Upload
andrea-tucci
View
216
Download
1
Embed Size (px)
Citation preview
SYSTEMATICS, PHYLOGENY AND FLORISTICS
New planktic species of Kirchneriella Schmidle (Chlorophyceae,Selenastraceae) from Brazilian freshwaters
Daniella da Silva • Celia L. Sant’Anna •
Andrea Tucci • Augusto Comas
Received: 30 July 2012 / Accepted: 1 April 2013 / Published online: 16 July 2013
� Botanical Society of Sao Paulo 2013
Abstract New planktic species of Kirchneriella Schm-
idle (Chlorophyceae, Selenastraceae) from Brazilian
freshwaters. In the samples collected from an artificial lake,
located in the metropolitan region of Sao Paulo, an inter-
esting algal population morphologically similar to Kirch-
neriella Schmidle was found. The crescent-shaped cells
were oriented, inside the colony, with their convex faces
toward the periphery of the colony resembling those of K.
roselata Hindak. However, the remaining portions of the
mother cell wall with more or less radial arrangement
similar to the connective mucilaginous threads of the genus
Selenodictyon Uherkovich & Schmidt ex Comas & Ko-
marek were also observed. After further studies based on
light microscope, the studied material proves to be mor-
phologically distinguishable from K. roselata, although it
never forms rosette-like groups, which are typical in K.
roselata. The remaining parts of the mother cell wall more
or less radially disposed near the cells (autospores just
released from the mother cell wall) are not connective
threads. Thus, a new species, K. brasiliana, is described in
this paper. According to literature, the main diagnostic
feature of Selenodictyon is the crescent-shaped cells
attached by their concave faces to mucilaginous stalks like
in Dictyosphaerium; however, remaining portions of the
mother cell wall more or less radially oriented were also
formed in some Kirchneriella populations. Therefore, the
authors consider that such feature could be misinterpreted
as connective mucilaginous threads, which raises doubts
regarding the validity of Selenodictyon.
Keywords Green algae � Fish pound � New species �Taxonomy
Introduction
Kirchneriella Schmidle 1893 was traditionally included in
the families Ankistrodesmaceae (Korsikov 1953), Chlo-
rellaceae (Komarek and Fott 1983), or in Selenastraceae
(Marvan et al. 1984), belonging to Chlorococcales. Recent
studies based on gene sequence analysis (Krienitz et al.
2001) show phylogenetic relationships with Sphaeropleales
sensu Deason et al. (1991). These studies also showed that
the family Selenastraceae constitutes a monophyletic group
belonging to the class Chlorophyceae (Krienitz et al. 2011).
In spite of the fact that the morphological features do not
correspond to molecular basis for distinguishing genera in
the Selenastraceae (Fawley et al. 2005), the genus Kirch-
neriella, in the traditional sense (as genus forma), is well
characterized by arcuate, crescent-shaped, or cylindrical-
fusiform (irregularly curved) and even sigmoid cells, dis-
posed in colonial mucilage. The cell ends are sharp, pointed
round, pointed, or round. The cells inside the colony can be
organized with their convex side turned to the colony center
or turned to the colony edge. The parietal chloroplast occu-
pies a large portion of the cell volume and pyrenoids may be
present or not. Reproduction occurs by autospores disposed
in series inside the mother cell wall or, more rarely, more or
less in parallel arrangement. Autospores are liberated by the
rupture of the mother cell wall, which later will be com-
pletely or partially gelatinized. In this last case, the remnants
of the mother cell wall remain inside the colonial mucilage.
D. da Silva � C. L. Sant’Anna (&) � A. Tucci
Instituto de Botanica, Nucleo de Pesquisa em Ficologia,
Sao Paulo, SP, Brazil
e-mail: [email protected]
A. Comas
Centro de Estudios Ambientales de Cienfuegos, Ministerio de
Ciencias, Tecnologıa y Medio Ambiente, Cienfuegos, Cuba
123
Braz. J. Bot (2013) 36(2):153–157
DOI 10.1007/s40415-013-0014-5
The presence or absence of the pyrenoid (under the light
microscope) was traditionally considered a good diacritic
feature, allowing the classification of species into two
independent genera: (1) Kirchneriella s. str., with pyrenoid
(type species K. obesa (W. West) Schmidle 1893); (2)
Pseudokirchneriella Hind. 1990, without pyrenoid (=Kir-
chneria Hindak 1988, Raphidocelis Hindak 1977 sensu
Marvan et al. 1984).
There is no precise designation of the type species of
Pseudokirchneriella, and for this reason we are considering
Pseudokirchneriella subcapitata (Korsikov) Hindak 1990
as a type species, based on the fact that Kirchneria, whose
type species is K. subcapitata (Hindak 1988), is considered
to be synonymous with Pseudokirchneriella.
According to molecular studies (Krienitz et al. 2001;
Krienitz and Bock 2012), small genera based only on
morphological features are not sustainable, and the
adoption of only one genus (Kirchneriella) in broad sense
could be the correct option. Other features such as the
presence or absence of a pyrenoid, single cells or in col-
onies, and autospores arrangement inside the mother cell
wall are not enough to distinguish the mentioned genera
separately.
In the samples collected from an artificial lake, loca-
ted in the metropolitan region of Sao Paulo City, Brazil,
an interesting algae, morphologically similar to Kirch-
neriella roselata Hindak, was found. The cells are ori-
ented with their convex faces toward the periphery of the
colony and after reproduction (autospores liberated by
rupture of mother cell wall), the remnants of the mother
cell walls remain more or less radially oriented inside
the colonial mucilage looking like connective mucilagi-
neous stalks similar to those of Selenodictyon brasiliense
Uherkovich & Schmidt ex Comas & Komarek. This
algal population was studied in order to reach its accu-
rate taxonomical definition and clarify its relations to
other similar taxa.
Materials and methods
The studied populations were collected from an artificial
lake used for fishing (some limnological parameters are
shown in Table 1), located in the City of Suzano, Sao
Paulo State, southeastern Brazil (S23�2502400 W46�2200500).The samples were obtained using van Dorn’s bottle
and plankton net (20 lm), in dry and rainy seasons in a
year.
The samples were preserved with 4 % formaldehyde
and studied in a Zeiss Axiosplan 2 microscope. China ink
was used to improve the mucilage observation. The clas-
sification systems of Komarek and Fott (1983) and Krienitz
et al. (2001) were adopted.
Results and discussion
Kirchneriella brasiliana sp. nov.
Diagnosis: Coloniae libere natantes, irregulariter sphaericae
usque ovales, multicellulares, praecipue per turmas 2–4
cellulares aggregatas circum superficiem coloniae formatas,
cum latis concavis ad centrum coloniae dispositae. Tegu-
mentum gelatinosum, homogeneum, tenues, hyalinum, sine
structura. Cellulae fusiformes, arcuatae, moderate contor-
tae, ad utrosque polos aequaliter angustatae, apicibus acutis.
Membrana cellularia levis, hyalina. Chloroplastum unum,
parietale, sine pyrenoide. Propagatio 2–4 autosporis; auto-
sporae e membrana matricali divisione centrali in partes
duas liberantur; frequenter vestigia membranae matricali
circa cellulae juveniles remanens, ut videtur similis con-
nectivis, re vera autem, e cellulis distantibus. Dimensiones
cellularum: 11.3–17.0 9 1.2–3.1 lm.
Habitatio (locus classicus): in plancto lacus, Suzano,
Brasilia.
Holotypus: Brazil, Sao Paulo State: Suzano, 12-III-
2002, D Silva s.n. (SP3913443).
