornuco­
pioid laminae or divisions from a coromon base, the
main divisions suborbicular or broadly ovate to oblong
oc linear-oblong, mostiy 1-3 cm long, strongly crispate,
monostromatic except at the extreme base, 20-30 J,J.
thick, rigid on drying and scarcely adhering to paper, the
margins cornrnonly lobulate or subdentate, the IobuIes
 192 Filho, Coll: The Genus Porphyra in the American South Atlantic Filh

often auricular, cucullate, cornute, or somewhat trum­

Monospores are also present at the margin where they
pet-shaped; cells mostly 13-25¡.¡. in maximum diameter
can germinate while still attached to the mother plant;
in surface view, their protoplasts irregularly angular or
we believe that these sporelings have arisen from mono­
sometimes fusiform, separated by spaces of 5-10 ¡.¡., the
spore germination because they develop differently
basal protoplasts flagellate-pyriform; cells in sections
from the carpospores. The plant is monoecious presen­
subquadrate, the superficial walls firm, 5-10 ¡.¡. thick;
ting spermatia and carpospores mixed together in a
monoicous; antheridial sori narrowly marginal, 32
marginal band, usually with the spermatia more external
antherozoids in each antheridium; sporocarps forming
than the carpospores; the carpospore mother cell is
submarginal sori, 8 carpospores in each sporocarp. On
divided by curve (convex-concave) walls, becoming
Balanus sp. etc., in the vicinity of Cabo Frio, Province
encased one in the other when young; there are 16 carpo­
of Rio de Janeiro, Brazil."
spores and 64 spermatia per mother cell, and it seems
that both can be formed by sister cells. In the State of
Material examined
Sao Paulo this species was found growing from July to
SPF 1974, part from the isotype, Cabo Frio (R.J.),
December, but at the end ofOctober it becomes rare;
August 8, 1915, Rose and Russell (20684).
it grows typically on rocks, and also on Chthamalus,
An exarnination of the original material has shown us at the supralittoral fringe in wave exposed places; how­
(
\,1 that Howe's description was in perfect agreement with
ever sorne plants were also found on wood arid in
the material. We have to add only that we don't know places where the salinity was a bit lowered by the
) what he meant by "subdentate margin" since we have presence of small fresh-water streams. In sorne specimens
we found a parasite fungus irregularly distributed in the
O/Ir not found marginal teeth. It is interesting also to note
that the species was said to be found growing on a spe­ basal portion of the thallus (see fig. 19).
cies of Balanus, a genus that grows down the infralittoral, This species is clearly distinct from all the other species

I since all the other Brazilian species of Porphyra are to referred to the American Atlantic up to now, since it is

I be found only at the supralittoral fringe. At this point

L they are commonly found growing on Chthamalus sp.
the only one with marginal teeth.

P. acanthophora Oliveira Filho et Coll, sp. nov. P. acanthophora varo bras;/iens;s Oliveira Filho et Coll,
Thallus larninis plus minusve orbiculatis, diametro maxi­ varo nov.

mo 9 cm, ad substratum basi fods, coriaceis, chartae A P. acanthophora varo acanthophora recedit thallo

herbarü haud adhaerentibus, 30-35 ¡.¡. crassis in medio vulgo fasciato, margine undulato, 15-25¡.¡. crasso in

ubi sterilibus, ad marginem dentibus minimis uni-multi­ parte media, ad chartam herbarii manifeste adhaeres­

cellularibus omatís, monoicis, 16 carposporis in spor­ cente.

angium exhibentibus, 8 in singulis stratis, cum sper­

matiis mixtis; spermatiis 64 pro antheridio in 4 strate Holotypus - Specimen collegunt Y. Ugadim 6-IX-1968

distributis; monosporangiis marginalibus. in litore maris Caiobá, Paraná State, Brazil, SPF 1216.

Isotypus ad BM et NY.

