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We redefine the fern genus Anisocampium based on the results of a phylogenetic analysis using two regions of the chloroplast genome. The redefined genus is separate from Athyrium s. str. and comprises four species, two of which were previously assigned to Athyrium or Kuniwatsukia. Anisocampium paucijugum is reduced to a synonym of A. cumingianum. The four accepted species in Anisocampium are morphologically distinct but the putative hybrid An. ×saitoanum suggests low reproductive isolation in the genus. Anisocampium can be distinguished from related genera by a combination of characters only.
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60 (3) • June 2011: 824–830Liu & al. •
Phylogeny and taxonomy of Anisocampium
The classification of the athyrioid ferns remains unset-
tled, even though molecular phylogenetic data have been in-
creasingly used to determine relationships. One ambiguous
genus is Anisocampium C. Presl. (1849), described based on
the imparipinnate leaves and round sori of An. cumingianum
C. Presl. Copeland (1947) added a second species, An. pauci-
jugum (Aderw.) Copel., and stated that Anisocampium shows
an athyrioid affinity.
In 1973, Pichi Sermolli proposed the genus Kuniwatsukia
as a replacement name for Microchlaena Ching (1938), a later
homonym of Microchlaena Wight & Arn. (Sterculiaceae).
Micro chlaena Ching was based on M. yunnanensis (C hrist)
Ch ing (= Microchlaena cuspidata (Bedd.) Ching). Iwatsuki
(1970) considered M. cuspidata to be related to An. cumingia-
num and Athyrium sheareri (Baker) Ching, based on vascular
anatomy of rhizome and stipe, leaf architecture, and chromo-
some number. Hsieh (1985) formally transferred Athyrium
sheareri to Anisocampium.
Recently, Fraser-Jenkins (2006) referred Anisocampium
cumingianum to Athyrium sect . Niponica Ching & Y.T. Hsie h ,
a section established for Athyrium niponicum ( M ett.) H a nce
(Hsieh, 1986). Athyrium niponicum has been recognized as a
member of Athyrium for mo re th an a centu r y afte r its tr ansfer
from Asplenium, but recent molecular analyses have shown
it to have a closer affinity with Anisocampium and Kuniwat-
sukia. In his floristic treatment of Chinese Athyrium, Wan g
& al. (1999) assigned ten species to sect. Niponica, in four
series: ser. Niponica with two species (Ath. niponicum, Ath.
brevisorum (Wall. ex Hook.) Moore) and ser. Biserrulata,
ser. Fallaciosa and ser. Yokoscentia with the remai ning eight
species. Series Niponica was characterized by the creeping
rhizomes, the abruptly narrowed apex of the bipinnate leaf,
the shortened basal pinnules of the stalked, basal pinnae.
Except for Athyrium niponicum, all species differ from the
remaining 160–220 species of Athyrium in leaf dissection
and shape of sori.
A putative hybrid between Athyrium niponicum and Ath.
sheareri (≡ An. sheareri) was reported by Serizawa (1986).
Hybridization suggests close affinities of the two species. The
free venation of Athyrium sheareri is similar to that of Athyrium
and differs from the anastomosing venation of Anisocampium
Recent molecular phylogenic studies have shown that Athy-
rium s.l. (including Anisocampium) is paraphyletic. Based on
rbcL seq uence s, Sano & al. (2000) foun d Ath. sheareri and Ath.
niponicum to form a clade sister to a clade comprising Cor-
nopteris Nakai and Athyrium that includes Pseudocystoperis
Ching. Based on the trnL-F regions, Wang & al. (2003) found
Kuniwatsukia cuspidata to be cluster ed with Ath. niponicum. In
a molecular analysis using a larger dataset, Adjie & al. (2008)
confirmed that Ath. niponicum, Ath. sheareri and Ath. cuspida-
tum form a monophyletic clade. Anisocampium cumingianum,
the type species of Anisocampium, was lacking in all previous
molecular phylogenetic analyses, however, leaving the relation-
ship between Anisocampium and Athyrium unclear.
In this paper, we describe the molecular phylogeny of the
species considered to be related to Anisocampium. Based on
the results of the analysis and comparative morphology, we
redefine the genus Anisocampium as comprising four species,
the Sino-Japanese An. niponicum and An. sheareri, the Sino-
Himalayan An. cuspidatum, and the tropical and subtropical
Asian An. cumingianum (Fig. 1).
Taxon sampling and source of sequences. —
A tot al of 20
athyrioid species of ferns, including eight species of Athyrium,
three of Cornopteris, two of Diplazium, three of Deparia, and
all four species of Anisocampium were sam pled . The species of
Molecular phylogeny and taxonomy of the fern genus Anisocampium
Yea-Chen Liu,1,2 Wen-Liang Chiou1 & Masahiro Kato3
1 Division of Botanical Garden, Taiwan Forestry Research Institute, 53 Nan-Hai Road, Taipei, Taiwan
2 Current address: Depart ment of Biological Resources, National Chiayi Universit y, Chiayi, Taiwan
3 Department of Botany, National Museum of Nature and Science, Tsukuba 305-0005, Japan
Authors for correspondence: Wen-Liang Chiou,; Masahiro Kato,
We redefine the fern genus Anisocampium based on the results of a phylogenetic analysis using two regions of the
chloroplast genome. The redef ined genus is separate f rom Athyrium s.str. and comprises four species, two of which were
previously assigned to Athyrium or Kuniwatsukia. Anisocampium paucijugum is reduced to a synonym of A. cumingianum.
The four accepted species in Anisocampium are morphologically distinct but the putative hybrid An. ×saitoanum suggests low
reproductive isolation in the genus. Anisocampium can be disti ngu is he d fro m re la te d ge ner a by a combi na tion of ch ara ct ers onl y.
Anisocampium; Athyrium; Kuniwatsukia; rbcL; trnL-F
Liu & al. •
Phylogeny and taxonomy of Anisocampium
60 (3) • June 2011: 824–830
Athyrium represent all five sections recognized by Ching and
Hsieh (H si eh , 1986), or six of the 14 sec tions pro posed by Wang
(1997). Three species of sect. