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Spore morphology of the representatives of the subfamily Ceratopteridoideae (J. Sm.) R.M. Tryon from the family Pteridaceae E.D.M. Kirchn. (Pteridophyta)

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p>Scanning electron microscopy (SEM) was used to perform a comparative study for four representatives of the subfamily Ceratopteridoideae (J. Sm.) R.M. Tryon from East Asia: Ceratopteris thalictroides Brongn., C. pteridoides (Hook.) Hieron., Acrostichum aureum Linn., and A. speciosum Willd. The analysis of the external morphology of the representatives of Ceratopteridoideae revealed a strong difference between Ceratopteris and Acrostichum . The external morphology of spores of Ceratopteris and Acrostichum exhibited the features characterizing the family Pteridaceae as a whole: tetrahedral spores with a three-ray laesure, from triangular-roundish to roundish, distinct exosporium with a pronounced surface ornamentation and absence of perisporium, the exosporium surface varying from smooth and rough to that covered with large tubercles and roller-like bulges. The external morphology of spores of Acrostichum is quite simple, although among other representatives of the family Pteridaceae, the spores of Ceratopteris have distinctive features: very large equatorial diameter (106–124 µm); spores are almost roundish; distinct cylindrical folds running parallel to the corners of the spore across its extensive distal surface. </p
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Ukrainian Journal of Ecology
Ukrainian Journal of Ecology,
2017, 7(2)
,
124-129,
doi:
10.15421/2017_29
ORIGINAL ARTICLE UDC 582.394
Spore morphology of the representatives of the subfamily
Ceratopteridoideae (J. Sm.) R.M. Tryon from the family Pteridaceae
E.D.M. Kirchn. (Pteridophyta)
A. V. Vaganov1, I. I. Gureyeva2, A. A. Kuznetsov2, A. I. Shmakov1, R. S. Romanets2, V. A. König3
1South-Siberian Botanical Garden, Altai State University
prospect Lenina, 61, Barnaul, 656049, Russia, E-mail: vaganov_vav@mail.ru
2Tomsk State University
prospect Lenina, 36, Tomsk, 634050, Russia. E-mail: gureyeva@yandex.ru
3 Scanware Electronic GmbH
Darmstädter Str. 9-11, Bickenbach, D-64404, Deutschland. E-mail: Viktor_Koenig@gmx.de
Scanning electron microscopy (SEM) was used to perform a comparative study for four representatives of the subfamily
Ceratopteridoideae
(J. Sm.) R.M. Tryon from East Asia:
Ceratopteris thalictroides
Brongn.,
C. pteridoides
(Hook.) Hieron.,
Acrostichum aureum
Linn., and
A.
speciosum
Willd. The analysis of the external morphology of the representatives of
Ceratopteridoideae
revealed a strong difference between
Ceratopteris
and
Acrostichum
.
The external morphology of spores of
Ceratopteris
and
Acrostichum
exhibited the features characterizing the family
Pteridaceae as a whole: tetrahedral spores with a three-ray laesure, from triangular-roundish to roundish, distinct exosporium
with a pronounced surface ornamentation and absence of perisporium, the exosporium surface varying from smooth and
rough to that covered with large tubercles and roller-like bulges.
The external morphology of spores of
Acrostichum
is quite simple, although among other representatives of the family
Pteridaceae,
the
spores of
Ceratopteris
have distinctive features: very large equatorial diameter (106–124 µm); spores are
almost roundish; distinct cylindrical folds running parallel to the corners of the spore across its extensive distal surface.
Key words: genus
Ceratopteris,
genus
Acrostichum,
subfamily
Ceratopteridoideae
, family Pteridaceae,
morphology of the
spores, scanning electronic microscopy (SEM).
Introduction
The representatives of
Ceratopteris
(4 species) and
Acrostichum
(23 species) are aquatic, semi-aquatic and helophytic ferns of
pantropical distribution (Nooteboom, 2012).
