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Spore morphology of Haplopteris C. Presl species (Vittarioideae, Pteridaceae) from China

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A comparative study of spores of four species of Haplopteris C. Presl, H. amboinensis (Fée) X.C. Zhang, H. forrestiana (Ching) E.H. Crane, H. linearifolia (Ching) X.C. Zhang, and H. mediosora (Hayata) X.C. Zhang, from China was performed using the method of scanning electronic microscopy (SEM). Spores of Haplopteris species are bilateral monolete ellipsoidal or ellipsoidal, but slightly narrowed in the middle. Perispore thin, easily breakable, smooth; exospore smooth, its surface is finely granulate or finely undulate (as seen at higher magnification). Haplopteris amboinensis have the largest spores (79.5×34.3×40.9 μм), H. mediosora – the smallest one 49.2×24.5×24.9 μм).
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Ukrainian Journal of Ecology
Ukrainian Journal of Ecology,
2017, 7(4)
,
290294,
doi:
10.15421/2017_118
ORIGINAL ARTICLE 582.394
Spore morphology of
Haplopteris
C. Presl species
(
Vittarioideae
,
Pteridaceae) from China
A.V. Vaganov1, I.I. Gureyeva2, A.A. Kuznetsov2, R.S. Romanets2
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
Submitted: 02.11.2017. Accepted: 14.12.2017
A comparative study of spores of four species of
Haplopteris
C. Presl,
H. amboinensis
(Fée) X.C. Zhang
, H. forrestiana
(Ching)
E.H. Crane,
H. linearifolia
(Ching) X.C. Zhang, and
H. mediosora
(Hayata) X.C. Zhang, from China was performed using the
method of scanning electronic microscopy (SEM). Spores of
Haplopteris
species are bilateral monolete ellipsoidal or ellipsoidal,
but slightly narrowed in the middle. Perispore thin, easily breakable, smooth; exospore smooth, its surface is finely granulate
or finely undulate (as seen at higher magnification).
Haplopteris amboinensis
have the largest spores (79.5×34.3×40.9 µм),
H.
mediosora
the smallest one 49.2×24.5×24.9 µм).
Key words:
Haplopteris, Vittarioideae;
Pteridaceae;
spore morphology; scanning electronic microscopy (SEM)
The genus
Haplopteris
C. Presl was re-defined by E.H. Crane (1997) to accommodate nearly all the Old World species formerly
recognised as ‘
Vittaria
’ (Lindsay, 2003). M. Christenhusz et al. (2011) included the genus
Haplopteris
together with eleven more
genera (
Adiantum
L.
, Ananthacorus
Underw. & Maxon
, Anetium
Splitg.,
Antrophyum
Kaulf.
, Hecistopteris
J. Sm.,
Monogramma
Schkuhr
, Polytaenium
Desv
., Radiovittaria
(Benedict) E.H. Crane,
Rheopteris
Alston,
Scoliosorus
T. Moore,
Vittaria
Sm.) in
subfamily
Vittarioideae
(C. Presl) Crabbe, Jermy & Mickel of large family Pteridaceae. E. Schuettpelz et al. (2016) on the rezults
of large molecular-phylogenetic study based on plastid markers (atpA, chlN, rbcL, rpoA) also support the recognition of eleven
vittarioid genera, some of them are another in compare with M. Christenhusz’s classification:
Ananthacorus, Antrophyopsis
(Benedict) Schuettp.,
Antrophyum, Haplopteris, Hecistopteris, Polytaenium, Radiovittaria, Rheopteris, Scoliosorus, Vaginularia
Fée
,
and
Vittaria
. According to molecular-phylogenetic data,
Haplopteris
is close related with
Radiovittaria
and
Hecistopteris
(Schuettpelz et al., 2016). Genera
Hecistopteris
and
Radiovittaria
occur only in tropical America; the genus
Haplopteris
includes
about 40 species, most of them occur in tropical Asia and the Pacific Ocean islands, about ten species grows in tropical and
subtropical Africa and the Indian Ocean islands (Schuettpelz et al., 2016; GBIF). Flora of China includes 13 species (Zhang et al.,
2013).
