C

1996 The

Japan

Mendel

Cytologia

Society

Life History and Cytology of the Freshwater

Chaetomorpha
exilissima sp. nov
(Cladophorales,
Elisa
1

Laboratory

of Nonvascular

Universidad
2

Accepted

March

R. Parodi1,2

Nacional

Plants,

1996

Alga

Chlorophyta)

and

del Sur,

61: 179-188,

Eduardo

J. Caceres1

Department

San Juan

of Biology

670,

8000

and

Bahia

Biochemistry,

Blanca,

Argentina

Argentinian Institut of Oceanography (IADO), Av. Alem 53, 8000

Bahia Blanca, Argentina

14, 1996

The genus Chaetomorpha Kutz. includes filamentous unbranched green algae with multinucleate cells possessing a parietal reticulate chloroplast with numerous pyrenoids. The
filaments are attached to the substratum by means of a disc-like holdfast that may produce
finger-like projections (Blair 1983). Some species remain attached during their entire life while
others produce free-floating plants in certain periods of the life-cycle (Fritsch 1956). All
species exhibit alternation of gametophytic and sporophytic generations.
The significance of this work resides on that with exception of the early work of Lagerheim
(1887), a more recent detailed study of the biology and morphology of a freshwater species of
the genus has been never done. Consequently, the aim of this paper is to provide precision on
the cytology of the different stages of the life history of Chaetomorpha exilissima , a new
freshwater species found in Argentina. Moreover, since the mitosis has never been studied in
detail in the genus, existing only images of certain individual stages of the process (Kapraun
1993), the present are the first complete observations of the mitosis in the genus Chaetomorpha.
Material
The

study

was

Independencia,
(3841'S,
fixed

situ

medium

to

Cultures

on

the

Blanca

Biologia

cultures.

No

mounted
acetate

a cool-white

2 hr

(Nunez

for

15 min

in

darkness,

of
1968).

(BBB).

1N

2%
The

used

was

carried

for

out

during

distilled
acetic

Nonvascular

with

20

min

in

washed

solution

of
was

were

made
of

in

70%
at

ferric
made

Bahia

induction

out

acid

water,

karyotype

the

carried

hydrochloric

bleached
to

of
Sur,

Observations

were

was

del

at

67

60,

on

stained

distilled

haematoxilin
using

the

temperature

under

15 : 9 hr

of

with
of

"Cario"

was

For
acid

to

subsequent-

stain

sodium

(Johansen
metasulphite

during
the

plants

acetic

Schiff

water

of
andthe

zoospores.

glacial

material

LD

second.

collected

and

with

per
Department

recently

The

cultures.

Argentina

gametes

was
natural

room

Plants,

a 1 : 3 : 3 mixture
with

in

gmol/m2

Blanca,

a 1 : 3 mixture
ethanol.

material

starting

20-21C

providing

the

and

sites,
at

lamps

Laboratory

of

Parque

Argentina

transported

dimensions

collection

in

Aires,

Part
was

or

located

Buenos

autumn.

cell

the

Nacional

fixation

in

acid
drop

the

pond,
de

collection

north-west

fluorescent
in

methods

Postfixation

hydrochloric
in

deposited

Herbarium

the
of

the

artificial

and

of

from

from

Universidad

special

rest

coming

with

observations,

ethanol.

for

water

an

Provincia

determination

provided

are

from
Blanca,

summer

The

immediate

sterilized

paratypes

hydrolyzed

1 N

in

Bahia

during

illumination

Biochemistry,

karyological

1940)
to

grew

methods

coming

de

counts.
for

illumination

and

absolute
ly

made

and

Biology
Bahia

Filaments

sunlight

by

material

Partido

chromosome
laboratory

were

Holotype

or

the

under

regime

for

in

Blanca,

6215'W).

in

and

conducted

Bahia

and

30 min,

addition

computer

of

and
iron

program

Elisa

180

R. Parodi

and

Eduardo

J. Caceres

Cytologia

61

(Coccuci and Miinch 1989). Centromeric position was inferred by the shape and curvature of
the chromosomes (Wik-SjOstedt 1970). The terminology proposed by Battaglia (1955) has
been adopted. Photomicrographs were taken with a Leitz SM Lux microscope and a Wild MPS
15 Semiphotomat camera.
Results

and discussion

Life history
Life

history

gametophyte

(n

nucleate

cells

cell

3-5

wall

(Figs.