Due to high polymorphism inherent in the representatives of
Ceratopteris
, the genus consists of an infinite number of forms
restricted to different geographical regions. Therefore, more than 12 species were described within the genus. However, many
pteridologists believe that the genus comprises only one variable species of
C
.
thalictroides
(L.) Brongn. (Lloyd, 1974).
The representatives of
Ceratopteris
are completely immersed in the aquatic environment and prefer such habitats as bogs,
ponds, and wet ditches. On the contrary, the representatives of
Acrostichum
grow on the shores of the seas, mainly in mangrove
forests, where water composition is brackish (Tryon, Lugardon, 1991; Zhang et al., 2013).
The Flora of China includes four species of the subfamily
Ceratopteridoideae
(J. Sm.) R.M. Tryon:
Ceratopteris thalictroides
Brongn.,
C
.
pteridoides
(Hook.) Hieron.,
Acrostichum aureum
Linn., and
A
.
speciosum
Willd. (Zhang et al., 2013). Moreover,
Ceratopteris thalictroides
and
Acrostichum aureum
show fragmented pattern of distribution along the tropical belt throughout
all continents, except for Antarctica.
The species
Ceratopteris pteridoides
is prevalent mainly in Central, North and South America. In Asia, it is commonly found in
China (Anhui, Hubei, Jiangsu, Jiangxi, and Shandong), Bangladesh, India, and Vietnam. The species
Acrostichum speciosum
is
endemic to China (including Hainan), Malaysia, Thailand, Vietnam, and Australia (Zhang et al., 2013).
125
Spore morphology of the representatives of Ceratopteridoideae
Ukrainian Journal of Ecology, 7(2), 2017
In the current fern system developed using molecular genetics methods, the representatives of the genera
Ceratopteris
and
Acrostichum
exhibit an extremely high degree of relationship (Hasebe et al., 1996; Smith et al., 2006; Schuettpelz et al., 2007;
Liu et al., 2008; Schuettpelz, Pryer, 2008; Christenhusz, 2011; Schneider et al., 2013; Zhang et al., 2017).
However, the authors of early scientific works on the taxonomy of ferns emphasized the peculiarity of the external morphology
of the representatives of
Ceratopteris
and defined the genus as a monotype that forms an independent family
Parkeriaceae
Hook. (Copeland, 1947; Pichi-Sermolli, 1977). Pichi-Sermolli refers the genus
Acrostichum
to the family
Pteridaceae
(Pichi-
Sermolli, 1977). Later, Rolla Tryon assigns the status of the subfamily
Ceratopteridoideae
to the tribe
Ceratopterideae
J. Smith
(Tryon, 1986). The data on molecular phylogeny obtained for the representatives of
Ceratopteris
and
Acrostichum
, as well as
the peculiar ecology of their habitat, allow us to recognize an independent subfamily
Ceratopteridoideae
along with the four
ones included in the family Petridaceae. In order to reveal the features of the external morphology of spores for the
representatives of the subfamily
Ceratopteridoideae
, we examined six specimens, representatives of four species from East
Asia (China, including Hong Kong and Hainan Island).
The spore morphology of the representatives of the subfamily
Ceratopteridoideae
has been poorly studied. Particularly relevant
is obtaining data on the external morphology of spores of Asian
Ceratopteris
and
Acrostichum
.
The SEM method was used to analyze the spore morphology of the representatives of
Ceratopteris
on specimens from Africa
(Liberia and Salvador), South America (Peru and Brazil), Cuba, India, and Australia
C
.
cornuta
(P.Beauv.) Lepr.,
C
.
pteridoides
,
C
.
thalictroides
, and
C
.
richardii
Brongn. (Tryon, Lugardon, 1991; Dettmann, Clifford, 1991). Similar to the studies of the spore
morphology conducted last century (Nayar, 1968; Sahashi, 1979), micrographs of
Ceratopteris
have few exposures of
microphotographs. In addition, poor quality of micrographs does not allow a detailed description of the external morphology
of spores that does not provide sufficient data and complete analysis of the exosporium surface. The study of the spore
morphology of
Acrostichum
was limited to specimens from tropical America (Mexico, Honduras and Guyana), Africa (Liberia,),
Southeast Asia (Philippines and Cambodia), and Fiji
A
.
danaeifolium
Langsd. a. Fisch., and
A
.
aureum
(Tryon, Lugardon, 1991).