Our previous studies of spores of some Pteridaceae groups are shown that spore morphology studied using SEM-method and
molecular-phylogenetic results are congruent for different genera such as
Onichium
(Vaganov et al., 2017a),
Pityrogramma
(Vaganov et al., 2017b),
Vaginularia
(Vaganov et al., 2017c). Morphology of spores is very important for systematics of
subfamilies Pteridoideae (Kuznetsov, 2014), Cryptogrammoideae (Vaganov, 2016) and Ceratopteridoideae (Vaganov et al.,
2017c).
Spores of several species of
Haplopteris
were investigated by Chen Ch.-W. et al. (2013, 2014, 2017). SEM-micrographs of some
Haplopteris
species are provided in the large monograph by A. Tryon and B. Lugardon (1991) under the genera
Antrophyum
(
Antrophyum
(
Haplopteris
)
ensiforme
Hk. in Benth. and
Vittaria
(
Vittaria
(
Haplopteris
)
zosterifolia
Willd. and
V.
(
Haplopteris
)
scolopendrina
(Bory) Thwaites). According to these studies, spores of
Haplopteris
are monolete and ellipsoidal or bean-shaped.
The perispore are either smooth or slightly papillate and scattered with spherules and rodlets, high variation in spore size is
observed in some
Haplopteris
species (Tryon, Lugardon, 1991; Chen et al., 2017).
The aim of this study is to provide details of spore morphology of
Haplopteris
species using scanning electronic microscopy
(SEM) to reveal features useful for systematics and phylogenetics.
Materials and methods
Spores were obtained from herbarium specimens of four species of
Haplopteris
:
H. amboinensis
(Fée) X.C. Zhang (Hunan
Province)
, H. forrestiana
(Ching) E.H. Crane (Yunnan Province),
H. linearifolia
(Ching) X.C. Zhang и
H. mediosora
(Hayata) X.C.
Zhang (Tibet Autonomous Region), stored in PE (Herbarium of Institute of Botany, Chinese Academy of Sciences (Beijing). Only
291
Spore morphology of Haplopteris C. Presl species
Ukrainian Journal of Ecology, 7(4), 2017
mature spores were used for SEM observations. Spores were mounted on SEM stubs using double-sided carbon adhesive tape
and coated with gold in a “Quorum Q150R S” sputter-coater. Stubs were viewed and photographed with the scanning
electronion microscope “Mini-SEM SNE-4500M” in the Laboratory of Structural and Molecular Analysis of Plants (Tomsk State
University, Tomsk, Russia). Spore surface was scanned in a high vacuum at voltage of 20 kV, through 2000× to 15000×
magnification. The length of major and minor equatorial diameter and polar axis, length and width of laesura were used as the
main morphometrical characters. All measurements were made on SEM-micrographs of spores in distal (major and minor
equatorial diameter), proximal (length and width of laesura) and equatorial positions (polar axis) using the computer program
“Image J”. For terminology, we primarily followed A. Tryon and B. Lugardon (1991).
Investigated specimens:
H. amboinensis
: “Pingjiang County, Hunan Province. Coll.: Gu Jia, Qi Shixin, Wang Lanying. No 124, 2008.01.26, No 01782709”.
H. forrestiana
: Yunnan Province. Mt. Huanglian Lvchun County Reservoir. Alt. (m): 1780. Col.: Qi Xinping. No Q091. Dt.:
2008.10.02. No 01757772”.
H. linearifolia
: “Zayu County, Tibet Autonomous Region. Alt.: 2500 m. Coll.: X.C. Zhang. 97°04´N 28°37´E. №4945. 2007.10.03.
№01792281”.