5,

6)

nuclei

of

the

differed

the

diam.,

were

Upon

18),

sporophyte

250-500 m
gm

long

thick

were

(Figs.

also

ovoid,

reported

7).

Fig.

and

proper
Successive

1.

zoospore.
flagellated

C.

meiosis

aerea

lose

mitoses

cell
and

exilissima.

2. Attached
zoospore.
gamete.
6. Syngamy.

9,

10)

which

vegetative

a conspicuous
1929,

elongation
cytokineses

Life

and
(Figs.

history.

1971)
and

M!:

In

the
11,

meiosis.

all

and

C.

S!:

and

and
diam.,

sequence

resulted

syngamy.

an
in

7 ,am

the

has
1972).

substrate

incipient
young

in

events

(Patel
to

all

thus

described
long,
of

melagonium

of

in
and

previously

holdfast
plants

1. 4-flagellated

3. Germling.
4. Free gametophytic
haploid
filament.
7. Zygote.
8, 9. Germination.
10. Free zoosporophytic
filament.

multi-

75 m

cytokinesis

themselves

production
12)

with

adult

parietal

10 gm

This

attached

The

synchronically
one

Zoospores,

eyespot.

Patel

flagella

long

after

cells.

cells.

. 1.

reticulate

occurred

and

Fig

free-floating

75-150 m

vegetative

their

were

in

chloroplast

zoosporangia

the

with

plants

diameter,

from
to

depicted

zoosporangia,

(Figs.

(Hartmann

with

in

developed

and

began

of

dimensionally

zoospores

Chaetomorpha

isomorphic

= 36),

30-50 m

identical

4-flagellate

the

(2n

Pairs

which

are

for

germination

is diagramatically

and

after

cells

zoosporangia

exilissima

2-4).

produced

vegetative

Germination
(Fig.

Chaetomorpha

morphologically

species

been

of

5. 2diploid

that

1996

Cytology

Figs.

2-12.

place

by

Chaetomorpha
growth

elongated

cell

numerous
acetic

pores.

pores.

Fig.

Metaphase
bars

undergone

disc-like

4.

to

zoospore

plate

of an
Figs.

the

its

8.
cells.

Cell
cells

Young

9,
9.

=20

takes

signifies

an

with

numerous

nuclei

(carmin-

released

through

opposite

being

zoospores

10.

in

Figs.

filament

through
which

an

opposite
incipient

attached

Initial

Pachitene.
Fig.

gm;

This
star

a sporophytic

multicellular

Figs.

nucleus.
bars

The

of

are
of

(arrows).
cell.

germling

Fig.

gametophytic
6, scale

3.

release

differentiation.

5, and

elongated

zoospores

unicellular
Fig.

of

haploid
3,

Fig.

after

holdfast.

Cytokinesis

an

Sporophytic

that

haploid

diameter

prior

in

181

Chaetomorpha

Sporophyte.

wall

Fig.
Note

gametophytic

= 50 gm;

2.

zoosporangium

irregular

Alga

cytokinesis.

heads).

Empty

the

Fig.
transversal

Zoosporangia.

6.

into

nuclei

8, scale

5.

Attached,

Note

sporophytic
12.

having

Fig.
7.

centripetal

(arrow

Fig.

developed

germling.

11,

after

Freshwater

exilissima.

pyrenoids
stain).

lateral

has

of

of the

stages
Fig.

13.

Karyotype.