Holotypus - Specimen collegunt Oliveira Filho et CoIl

25-X·1973 in litore maris Ubatuba, Sao Paulo State,
Brazil, SPF 5170. Isotypus ad BM et NY.
Aditional material examined:
Sao Paulo State: Ubatuba - SPF 1963-1965, 1970,
4906,4907,4909,4910,4912,4913. Santos - SPF
1160, 1202,4914-4916. Rio Grande do Sul State:
Torres - SPF 4873.
Plants forming rosulated tufts, with several orbicular
blades united by a basal holdfast; frond brownish-red,
when young, to greenish at maturity, measuring about
9 cm in the larger diameter; texture membranaceous,
firm, on drying scarcely adhering to paper; thallus
monostromatic measuring about 50-60 ¡.¡. thick near the
base, 30--35 ¡.¡. in the rniddle portions to 23-30 ¡.¡. near
the terminal portions of sterile blades; cells showing one
single star-shaped plastid; marginal microscopic teeth,
l-multicellular, are present in the sterile borders where
,laminar or vesicular proliferations are also cornmon.
Aditional material examined:
Ceará State: Paracuru - SPF 4896-4898. Barra do
Ceará - SPF 4893, 4894, 4899, 4900. Barra do Rio
Cocó - SPF 4889, 4901, 4903. S-ao Paulo State: Uba­
tuba - SPF 1961, 1966, 1968. Santos - SPF 1008.
Paraná State: Caiobá - SPF 1216. This variety is
distinct from the type variety by its habit, size and
thickness. Here the blades are strap-shaped, usually
isolated, with undulated to folded margins and reaching
up to 20 cm long and 2-12 cm broad; the colour varies
from rose to vinaceous; its texture is more delicate
than the type variety and on drying adheres well to
paper; it is about 50 ¡.¡. thick near the base and 15-25¡.¡.
from the middle to the apex; the marginal teeth are
regularly distributed. It is less frequent than the varo
acanthophora though with a larger distribution; its
discontinuous distribution is certainly related to the fig. 1
Fig.3
lack of collections during the winter time at several
Fig.4
points of the Brazilian coast. Fig.6

80tanica Muina / Vol. XVIll/1975 / Fase. 4 Botar

 Filho, Coll: The Genus Porphyra in the American South Atlantic 193

,/
1 4

0­

[1S 5 .

5 e",

3 5 "

5CM

Fig. 1-2. Porphyra spiralis sp. nov. var; 8pira/is varo nov. isotype and holotype, respectively.
Fig. 3. Porphyra spiralil varo amplifolÚl varo nov., holotype.

Fig. 4-5. Porphyra acanthophora sp. nov. varo brasilienm varo nov., holotype and isotype, respectively.

Fig.6. Porphyra acanthophora varo acanthophora varo nov., holotype.

194 Filho, Coll: The Genus Porphyra in the American South Atlantie

7 8

'l '1. :,~,

y ~ -..

'~ tí lI'

.
..,1
t
/
'
~' 11
'
J

.. 1
\\
,¡ "
.,.... (...
J
\, '

~ (
.,_" , 1
,/;,'# ,
.-
Fig, 7, P. spiralis, detail of sterile margin X 480.
Fig. 8. P. spiralis varo spÚ'alis, frrst division of fertile eells leading to spermatium formation X 528.
Fig. 9. P.spiralis varo spiralis fertile margin showing spermatia X 200.
Fig. 10. P. spiralis varo amplifolia, detail of spermatium formation X 528.
Fig. 11. P. spiralis varo spiralis, detall of spermatia stained with haematoxylin X1.200.
Fig. 12. P. spiralis varo spiralis, eross seetion of the thallus showing a carpogonium and earpospore formation X 800.
Fig. 13. P. spiralis varo spiralis, eross seetion of a fertlle portion showing carpospores disposition X 600.