Material and Methods
The spores for the study were selected from the herbarium material deposited in the Herbarium of the Institute of Botany of
the Academy of Science of China (PE, Beijing). The spores were fixed on a slide using an electrically conductive adhesive tape,
sputtered with gold using a sputter coater Quorum Q150R S, and examined under a scanning microscope Mini-SEM SNE-4500M
(Korea) located in the laboratory of structural and molecular analysis of plants, TSU. The spores surface was scanned in high
vacuum at voltage of 1030 kV and magnified from 2,500 to 10,000 times. The spore dimensions and the element
ornamentation were determined using photographs taken with a scanning microscope.
Results and Discussion
The description and original electron micrographs of spores of the species
Ceratopteris
and
Acrostichum
are provided below.
1.
Ceratopteris thalictroides
Brongn. (Fig. 1, ae). Spores in the proximal-polar and distal-polar positions are triangular-roundish
to roundish. The equatorial diameter is (101) 112.8 (124) μm. The polar axis is 101.7 (from 92.3 to 110) μm. In the equatorial
position, the distal side of the spore is almost spherical, the proximal side is expressed only at the apex flattened. The rays of
laesura are straight (24) 28.7 (32.4) μm long, (4.45) 4.62 (4.95) μm wide, rising above the sporoderm. Roller-like folds running
from the spore corners across the extensive distal surface of the spore are highly risen above the sporoderm (up to the height
of laesuras) and reach the width of (3.47) 3.83 (4.29) μm. The distance between the adjacent roller-like folds is (3.0) 5.11 (7.4)
μm. In the depressions between roller-like folds and laesuras, the exosporium surface is uniformly covered with elongated
excrescences of (0.237) 0.312 (0.379) μm in diameter.
Investigated specimen: Deep Bay, Hong Kong. Edge of rice field. Shiu Ying Hu 5777. Oct. 6, 1968. 00587547.
2.
Ceratopteris pteridoides
(Hook.) Hieron. Christ (Fig. 2, ae). Spores in the proximal-polar and distal-polar positions are
triangular-roundish to roundish. The equatorial diameter is (106) 114.8 (121) μm. The polar axis is (92) 93.4 (96) μm. In the
equatorial position, the distal side of the spore is almost spherical, the proximal side is expressed only at the apex flattened.
The rays of laesura are straight (30.5) 32.6 (34.5) μm long, (2.6) 3.2 (4.2) μm wide, rising above the sporoderm. Cylindrical folds
running from the spore corners across the extensive distal surface of the spore are highly risen above the sporoderm (up to
the height of laesuras) and reach the width of (3.3) 3.73 (4.5) μm. The distance between the adjacent roller-like folds is (2.8) 5.84
(10) μm. In the depressions between roller-like folds and laesuras, the exosporium surface is uniformly covered with tubercles
of (0.163) 0.23 (0.317) μm in diameter.
Investigated specimen: Vicinity of Hanyang, Hupeh. Alt. 50. Habitat in water. Habit herb or water fern. S.W. Teng №302. Nov. 10,
1932. №00587529.
3.
Acrostichum aureum
Linn. (Fig. 3, a, b). Spores in the proximal-polar and distal-polar positions are triangular-roundish, non-
lociniate. The equatorial diameter is (48) 50 (52) μm. The rays of laesura are straight, (14.7) 16.3 (18.2) μm long, (2.2) 2.44 (2.9)
μm wide, slightly risen at the apex above the surface of the spore. The exosporium surface on the distal and proximal sides is
covered with slightly protruding tubercles of (1.0) 1.28 (1.5) μm in diameter, rough.
Investigated specimen: Fairly common; moist, level land, sandy soil, seashore, semi-woody, erect. Shan Mong, Yai-hsien District.
Coll. Lau S.K. 5998. Flora of Hainan, 12th Hainan Expedition.