Description
Kirchneriella brasiliana sp. nov. (Fig. 1)
Colonies free floating, rounded to ovoid, formed by groups
of 2–4 cells arranged in the colony periphery; cells lunate,
pointed at both ends, concave side turned to the center of
Table 1 Physical, chemical, and biological variables of the studied
lake, Sao Paulo State, Brazil (according to Mercante et al. 2004,
2005)
Variables Collecting
Dry season Rainy season
Water temperature (�C) 21.9 30.0
Deep (m) 1.0 1.0
Euphotic zone (m) 0.6 0.6
Conductivity (lS cm-1) 30.0 30.0
pH 5.8 5.6
Dissolved oxygen (mg L-1) 6.1 11.0
Turbidity (NTU) 78.0 72.0
N–NO2- (lg L-1) 7.0 9.1
N–NO3- (lg L-1) 58.3 45.5
N–NH4? (lg L-1) 86.0 29.0
P–PO4- (lg L-1) 27.5 14.5
PT (lg L-1) 161.7 162.3
Chlorophyll a (lg L-1) 68.7 106.5
Trophic state index 69 69
154 D. da Silva et al.
123
colony; mucilaginous envelope thin, difluent; chloroplast
parietal, without pyrenoid; reproduction by autospores;
remaining parts of the mother cell wall in the center of the
colony, displaying radial orientation near the cells. Cells
length: (11.3–) 14.7 lm (–17). Cells diameter: (1.2–)
2.0 lm (–3.1).
Locality: Brazil, Sao Paulo, Municipality of Suzano,
fish-pond, plankton material, 08-X-2001 (dry season),
D Silva s.n. (SP3913442); 12-III-2002 (rainy season),
D Silva s.n. (SP391343).
The Brazilian material is morphologically similar to
K. roselata described from Eslovaquia (Hindak 1984), and
also registered in Ukraine (Tsarenko 1990) and Bulgaria
(Stoyneva 1998). K. roselata is an infrequently observed
species in the plankton of waters with different trophic
conditions (Stoyneva 1998). However, this taxon is
characterized by cells sharply curved, thin, normally joined
at one end, forming rosette-like groups, which present their
concave faces toward the center of the colony. When
they are present, remnants of the mother cell wall, par-
tially gelatinized, are scarce and spread without a radial
disposition.
Actually, the studied material is morphologically related
to this species, but the cells of the studied colonies never
form rosette-like groups, which are typical in K. roselata
and remnants of the mother cell wall, more or less radially
oriented in colonial mucilage, are usually present.
The Brazilian population is also similar to Selenodictyon
brasiliense when considering the orientation of the cells in
the colony, but in particular, because of the remaining
portions of the mother cell wall that present more or less
radial disposition near the cells (autospores just released).
Fig. 1 Kirchneriella brasiliana. 1, 2 General aspect of the colonies showing the cells disposition. 3 Detail of mother cell wall rests radially
disposed (arrow). 4 Scheme of one colony with mother cell wall rests (arrow). Bar 10 lm
New planktic species of Kirchneriella 155
123
However, they are not connective threads which are
obligatory in Selenodictyon, according to its diagnosis
(Comas and Komarek in Comas 1992).
Considering all taxonomical characteristics of the
studied population, we concluded that the material could be
considered as an independent species of Kirchneriella
(Table 2 shows the main morphological features of the
species compared with other related taxa).
Due to the similarity of the new species with S. brasiliense,
some taxonomic comments are necessary: the genus Sele-
nodictyon, with its type species S. brasiliense, was pub-
lished improperly by Uherkovich and Schmidt (1974)
(nom. nud.), based on plankton samples from Lago Cas-
tanho, in the State of Amazon, north of Brazil.
Supported by the work of van der Heide (1982) based on
samples from Lake Brokopondo, Suriname, as well as by
materials from Cuba that could correspond to Selenodict-
yon, Comas and Komarek (Comas 1992) validated the
genus Selenodictyon. With another concept, they grouped
in this genus some species with elongated to crescent-
shaped cells, two or more times longer than wide, attached
by their concave faces to mucous filiform stalks similar to
those of Dictyosphaerium Nageli. The species have one
parietal chloroplast with or without pyrenoid. Selenodictyon
brasiliense presents pyrenoid, according to the illustration of
Uherkovich and Schmidt (1974).
According to this conception, the genus Selenodictyon
was represented by three species: S. brasiliense, S. navicu-
liforme Comas & Komarek and S. elongatum (Hindak)
Comas & Komarek.