H. mediosora
: “Zayu County. Tibet Autonomous Region. On rocks. Alt.: 2800 m. Coll.: X.C. Zhang. 97°01´N 28°46´E. No 4978.
2007.10.05. No 01792121”.
Results and discussion
Haplopteris
has bilateral monolete spores. The spores of all examined taxa of
Haplopteris
are bean-shaped in lateral view,
ellipsoidal or ellipsoidal compressed laterally in the middle part in distal and proximal view, and round in polar view. Perispore
is thin, smooth, and easily breakable, exospore without sculpture.
1.
H. amboinensis
(Fig. 1, AD; table 1). Spores in distal and proximal view are ellipsoidal, slightly narrowed in the middle. Major
equatorial diameter 73.9–88.2 µм, minor equatorial diameter 33.5–36.2 µм, polar axis 40.0–41.5 µм, laesura length 51,554,7
µм, laesura width 2.6–3.1 µм. Exospore surface is finely granulate.
2.
H. forrestiana
(Ching) E.H. Crane (Fig., EH; table 1). Spores in distal and proximal view are ellipsoidal. Major equatorial
diameter 71.5–76.8 µм, minor equatorial diameter 36.9–40.9 µм, polar axis 31.438.6 µм, laesura length 52.0–59.0 µм, laesura
width 1.8–2.6 µм. Exospore surface is finely granulate.
3.
H. linearifolia
(Fig. 2, AD; table 1). Spores in distal and proximal view are ellipsoidal. Major equatorial diameter 51.7–61.3 µм,
minor equatorial diameter 29.2–32.9 µм, polar axis 21.7–32.9 µм, laesura length 40.2–43.5 µм, laesura width 1.3–1.8 µм.
Exospore surface is finely undulate.
4.
H. mediosora
(Fig. 2, EH; table 1). Spores in distal and proximal view are ellipsoidal, slightly narrowed in the middle. Major
equatorial diameter 47.7–51.3 µм, minor equatorial diameter 23.1–26.3 µм, polar axis 24.6–25.2 µм, laesura length 32.137.6
µм, laesura width 1.1–2.5 µм. Exospore surface is finely granulate.
Table 1. Spore size of
Haplopteris
species, mean (minmax)
Species
Major equatorial
diameter, μm
Minor equatorial
diameter, μm
Polar axis, µм
Laesura length, µм
H. amboinensis
79.5 (73.988.2)
34.3 (33.536.2)
40.9 (40.041.5)
53.1 (51.554.7)
H. forrestiana
74.7 (71.576.8)
38.7 (36.940.9)
34.7 (31.438.6)
55.2 (52.059.0)
H. linearifolia
58.1 (51.761.3)
30.6 (29.232.9)
27.5 (21.732.9)
41.6 (40.243.5)
H. mediosora
49.2 (47.751.3)
24.5 (23.126.3)
24.9 (24.625.2)
34.9 (32.137.6)
In contrast with most Pteridaceae species having tetrahedral trilete spores with very distinctive sculpture, that were described
in multiple papers (Sladkov, 1961; Nayar, Devi, 1967; Tryon, Lugardon, B., 1991; Demske et al., 2013; Kuznetzov et al., 2014;
Vaganov, 2016; Vaganov et al., 2017a, b, c, d),
Haplopteris
species has bilateral monolete spores. Spores of four examined
species are very similar in shape to each other. Spores of
Haplopteris
forrestiana
and
H. linearifolia
are ellipsoidal in distal and
proximal position, spores
H. amboinensis
and
H. mediosora
are ellipsoidal, slightly narrowed in the middle. It should be noted,
that monolete spores of Pteridaceae species, particularly
Haplopteris
, are different with monolete spores of species from other
families of leptosporangiate ferns by length-to-width ratio (major-to-minor equatorial diameter) of the spore. In
Haplopteris
length of spore is 22.5 times larger, than its width, in
Cystopteris
Bernh. and
Gymnocarpium
Newman (Cystopteridaceae)
length-to-width ratio is 1.51.8 (according to data, given by Gureyeva & Kuznetsov, 2015), i.e. spores of
Haplopteris
species are
longer and subtler.