Ukrainian Journal of Ecology, 7(2), 2017
Ukrainian Journal of Ecology
126
Fig. 1. SEM views of spores of
Ceratopteris thalictroides:
a proximal side; b distal side; c
,
d spore in equatorial position; e
,
f fragment of proximal surface.
4.
Acrostichum speciosum
Willd. (Fig. 4, a-c). Spores in the proximal-polar and distal-polar positions are triangular-roundish,
non-lociniate. The equatorial diameter is (38) 46.6 (51.8) μm. The rays of laesura are straight (9.9) 10.7 (11.7) μm long, (0.8) 1.1
(1.3) μm wide, slightly risen at the apex above the surface of the spore. The exosporium surface on the distal and proximal sides
is covered with slightly protruding tubercles of (0.3) 0.64 (0.9) μm in diameter, rough.
Investigated specimen: Hainan, Wenchang, Qinglan Reserve 19о37.30`N, 110о44.79`E. Habitat: on ground in mangrove forests.
Alt.: 0 m. Coll. A. No.: S.Y. Dong a. X.C. Zhang 682. 2003-03-17. №00543730.
127
Spore morphology of the representatives of Ceratopteridoideae
Ukrainian Journal of Ecology, 7(2), 2017
Fig. 2. SEM views of spores of
Ceratopteris pteridoides:
a proximal side; b distal side; c
,
d spore in equatorial position; e
,
f
fragment of proximal surface.
The analysis of the external morphology of the spores of
Ceratopteridoideae
revealed a few features characterizing the family
Pteridaceae: tetrahedral spores with a three-ray laesura, triangular-roundish to roundish, distinct exosporium with expressed
ornamentation, without perisporia, the exosporium surface varying from smooth and rough to that covered with large tubercles
and roller-like bulges. A strong difference in the external morphology of spores of
Ceratopteris
and
Acrostichum
has been
revealed. Thus, the distinctive features of the external morphology of spores among the genera of the subfamily
Ceratopteridoideae
vary within a range that is much greater if compared to the subfamily
Cryptogrammoideae
S. Linds., which
also includes a small number of genera (
Coniogramme, Cryptogramma
and
Llavea
). The other three subfamilies of the family
Pteridaceae include from 12 to 20 genera, and this explains a strong variation in the morphological features of spores.
Ukrainian Journal of Ecology, 7(2), 2017
Ukrainian Journal of Ecology
128
Fig. 3. SEM views of spores of
Acrostichum aureum:
a spore in proximal-equatorial position; b fragment of proximal surface.
The external morphology of spores of
Acrostichum
is quite simple. Its primitive structure makes the morphology similar to that
of the representatives of various subfamilies of the family Pteridaceae. The morphology of spores of
Acrostichum
resembles
the shape and nature of the spore surface of the representatives from the genera
Pellaea
Link,
Adiantum
L.,
Coniogramme
Fée,
Llavea
Lag., and
Anopteris
(Prantl) Diels (Tryon, Lugardon, 1991; Vaganov et al., 2010; 2011; Vaganov, 2016).
Fig. 4. SEM views of spores of
Acrostichum speciosum: а
spore in proximal-equatorial position; b distal side; c
fragment of
distal surface.
On the contrary, the external morphology of spores of
Ceratopteris
is extremely original and not similar to that of the
representatives of the family Pteridaceae. In addition, fairly large overall dimensions of the equatorial diameter of spores of the
129
Spore morphology of the representatives of Ceratopteridoideae
Ukrainian Journal of Ecology, 7(2), 2017
representatives of the genus
Ceratopteris
, the distinctive features of the external morphology and very peculiar ecology of the
fern habitat show its unique position in the system of the family Pteridaceae.
The study conducted using the scanning electron microscopy revealed no affinity between the ferns of the genera
Ceratopteris
and
Acrostichum
since a small number of similar features can be observed in the external morphology of spores.
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Citation:
Vaganov, A.V., Gureyeva, I.I., Kuznetsov, A.A., Shmakov, A.I., Romanets, R.S., König, V.A. (2017). Spore morphology of the representatives of the
subfamily Ceratopteridoideae (J. Sm.) R.M. Tryon from the family Pteridaceae E.D.M. Kirchn. (Pteridophyta).