Taking into account the presence of Kirchneriella popu-
lations in materials from Cuba and Mexico (Veracruz), in
which crescent-shaped cells are disposed similarly to those
of Selenodictyon, with remaining portions of the mother
cell wall more or less radially disposed inside the colonial
mucilage, which could be misinterpreted as connective
threads similar to Dictyosphaerium/Selenodictyon, Comas
and Perez Baliero (2002) pointed out serious doubts in
relation to the existence of the genus Selenodictyon,
keeping, ‘‘ad interim’’, only one species: S. brasiliense.
According to the authors, the other Selenodictyon species
were not justified.
After our observations in Brazilian samples from Sao
Paulo State, we also consider that Selenodictyon is not a
good genus, and possibly it could be a particular species of
Kirchneriella s.l., whose cells’ concave face is disposed
toward the center of the colony. The remnants of the
mother cell wall, which are radially disposed inside the
Table 2 Comparison of the diacritic features of K. brasiliana and related species of the genera Selenodictyon and Kirchneriella
Selenodictyon
brasiliense
Uherk. &
Schmidt
(Uherkovich and
Schmidt 1974)
Selenodictyon
brasiliense
Uherk. &
Schmidt
(Komarek and
Fott 1983)
Selenodictyon
brasiliense
(Uherk. and
Schmidt) Com. &
Kom. (Comas
1992)*
Kirchneriella roselata
Hind. (Hindak 1984)
Kirchneriella
irregularis G.M.
Smith (Korsikov
1953)
Kirchneriella
brasiliana sp.
nov.
Cell form Lunate, pointed at
both ends
Lunate, pointed at
both ends
Lunate, pointed at
both ends
Fusiform, curved, slightly
sigmoid
Fusiform,
curved,
regularly
attenuated to
the bluntly
rounded or
obtuse ends
Lunate, pointed at
both ends
Cells
distribution
in the
colony
Groups of 2–4
cells arranged
in the colony
periphery;
concave side
turned to the
center of colony
Groups of 2–4
cells arranged
in the colony
periphery;
concave side
turned to the
center of colony
Groups of 2–4
cells arranged
in the colony
periphery;
concave side
turned to the
center of colony
Groups of 2–4 cells with
their ends close to each
other, forming rosette-
like groups; concave
side of the cells turned
to the center of colony
Groups of 4 cells
irregularly
disposed in the
mucilage; cells
with the
convex side
outwards
Groups of 2–4
cells arranged
in the colony
periphery;
concave side
turned to the
center of colony
Chloroplast
and
pyrenoid
_ Chloroplast
parietal with
one pyrenoid
(according to
the figures)
Chloroplast
parietal with
one pyrenoid
Chloroplast parietal
without pyrenoid
Chloroplast
parietalprobably with
one pyrenoid
Chloroplast
parietal without
pyrenoid
Cell length 6.5–7.0 lm 6.5–7.0 lm 6.5–7.0 lm 7.5–12 lm 6.0–21.0 lm (11.3) 14.7 lm
(17.0)
Cell wide 1.5 lm 1.5 lm 1.5 lm 1.5–2.0 lm 3.0–6.0 lm (1.2) 2.0 lm (3.1)
Mucilage _ Thin, hyaline Thin, hyaline Homogenous, hyaline _ Thin, hyaline
* Valid publication according to Botanical Code of Nomenclature
156 D. da Silva et al.
123
colony, must have been erroneously considered as con-
nective threads by Uherkovich and Schmidt (1974) and
Comas and Komarek (Comas 1992).
However, it is not possible to transfer S. brasiliense to
the genus Kirchneriella as a new combination because the
holotype of S. brasiliense is just a figure (Uherkovich and
Schmidt 1974, Taf. IV, Fig. 84), in which it is possible to
observe cells attached to the mucilaginous threads. As there
is no preserved sample of S. brasiliense, it is not possible to
confirm whether these ‘‘mucilaginous threads’’ were mis-
interpreted. Because of that and the presence of pyrenoids
in Fig. 84 that never occur in the material from Sao Paulo
State, it is impossible to consider S. brasiliense as a syno-
nym of K. brasiliana. The real taxonomic position of
S. brasiliense will be defined only if other populations are
found again.