7, 9-13,

scale

lateral
holdfast

gametophytic
of

10.

with

the

meiosis

Leptotene.
Figs.
bars

of
Figs.

2, 4,
= 5 m.

and

Elisa

182

became

attached

disc-like

holdfast

to

remained

attached.

flagellate

with

escaping
of

the

by
(Fig.

migrated

to

Cdceres

14

a single
were

laterally

the

by
cells

fuse

with

with

cell

these

germlings

200

pore.

and

losing
the

(Parodi
the

basal

even

Cdceres

and
the

the

in

this

in

in

Rhizoclonium
both

ending
whilst

and

gametes
14),

is

maintain

gametes
conjugated

contents

of

similar

one
to

same

size

15

16

cell

that

(Parodi
the

2-

germination

hieroglyphicum
cells

mature

between

very

in

ovoid,

Isogametes
the

61

they

when

subsequent

normally

rounded;
process

diam.,
diameter

(Fig.

gametangium.

The

case

diam.

and

occurred

same

in

uniform

diam.

formation

becoming

and

20 m

no

60 m
6 m

zygote

Cytologia

long,
had

long,

other.

1995)
that

um

Syngamy

from

of

J. Cdceres

450

long,

Syngamy,

mobility

difference

gm
um

15-18).

contents
and

only

10

(Figs.

individual
after

Eduardo

were

lateral

and

of

eyespot,

observed

same
15)

surera
1993)

The

conspicuous

through

produced

substrate
8).

Gametangia
a

zygote

Cladophora

the

(Fig.

R. Parodi

17

18

Figs. 14-18. Chaetomorpha exilissima. Sexual reproduction.

Scale bars =10 gm. Fig. 14.
Gametangium. The zoogametes are being released through a lateral pore. Fig. 15. Syngamy.
The eyespots of both gametes are indicated by arrows. Fig. 16. Copulating gametes. The
contents of one cell migrate to fuse with the contents of the other. Note the eyespots of both
gametes (arrows). Fig. 17. Germinating zygote with a short germinating tube. The two eyespots
derived from two gametes are visible (arrows). Fig. 18. Germinating zygote. The eyespots have
migrated to the germination tube (arrows).

of
and

while

1996

Cytology of the Freshwater

Alga Chaetomorpha

183

in C. surera and R. hieroglyphicum one of the cells growths considerably. It is the first time that
such tipe of sexual reproduction is described in the genus Chaetomorpha. During the first stages
of zygote germination, a long germinating tube was produced, into which both eyespots
migrated (Fig. 18, arrows).
Mitosis

The progress of the mitosis has been followed in sporophytic plants. The normal number
of nuclei per cell reached 40-50 (Fig. 4). Nuclear divisions occurred simultaneously not in all
but in several nuclei of the cell. During the process six stages could be distinguished: prophase,
metaphase, early anaphase, late anaphase, early telophase and late telophase. In interphase
nuclei reached up to 7 gm in diameter, were spherical and exhibited single conspicuous nucleoli
circled by numerous chromatic condensations similar to chromocenters. During prophase (Fig.
19) the number of chromatic condensations increased around the nucleolus, resembling the case

20'

21

22

23

24

25

Figs. 19-25. Chaetomorpha exilissima. Sporophytic mitosis. Scale bars = 5 um. Fig. 19. Prophase. The nucleolus is circled by peripheral chromatic condensations. Fig. 20. Metaphase plate
with 36 chromosomes. The nuclear envelope remains intact (arrows). Fig. 21. Diagrammatic
depiction of the metaphase plate of Fig. 20. Fig. 22. Early anaphase. Chromosomes appear to
diverge from the poles. Fig. 23. Late anaphase. The chromosomes are positioned parallel to
each other and some of them are still underway. The nuclear envelope is unbroken (arrows) and
has began to constrict itself in the middle portion. Fig. 24. Telophase. The middle constriction
of the nuclear envelope (arrows) has almost separate the daughter nuclei. The chromosomes are
beginning to disorganize. Fig. 25. Two daughter nuclei already in interphase. The nucleolus is
by this time not discernible.