Ukrainian Journal of Ecology, 7
(2), 124129.
This work is licensed under a Creative Commons Attribution 4.0. License
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The work presents the result of using comparative morphological analysis of spore of three subfamilies from the family Pteridaceae E. D. M. Kirchn. (Cryptogrammoideae S. Linds., Pteridoideae C. Chr. ex Crabbe, Jermy a. Mickel, Ceratopteridoideae (J. Sm.) R. M. Tryon) by evolutionary biology method by Mesquite software. This approach allows comparing in detail the standardized qualitative features of the morphology of spores with the data on phylogenetic relationship of representatives, which as a result helps to identify intra-and intertaxonomic groups of relationship, to search for evolutionary connections, to obtain new knowledge for solving problems in fern taxonomy. Obtained data allow supplement natural classification of the Pteridaceae family. Введение. // В работе представлен результат применения сравнительно-морфологического анализа спор представителей трех подсемейств семейства Pteridaceae E. D. M. Kirchn. (Cryptogrammoideae S. Linds., Pteridoideae C. Chr. ex Crabbe, Jermy a. Mickel, Ceratopteridoideae (J. Sm.) R. M. Tryon) методом эволюционной биологии в мо-дульной программной среде Mesquite. Данный подход позволяет детально сопоставить стандартизованные каче-ственные признаки морфологии спор с данными о филогенетическом родстве представителей, что в итоге спо-собствует выявлению внутри-и межтаксономических групп родства, поиску эволюционных связей, получению новых знаний для решения проблем в таксономии папоротников. Полученные данные позволят дополнять естественную классификацию семейства Pteridaceae. Ключевые слова. ДНК, морфология, молекулярная филогения, папоротники, систематика, сканирующая электронная микроскопия, эволюция растений, Ceratopteridoideae, Cryptogrammoideae, Pteridaceae, Pteridoideae.
... Spore morphology of species of the large fern family Pteridaceae has been discussed in the numerous works (Hennipman, 1975;Tryon, Lugardon, 1991;Palacios-Rios et al., 2016;Chao, Huang, 2018;Irfan et al., 2021) including ours (Kuznetsov et al., 2014;Vaganov, 2016;Vaganov et al., 2017aVaganov et al., -d, 2018a (Gureyeva, Kuznetsov, 2015;Vaganov et al., 2017a-c). Data of spore morphology of some species of Austrogramme, Syngramma, and Taenitis obtained using scanning electron microscopy were published in 1991 by A. Tryon and B. Lugardon (11 species in sum). ...
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... Comparison between SEM spore sizes retrieved from Nigeria (this study) versus those reported in Asia. We compared our records with the following records from Asia by Genus -Acrostichum (Vaganov et al. 2017;China); Ceratopteris (Vaganov et al. 2017;China); Ctenitis (no record found); Dryopteris (Lee & Park 2014;Korea); Lygodium (Haq 2017;Pakistan); Nephrolepis (Kao et al. 2014;Taiwan); Phymatosorus (Sofiyanti 2019;India); Pityrogramma (Vaganov et al. 2017b;China); Pteris (Chao & Huang 2018;China With only one species sampled from the families Lygodiaceae (Lygodium microphyllum), Nephrolepidaceae (Nephrolepis biserrata), and Polypodiaceae (Phymatosorus scolopendria), the intra-generic spore characteristics could not be explored. Nevertheless, our study contributed to the existing spore description for these genera. ...
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... It was shoved on numerous example that spore morphology and molecular-phylogenetic results are congruent for different genera such as Gymnocarpium and Cystopteris (Gureyeva, Kuznetsov, 2015), Onichium (Vaganov et al., 2017 a, b, c), Pityrogramma (Vaganov et al., 2017b) therefore the morphology of spores has phylogenetic signification. Spore morphology is very important for systematics of Pteridaceae subfamilies: Cryptogrammoideae (Vaganov, 2016) and Ceratopteridoideae (Vaganov et al., 2017c). ...
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