P. spiralis Oliveira Filho et Coll, sp. nov. Holotypus - Specimen collegunt L. Behar 14. IX. 1973

Plantae dense caespitosae, parte basali prehensili incons­ in litore maris ad Praia da Costa, Espirito Santo State,

picue evoluta. Frons figura fasciae angustae, 0,1-0,5 cm Brazil, SPF 4837. Isotypus ad BM et NY.

latae, contortae undulataeque, margine muticae, mem­

branaceae, chartae herbarii facile adhaerentis, 20-25 p. Aditional material examined:

crassitudine in medio, monoicae et vulgo protandricae, Espirito Santo State: Praia da Costa - SPF 4838-4849.

16 carposporis in sporangium in 2 stratis ex 8 sporis Ponta da Fruta - SPF 4864-4866, 4870, 4874, 4875.
compositis cum 64 spermatiis intermixtis; spermatiis Sao Paulo State: Ubatuba - SPF 4828-4830. Santos
16 in singulis stratis; mOnosporae desunt. - SPF 4826,4827. Plants tufted, with several narrow

Botaniea Marina I Vol. XVlI1/19751 Fase. 4

e
Filho, ColI: The Genus Porphyra in the American South Atlantie 195

.. ~,
~
'1···.
~'¡
"[
,f.. ". "~'o
//'' Y:'...o!' ¡." 21
~
,•....
, .'
~
" . . .
. 41:
.......

.' -. ( ~:'
• ';,o..... '-.,,:-,1/
~.í -'
':~Jf.l'. .
.. ~'.'
. .~:.
'¡.¡...
'~ 'V­
I .......

, '/
r .. '
J'
,
... ,

,/, >" •

. '. '9
/

Fig. 14-15. P. acanthophora varo acanthophora, detail of the dentate margin X 128 and 280 respectively.
Fig. 16. P. acanthophora varo acanthophora, marginal portion with proliferations and monospore-germlings X 128.
Fig. 17. P. acanthophora varo brasiliensis. detail of earpospore differentiation X 480.
Fig. 18. P. acanthophora varo brasiliensis, superficial view of a fertile area with carpospores X 96.
Fig. 19. P. acanthophora var. acanthophora, detail of an area with the parasite fungus X 384.
Fig. 20. P. acanthophora varo brasiliensis, frontal view ofvegetative cells X 320.
Fig. 21. P. acanthophora varo acanthophora, detail of a sporoearp in cross section X 800.

73 strap-shaped blades, 0.1-0.5 cm wide to 7-10 cm high,
attached by an inconspicuous holdfast; the blades are
twisted or have a folded margin; texture delicate, on
drying adhering well to paper; assuming a violaceous
colour when freshly dried. The thallus is monostromatic
49. and the cel1s present one plastid each; the thickness of
;. the thallus ranges from 38 ¡J. at the basal portions to
20-25 ¡J. at the middle and 19-23 Il at the tenninal
sterile portions; fertile portions measure about 30-35 Il
thick; in cross sections the ce1ls are square and regularly
disposed; the thallus margin is smooth and without any
kind of teeth or proliferations. The species is mono­
ecious presenting the spennatia and carpospores in an
almost continuous marginal band, except near the base;
there are 16 carpospores per mother ce1l, measuring
15-17 11 in diameter, and 64 spennatia. Monosporangia
were not found. Theplants grow on rocks or Chthamalus,
in wave exposed places usually on a leveI slight1y higher