References
Comas A (1992) Beitrage zur Kenntnis der Botryococcaceae (Chlo-
rellales) aus Kuba. Algol Stud 66:17–23
Comas A, Perez Baliero MC (2002) Chlamydophyceae (Chlorophy-
ceae) from Merin lagoon (Brazil-Uruguay, South America) with
special references to the Family Botryococcaceae. Algol Stud
107:49–65
Deason TR, Silva PC, Watanabe S, Floyd GL (1991) Taxonomic
Status of the species of the green algal genus Neochloris. Plant
Syst Evol 177:213–219
Fawley MW, Dean ML, Dimmer SK, Fawley KP (2005) Evaluating
the morphospecies concept in the Selenastraceae (Chlorophy-
ceae, Chlorophyta). J Phycol 42:142–154
Hindak F (1977) Studies on the Chlorococcal algae (Chlorophyceae)
I. Biologicke Prace. Bratislava 23:190
Hindak F (1984) Studies on the Chlorococcal algae (Chlorophyceae)
III. Biologicke Prace. Bratislava 34:264
Hindak F (1988) Studies on the Chlorococcal algae (Chlorophyceae)
IV. Biologicke Prace. Bratislava 34:264
Hindak F (1990) Studies on the Chlorococcal algae (Chlorophyceae)
V. Biol. Prace. Bratislava 36:225
Komarek J, Fott B (1983) Chlorococales. In: Hubber-Pestalozzi G
(ed) Phytoplankton des Susswassers, Systematik u. Biologie.
Teil 7, Das Stuttgart, pp 1–1044
Korsikov OA (1953) Pidklas Protokokovi (Protococcinae), Viznacnik
prisnovod. Vodorostej Ukrainskoi RSR. Akad Nauk URSR
5:1–439
Krienitz L, Bock C (2012) Present state of the systematics of
planktonic coccoid green algae of inland water. Hydrobiologia
698:295–326
Krienitz L, Ustinova I, Friedl T, Huss VAR (2001) Tradicional
generic concept versus 18S rRNA gene phylogeny in the green
algal family Selenastraceae (Chlorophyceae, Chlorophyta).
J Phycol 37:852–865
Krienitz L, Bock C, Nozaki H, Wolf M (2011) SSU rRNA gene
phylogeny of morphospecies affiliated to the bioassay alga
‘‘Selenastrum capricornutum’’ recovered the polyphyletic origin
of crescent-shaped chlorophyta. J Phycol 47:880–893
Marvan P, Komarek J, Comas A (1984) Weighting and scaling of
features in numerical evaluation of coccal green algae (genera of
the Selenastraceae). Algol Stud 37:363–399
Mercante CTJ, Cabianca MA, Silva D, Costa SV, Esteves KE (2004)
Water quality in fee-fishing ponds located in the metropolitan
region of Sao Paulo city, Brazil: an analysis of the eutrophication
process. Acta Limnol Bras 16:95–102
Mercante CTJ, Costa SV, Silva D, Cabianca MA, Esteves KE (2005)
Qualidade da agua em pesque-pague da regiao metropolitana de
Sao Paulo (Brasil): avaliacao atraves de fatores abioticos
(perıodo seco e chuvoso). Acta Sci Biol Sci 27:1–7
Stoyneva MP (1998) On the distribution of Pseudokirchneriella
danubiana and P. roselata in Bulgaria. Biologia 53:433–435
Tsarenko PM (1990) Short key of the Chlorococcalean algae of the
Ukrainian SSR. Naukova dumka, Kiev
Uherkovich G, Schmidt A (1974) Phytoplankton taxa in dem zentral-
amazonischen Schwemmlandsee Lago do Castanho. Amazoni-
ana 5:243–283
van Der Heide J (1982) Lake Brokopondo. Filling phase limnology
of a man-made lake in the humid tropics. Univ. Amsterdam,
Amsterdam
New planktic species of Kirchneriella 157
123