184

Elisa

R. Parodi

and

Eduardo

J. Cdceres

Cytologia

61

of Chaetomorpha melagonium (Patel 1972), Cladophora surera (Parodi 1990) or Boodlea

composita (Kapraun 1993). Nuclear content appeared granular with a deep stained marginal
heterochromatic band against the nuclear envelope. In metaphase the nucleolus disorganized
and 36 chromosomes gathered in a very close equatorial plate (Figs. 20, 21). The nuclear
envelope remained intact (Fig. 20, arrows) during this and all the stages of the mitosis (Figs.
22, 24 arrows). Although its persistence is indubitable (Figs. 23, 24) and its behavior very
similar to those of Cladophora (McDonald and Pickett-Heaps 1976, Scott and Bullock 1976,
Parodi 1990) it would be necessary studies with electron microscope to find accuracy on the
behavior and also on the presence or absence of centrioles. During early anaphase the
chromosomes converged to the nuclear poles without absolute synchronism (Fig. 22). During
late anaphase they positioned themselves paralell to each other (Fig. 23) and the nuclear
envelope suffered a narrowing in the middle portion (Fig. 23, arrows). During early telophase
(Fig. 24) the chromosomes began to disorganize, the nuclear envelope increased the middle
constriction (Fig. 24, arrows) and the daughter nuclei began to separate. During late telophase
nuclei completed separation (Fig. 25) while the nucleolus was not yet visible.
Karyotype

The gametic chromosome dotation Ch. exilissima is n =18 showing median (m) and
submedian (sm) centromeres in a proportion, sm :m = 1 : 2, (Fig. 13) as normal in many species
of Cladophora (Sinha 1963, Wik-SjOstedt 1970, Kapraun and Gargiulo 1987a, b, Parodi and
Caceres 1995), Rhizoclonium (Parodi and Caceres 1992) and Chaetomorpha (Hinson and
Kapraun 1991, Kapraun 1993). Chromosome size extend from 0.7-1.4 ,um. Godward (1966)
has suggested 6 as the basic chromosome number in Cladophorales concept corroborated by
Kapraun (1993) and Parodi and Caceres (1992, 1995). Therefore, Ch. exilissima (n = 18) may
be interpreted as an apparent triploid n = 3x. Nevertheless, the proportion between these two
types of chromosomes is in Ch. exilissima different than in the rest of the studied species.
According with Wik-SjOstedt (1970) and Kapraun and Gargiulo (1987a) this proportion is in
Cladophora 1: 1, being also the case of Chaetomorpha (Hinson and Kapraun 1991) since the
basic dotation of six chromosomes includes three sm and three m chromosomes. Consequently,
Ch. exilissima could be better interpreted as an aneuploid n = 18= 4x -6 via reduction of the sm
dotation.
Taxonomy

Diagnostic characters. The zoosporangia of C exilissima are produced atter cytokinesis

from vegetative cells and thus differ in morphology and size from the latter. This is a new
characteristic for the genus since in all previously described species the zoosporangia are
identical to the vegetative cells. Also liberation of zoospores exclusively through two opposite
pores on lateral walls in an unusual attribute in the genus. It has been mentioned only for C.
melagonium Kutz. (Patel 1972) and C. aerea (Dillwyn) Kutz. (Sar 1985) but not as exclusive
location of the pores for zoospore liberation.
When we compare C. exilissima with those species described in the literature as fresh-water
inhabitants (Table 1) the most resembling but nonetheless clearly different is the exclusive
fresh-water species C. herbipolensis (Lagerheim 1887). In the latter species the zoospores are
normally liberated through only one pore in the zoosporangium (exceptionally through two or
even three but alternate) and zoosporangia are formed without a previous cytokinesis of
vegetative cells. The vegetative cells are normally shorter (75-255 gm), wider ( [36] 45-130
um in diameter) and with lower relation length: wide (1/2-4). Only in the appearance and the
dimensions of the basal cell do the two species bear any similarity. When comparing with the
marine species of Chaetomorpha reported to have slender filaments of variable diameter (Table

1996

Cytology

Table

1.