:e.4 Botanica Marina / Vol. XVIII / 1915/ Fase. 4 17 A

196
than P. acanthophora varo acanthophora with wich it is
usually associated in nature.
is species is different from the other species referred
up to now to the American Atlantic mainly.because of
its thin, narrow strap-shaped thallus with folded margino
It also differs from the other monoecious monostro­
matic species by the number of carpospores per sporo­
carp, with exception of P. capensis Kuetz and P. acan­
thophora. However the first one is much thicker and the
second has a dentate margino
P. spiralis varo amplifolio Oliveira Filho et Coll, varo nov.
Discemitur a P. spirali varo spirali forma thalli; thallo
evidenter expansiore, 1-5 cm lato, margine plano vel
undulato sed non contorto.
Holotypus - Specimen collegunt L. Behar 22. IX. 1973
in litore maris ad Ponta da Fruta, Espirito Santo State,
Brazil, SPF 4868. Isotypus ad BM et NY.
This variety can be distinguished from varo spiralis by
its much wider thallus, that ranges from 1-5 cm. The
plants reach 13 cm high and the thallus is strap-shaped
or irregularly cleft leaf-like; the margins are undulated,
but not twisted.
Identification Key
1a. With marginal teeth o . o. 2
b. Without marginal teeth . o o 3
2 a. Thickness less than 30 IJ. (median portion, sterile
blades); blades isolated or a few together, strap·
shaped, with folded or undulated margins, up to
20 cm high . o . o . P.acanthophora var. brasiliensis
bo 30-60 IJ. thick (median portion, sterile blades);
blades orbicular in dense rosulated tufts, with un­
folded margin; up to 9 cm large o oo
. . . . . . . . . . o . P. acanthophora var. acanthophora
3a. Plants small (1-3 cm high), with suborbicular
blades; rigid, not adhering to paper ..... P. roseana
b. Plants larger(7-13 cm high), with strap-shaped
blades, adhering well to paper
4a. Blades very narrow (0.1-0.5 cm wide), twisted or
with strongly folded margins P. spiralis var. spiralis
b. Blades not so narrow (1-5 cm wide), with undulated
or smooth margins, usually partially cIeft ... o o ...
.. o oo P. spiralis varo amplifolia
Discussion
Conceming the four species of Porphyra referred up
to now to Brazil, in short we can say that:
(1) P. vulgaris C. Ag. and P. laciniata (Lightf.) C. Ag.
varo umbilicata C. Ag. are in the synonyrny of Po umbi­
Filho, Coll: The Genus Porphyra in the American South Atlantic
licalis (L.) Kuetz. and that this species does not seem
to occur in Brazil.
(2) P. roseana Howe was well described by Howe (1928)
as we could see while examining the original material.
Though it was never found again in Brazil or in other
places, it shows a group of characteristics that certainly
warrant its status as a valid taxon.
(3) P. atropurpurea (Olivi) De Toni involves a still un­
solved nomenclatural problem conceming its relations
with P. leucosticta Thuret. As one can see from the
original description of P. leucosticta (Thuret, in Le
Jolis, 1863), Thuret either didn't know or ignored
Ulva atropurpurea described by Olivi (1791). De Toni
(1897) was the first to note the similarity between both
specíes, when this author proposed the new conbination
P. atropurpurea (Olivi) De Toni, he included P. leucos­
ticta Thuret as its synonym. This apparently was accep­
ted by several phycologists (Collins and Hervey, 1917;
Howe, 1918; Taylor, 1937). Later other authors (Hamel,
1924; Taylor, 1957, 1960) did the opposite, Le., con­
sidered P. atropurpurea as a synonyrn of P. leucosticta
which is certainly a mistake, since Olivi's epithet is
older than the one'of Thuret. However, despite the
extensive description of Ulva atropurpurea given in the
original publication by Olivi (l.c.) it is still not clear
enough to warrant referring both names as a single taxon.
As we canjudge this could only be done after examining
the plants studied by Olivi, since Thuret's species is
clearly defined. However we could not identify any of
the existent Brazilian material either as P. leucosticta
or asP. atropurpurea as was done by Joly (1965),
Ferreira·Correia and Pinheiro-Vieira (1969), Braga
(1971) and Ugadim (1974).
When trying to identify the material coHected in Brazil,
we attributed a great importance to the presence or
absence of microscopical marginal teeth, even being
. aware that this criterium could be an artificial one.
However in this confused and stilllittle understood
genus we found it better to choose an objective
criterium in order to divide the Brazilian plants in two
groups which we consider as two new species namely:
4 P. spiralis and P. acanthophora. In each one of these spe­
cíes we subdivided the plants in another two groups
(of variety rank) using this time quantitative criteria as
thallus shape (relation between lengh and width), thick­
ness and indirect amount of mucilage. Based on these
characters it was easy to allot the majority of the specí­
mens in one or another of these four taxa, but not aH of
them. There are a few specimens from the State of
Paraná (SPF 1095, 1269, 1298a-b) without marginal
teeth, that cannot be placed under the varieties of
P. spiralis since the plants grow isolatedly and the thallus
is a broad leaf instead of strap-shaped. However we
have not enough material to justify the proposition of
a new species or of a third variety. We also found just a
few specimens of P. spiralis with intermediate charac-
Botanica Marina / Vol. XVIII /1975 / Fase. 4