Spores of examined species demonstrate differences in size: spores in
H. amboinensis
and
H.
forrestiana
are 1.31.6 times
larger than those in
H. linearifolia
and
H. mediosora
. Differentiation in spore size may reflect the different ploidy levels of these
species:
H. amboinensis
and
H.
forrestiana
may have a higher ploidy level than others. The largest spores are characteristic for
H. amboinensis
, the smallest spores are peculiar to
H. mediosora
. Similar results were obtained by Chen et al. (2017) for the
same species of
Haplopteris
(
H.
forrestiana, H. linearifolia, H. mediosora
, as seen on the figure 5 (Chen et al., 2017, p. 552). Two
of studied species,
H. forrestiana
and
H. linearifolia
, belongs to
H. taeniophylla
complex, which comprises of closely related
species exhibiting a little genetic differentiation (Chen et al., 2017). Spores of these species are similar in shape but different in
size.
Ukrainian Journal of Ecology, 7(4), 2017
Ukrainian Journal of Ecology
292
Acknowledgements
We are grateful to curators of the Herbarium of Institute of Botany, Chinese Academy of Sciences (Beijing) (PE) for the possibility
of obtaining of spores from the herbarium material. The study was partly supported by RFBR (project No 16-04-00513 А) and
Tomsk State University competitiveness improvement program.
Fig. 1. SEM-micrographs of spores of
Haplopteris
amboinensis
(Fée) X.C. Zhang (A–D) and
H. forrestiana
(Ching) E.H. Crane (E
H)
A, E distal side of spore; B, F proximal side of spore, laesura in the centre; C, G equatorial-proximal view of spore; D, H
lateral view of spore. All spores have fragmented perispore. Spherical spores of fungy on the surface of spores (E, G, H). Scale
bars: AD, F 30 µм; E, G, H 20 µм.
293
Spore morphology of Haplopteris C. Presl species
Ukrainian Journal of Ecology, 7(4), 2017
Fig. 2. SEM-micrographs of spores of
Haplopteris linearifolia
(Ching) X.C. Zhang. (AD) and
H. mediosora
(Hayata) X.C. Zhang (E-
H)
A, E distal side of spore; B equatorial-distal view of spore; C equatorial-proximal viev of spore; D, H lateral view of spore,
laesura from above; E distal side of spore; F proximal side of spore, laesura in the centre; G, H spore in equatorial position.
AD fragmentes of perispore are visible. Spherical spores of fungy on the surface of spores (A, C). Scale bars: AH 20 µм.
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Citation:
Vaganov, A.V., Gureyeva, I.I., Kuznetsov, A.A., Romanets, R.S. (2017 ). Spore morphology of Haplopteris C. Presl species (Vittarioideae,
Pteridaceae) from China.
Ukrainian Journal of Ecology, 7
(4), 290294.
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Comparative investigation of the spore surface ultrastructure of six specimens of Pityrogramma calomelanos (L.) Link (Pteridoideae, Pteridaceae) was carried out with using of scanning electronic microscopy (SEM). Specimens of Pityrogramma calomelanos were collected in the different parts of its range: in South China (Hainan Island), Nepal, Honduras, Ecuador, and Jamaica. It is established that for the spores of specimens collected in different parts of the range, the stability of the sculpture features and the low variability in size are characteristic. Spores are tetrahedral, trilete; laesura arms are thin and straight, equatorial diameter 48.1 (37.7-57.3) μm; distal side of spore is hemispherical, proximal side is convex or broadly conical. The stable features of the spore sculpture are the following: prominent equatorial flange protruding at 4.1 (2.5-7.8) μm, four uninterrupted ridges arranged parallel to equatorial flange and to each other on both proximal and distal sides of the spore (per two on the periphery of each side), thin and straight laesura arms, and reticular sculpture from irregularly arranged folds (rugulae) with the lumina irregular in shape on the both proximal and distal sides of the spore.