Comparison

of Chaetomorpha

of the Freshwater

exilissima

with

in fresh-water

Table

2.

Comparison

of the cell dimentions

slender

filaments

Alga

Chaetomorpha

exclusive

species

and with

other

found

environments

of Chaetomorpha
and

fresh-water

185

variable

filament

exilissima

with

marine

species

with

diameter

2) the most resembling is the long-celled C. cannabina (Areschoug.) Kjellman (Taylor 1960,
Blair 1983), but C. exilissima has even thinner and longer cells. The epiphytic C. minima
(Taylor 1960), has even thinner filaments, but the proportions of the cells dimensions and the
habit are very different. C. implexa Kiitz. (Kiitzing 1849), C. gracilis Kiitz. (Taylor 1960) are
also short-celled.
Habitat. This constitutes a new report of a freshwater species of Chaetomorpha on the
American continent. C. linum Kutz., C. implexa Kutz. (Bennett 1887) and C. gracilis Kutz.
(Carter 1919), have been mentioned as freshwater species, though they have been reported also
in brackish waters and were originally described as marine (Kutzing 1843, 1845, 1849). It is
doubtful whether the freshwater Chaetomorpha sp. mentioned for Argentina (Kiihnemann et
al. 1949) is in fact a Chaetomorpha representative since the basal cell described by the authors
lacks the for the genus characteristic disc-like holdfast (Boney 1966) but shows and elongated
rhizoid very similar to that of Rhizoclonium hieroglyphicum (Ag.) Kutz. (Parodi 1990, Parodi
and Caceres 1993).
We find sensible to hypothesize C. exilissima is an exclusive fresh-water species. It is
known that in regions close to the sea, marine organisms under certain conditions can colonize
fresh-water environments and this has been the case of certain Chaetomorpha species able to live
in both marine or brackish environments as C. aerea (Patel 1971, Sar 1985) and C. linum (Patel
1971). But this appears not the case, considering the isolation of the place in which the species
has been found and the absence of Chaetomorpha populations in the nearby marine environments. Moreover, the artificial pond filled with tap water in which C. exilissima growths has no
connection with open seas about 50 km far away. Freshwater species of Chaetomorpha are very
rare to the extent that in many phycological textbooks the genus is considered as exclusiverly

Elisa

186

marine

(Smith

1980).

Thus,

1950,

(Lagerheim
1887)

1955,

1887).
and

C.

freshwater

Fritsch

Chaetomorpha

has

Up

to

this

herbipolensis

species

in

Chaetomorpha,

mentioned

the

provided

illustration

R. Parodi

1956,

Bold

considered

time

only

and

C.

doubts

the

the

1985,
last

Mont.

1887)

have

Lagerheim

Conferva

author

Cytologia

1978,

since

blancheana

Although

with

J. Caceres

Wynne

(Lagerheim

genus.

by

Eduardo

been

Lager.

the

and

in

his

work

ansonii

recognizable

1981,

true

1856,

described

(1887),

a clearly

1975,
as

(Montagne
been

(Chaetomorpha?)

shows

Round

century

Lee

seeweed
Lagerheim

as

exclusively

on

freshwater

Ag. brevis

specimen

of

61

Nordst,

Microspora.

Diagnosis

Chaetomorpha
Aquae
sporophytum
interalia

2 flagellatibus,

Zoosporae

oviformes,
elabentes.

450 m

long,

Isogametae
lateralibus

Blanca.

70-120 m,

orificiis

locality:

30-50 m;
lat.

cellula

gm,

reticulatis.

adnatus,
hapteroidea

oviformes,

long
25

Gametophytum

obscure

200 m

stigma

juniorus

lat.,

cell.