Filho, Coll: The Genus Porphyra in the American South Atlantic
teristics between varo spiralis and varo amplifolia. These
individuals could be hybrids, ecads or even be in the
normal variation of the species, but at the present we
have no evidence to enlighten this point.
t is interesting to note that recent papers on Porphyra
(Krishnamurthy, 1972 and Kurogi, 1972) indicate the
existence of a large number of endemic species, with a
very restricted distribution, rather than the existence of
cosmopolitan species. This could mean that ir the
revisions were done in much larger areas, severa1 species
would certainly be grouped together. The Brazilian
coast is very large and there are still sorne points from
hich we did not collect material, which makes us
Literature
Agardh, J. G., 1882. TilI Algernes Syste~atik, Ard. 3. VI.
U1vaceae. Lunds Univ. Arsskr. 19: 1-177.
Collins, F. C. and A. B. Hervey, 1917. The Algae of Bermuda.
Proc. Amer. Acad. A rtsSci. 53: 1-195 + pis. 1-6.
De Toni, J. B., 1897. Sylloge Algarum. Omnium huiusque cog­
nitarum.4(1): 1-776. Patavii.
Drew, K. M., 1954. Studies in the Bangioideae III. The life history
of Porphyra umbilicalis (1.) Kuetz. varo laciniata (Lightf.)
J. Ag. A- The Conchocelis-phase in culture. Ann. Bot., Lond.,
N.S. 18(70): 183-211 + 3 pis.
Ferreira·Correia, M. M. and F. Pinheiro-Vieira, 1969. Terceira
Contribuic;40 ao inventário das algas marinhas bentonicas
do nordeste brasilieiro.Arq. Cien. Mar 9(1): 21-26.
Greville, R. K., 1833. Algae, 2: 447-450.1n: A. de Saint
Hilaire: Voyage dans le district des diamans et sur le litoral
du Brésil ... vol. 2, pp. 1-456. Paris.
Hamel, G., 1924. Floridées de France. Bangiales. 46 pp. Paris.
Howe, M. A., 1918. Algae, pp. 489-540. In Britton, N. L.,
Flora of Bermuda. IX + 585 pp. New York.
Howe, M. A., 1928. Notes on sorne marine algae from Brazil and
Barbados. J. Wash. Acad. Sci. 18(7): 186-194.
Joly, A. B., 1957. Contribuic;40 ao conhecimento da fiora fico
lógica marinha da Baía de Santos e arredores. Bol. Fac. ­
Filos. Cienc. Letr. S. Paulo. 217 Bot. 14: 1-196 + Mapa +
19 prs. . Taylor, W. R., 1957. Marine algae ofthe Northeastern coast of
Joly, A. B., 1965. Flora marinha do litoral Norte do Estado
de Sao Paulo e regiOes circunvizinhas. Bol. Fac. Filos. Cienc.
Letr. S. Paulo, 295 Bot. 21: 1-393.
Joly, A. B. and N. Yamaguishi, 1963. The life history of Por­
phyra atropurpurea (Olivi) De Toni, I. Bol. Fac. Filos.
Cienc. Letr. S. Paulo 19: 117-131 + 4 prs.
Krishnamurthy, V., 1972. A revision of the species of the
algal genus Porphyra occurring on the Pacific coast of
North America. Pacif, Sci 26(1): 24-49.
Kurogi, M., 1972. Systematics of Porphyra in Japan, pp. 167­
195 + 4 pis. In: Cantrib. System. Benthic Mar. Algae North