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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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The ultrastructure of the spore surface of Onychium ipii Ching (Pteridoideae, Pteridaceae) was investigated by using of scanning electron microscopy. Spores of Onychium ipii are trilete, tetrahedral, with hemispherical distal side and convex proximal one. Equatorial diameter 42.4-48.1 μm, polar axis 35.7-40.6 μm. Distal and proximal sides are separated by an equatorial flange, which protrudes on 4.3-4.8 μm all around the spore. Tubercles of different size form the interrupt "laesura lips" situated on both sides of laesura arms. Proximal side of spore with three stright ridges 3.1-4.5 μm width arranged parallel to spore margins and formed triangle in outline. Fused tubercles on distal side form sinuous folds with a few small areolae. Spores of Onychium ipii in compare with spores of O. moupinense Ching (Figure 3, D-F; Tables 1-2) are larger; their equatorial flange is more prominent; laesura arms are narrower; "laesura lips" are interrupted; stright ridges on the proximal side of spores Onychium ipii are broader, than the same in O. moupinense spores; sinuate folds on the distal side of Onychium ipii spores are more frequent than the same in O. moupinense spores. Spores of Onychium ipii and O. moupinense have the differences in the character of spore ornamentation and form. These features are the addition argument for recognition of Onychium ipii and O. moupinense as separate species.
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We used the method of scanning electron microscopy (SEM) to study the representatives of the genus Anogramma from South-West Asia and the Himalayas. To confirm the presence of Cerosora microphylla (Hook.) R.M. Tryon in China (hence, the absence of Anogramma microphylla (Hook.) Diels), we revealed the features of external morphology of spores that are not inherent in representatives of the genus Anogramma: absence of an equatorial ridge, roundish-triangular shape of the spores in the proximal-polar and distal-polar positions; absence of roller-like bulges extending upward in the form of ridges along the contour of the distal side of the spore; roller-like folds extending upward in the form of ridges both on the proximal and distal sides of the spore. In addition, one feature revealed for C. microphylla – spherical tubercles between the proximal and distal roller-like folds located along the spore equator – indicates its high species-specificity.
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Using the method of scanning electronic microscopy (SEM), a comparative study of ten representatives of subfamily Pteridoideae C.Chr. ex Crabbe, Jermy & Mickel family Pteridaceae E.D.M.Kirchn. was carried out. A comparative study of morphological characters of investigated spores has revealed characters that allow considering the relatedness of the studied species to one subfamily - Pteridoideae. Those characters include spore form - triangular-roundish, laesura rays are straight, merged into sporoderm (laesura lips), tubercles on the spore surface, and in some cases "cerebriform" folds, exosporium surface without excrescences. For species of the genera Afropteris and Taenitis , we described characters that allow considering them apart from the complex of characters belonging to the species of genera Actiniopteris, Anopteris, Onychium, Jamesonia and Pteris.
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Spores of Cystopteridaceae from northern Asia were examined using scanning electron microscopy. To evaluate the utility of spore morphology in the taxonomy of each genus, we examined spores of 14 species: seven species each of Gymnocarpium and Cystopteris. Among these are 12 species occurring in northern Asia and two species from other regions for comparative studies. The study focused particularly on perispore characters and spore size. Spores of all species examined are monolete, bean-shaped, with a range in spore size of 26–56 × 18–37 μm for Cystopteris and 25–48 × 16–34 μm for Gymnocarpium. The perispore is morphologically diverse within Cystopteris, but less so within Gymnocarpium. The perispore of the Cystopteris spores is characterised by folds and spines that are separate or form complex sculptural elements. Sacci, ridges and flanges, sometimes on the same spore, are characteristic of the perispore of Gymnocarpium. Spores have straight laesura over which the perispore forms a crest. The crest represents a high and flat fold, which is entire, foveolate or reticulate.