Gametangia
chloroplastic

Thallus

numerosae,

Provincia

cell.

isomorphes.

multicellularia,

cell.

4-flagellatibus,

20 m

Holotvpus:
Type

lat.

sp. nov.

alternantes,

setiformia

disciformi.

oviformes,

orificiis

Fila

natans,

long

hapteroidea

et Caceres

Generationes

= 36).

libere

adnatus,

cellula

Parodi

hospites.

(2n
algas

juniorus

exilissima

dulcis

cell.
cellula

long,

40-55

basali

long,

60 m

long

10 m

cell.

2 flagellatibus,

Isogametae

7 gill
lat.

Gametangia

stigma

450 m

lat.

75-150 m
long,

70-120m,

disciformi.

(n
Thallus

250-500m,

Zoosporangia

notabili,

viridia.

notabili,

in

cell.

10 m

nucleatis,
20 m

lat.,

numerosae,
long,

diam.

lat.

oppositis

cellula

long,

long,

75 m
per

25 m;

200 m

=18),
adultus

basali

60 m

6 m

lat.,

lat.
per

elabentes.

Parodi

10, 18 May

Artificial
de

Buenos

1988 (BBB)

pond,

Parque

Aires.

Argentina

(Fig.

2).

Independencia,

Bahia

(3841'S.

6215'W).

Blanca,

partido

de

Bahia

Summary

The life history and cytology of the freshwater alga Chaetomorpha exilissima sp. nov.
(Cladophorales, Chlorophyta) has been studied in detail. The adult gametophyte (n =18) and
sporophyte (2n = 36), isomorphic plants were free-floating with multinucleate cells. Pairs of
zoosporangia were produced after one cytokinesis from vegetative cells and thus differed
morphologically from the latter. In all previously described species the zoosporangia are
identical to the vegetative cells. Liberation of zoospores occurs by two opposite pores in the
zoosporangium. The progress of the mitosis has been followed in sporophytic plants. The
present are the first complete observations of the mitosis in the genus Chaetomorpha. During
the process six stages could be distinguished: prophase, metaphase, early anaphase, late
anaphase, early telophase and late telophase. The nuclear envelope remained intact during all
the stages of the mitosis. The gametic chromosome dotation showed median (m) and
submedian (sm) centromeres. Ch. exilissima with a proportion, sm :m =1 : 2, could be
interpreted as an aneuploid n =18 = 4x - 6 via reduction of the sm dotation. Considering the
formation of zoosporangia by cytokinesis of the vegetative cell, the liberation of zoospores by
two opposite pores in the zoosporangium and the isolation of the place in which the species has
been found with absence of Chaetomorpha populations in the nearby marine environments, we
hypothesize this populations is representative of a new exclusive freshwater species. The most
resembling but nonetheless clearly different species is the exclusive freshwater C. herbipolensis.
Acknowledgments

This research was supported by the Consejo Nacional de Investigaciones Cientificas y

1996

Cytology

of the

Freshwater

Alga

Chaetomorpha

187

Tecnicas de la Republica Argentina (CONICET), grants "PID 3-110-500/88" and "PID 3111/
92. E.R.P. is research member of CONICET. E.J.C. is research member of the Comision de
Investigaciones Cientificas de la Provincia de Buenos Aires, Argentina (CIC).

References
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Bennet, A. W. 1887. Freshwater algae of north Cornwall. J. R. Mic. Soc. 2: 7-29.
Blair, R. 1983. Taxonomic treatment of the Chaetomorpha and Rhizoclonium species (Cladophorales, Chlorophyta) in
New England. Rhodora 85: 175-211.
Bold, H. C. and Wynne, M. J. 1978. Introduction to the Algae. Structure and Reproduction. Prentice Hall, New
Jersey, 706 pp.
-

1985 . Introduction

Boney,

A. D.

Carter,

N.

Cocucci,

to the Algae.

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