Pacif. ed. l. Abbott and M. Kurogi. pp. 1-279. Japan.
Kylin, H., 1944. Die Rhodophyceen der schwedischen Westküste.
Acta Univ. Lund., N. F.• Avd. 2,40(2): 1-104 + 32 taf.
Botanica Marina (Vol. XVlIl ( 1975 (Fasc. 4
-
197
think that the number of the taxa could eventúally be
increased in the future.
Acknowledgments
We wish to acknowledge Dr. A. B. Joly for the critica1
revision of the manuscript; Mrs. M. do Carmo Lockley
for the revision of the english text; Dr. C. T. Rizzini for
the latin diagnosis; Dr. R. Laval1ard for the french
surnmary; Mr. P. Windish for the german surnrnary; MSc.
L. Behar, Mr. L. Baptista and Mes. F. P. Vieira for the
collection of material; and to Dr. R. Starr for providing
a copy of Ülivi's papero
Le Jolis, M. A., 1863. Liste des algues marines de Cherbourg.
Mém. Soc.lmp. Sci Nat. de Cherbourg 10: 1-168 + 6 pis.
Martens, G. von, 1870. Conspectus algarum Brasiliae hactenus
detectarum. Vidensk. Medd. dansk naturh. Foren. Kbh. 2:
297-314.
Martius, K. F. P. von, F. G. Eschweiller and C. G. Nees von
Esenbeck, 1833. Flora Brasiliensis seu enumeratio plantarum
in Brasilia ... 4, 1: I-IV + 390 pp. (Algae 1-50). Stuttgart.
Moebius, M., 1890. Algae brasiliensis a cl. Dr. Glaziou collec­
tae. Notarisia 5: 1065-1090 + 1 pI.
Montagne, J. F. C., 1839. Cryptogamae brasiliensis seu plantae
a
celulares quas in itinere per Brasiliam céleb. Auguste de
Saint·Hilaire collectas recensuit observationibusque nonnulis
ilustravit. Ann. ScL nat., Bot. 2e. sér. 12: 42-44.
Olivi, G., 1791. "Sopra una nuova spezie di Ulva delle Lagune
Venete." Saggi Scient. Lett. Padova 3(1): 144-154, pI.
1-3.
Piccone, A., 1886. AIghe del viaggio de circumnavigazione della
"Vettor Pisani". 97 pp. + 2 pis. Génova.
Piccone, A., 1889. Nuove alghe del viaggio de circumnavigazio·
ne della "Vettor Pisa ni". Accad. Lincei 4(6): 10-63.
Taylor. W. R.. 1937. Marine algae of the Northeastern coast of
North America. Univ. Mich. Stud. ScL Ser. 13, VII + 427
pp. Ann Arbor.
North America. Univ. Mich. Stud. ScL Ser., 2nd ed. X +
509 pp. Ann Arbor.
Taylor, W. R., 1960. Marine Algae of the Eastern Tropical and
Subtropical Coasts of the Americas. Univ. Mich. Stud. Sci
Ser. 21, IX + 870 pp. Ann Arbor.
Ugadim, Y., 1974. Algas Marinhas Bentonicas do litoral sul do
Estado de Sao Paulo e do litoral do Estado do Paraná. Bol.
Bot1mica Univ. S. Paulo 2: 81-122.
Yoneshigue-Braga, Y., 1971. Flora marinha bentonica da Baía
da Guanabara e cercanías. 3. Rhodophyta 1. Inst. Pesq. Mar.,
R. de Janeiro 55: 1-36, pis. 1-14+ mapa.
Zeller, G., 1876. AIgae Brasiliensis.1n: Symbolae ad FloIam
Brasiliae centralis cognoscendam, ed. Warming, E. Vidensk.
Medd. dansk naturh. Foren. Kbh. pp. 426-432.
17 A*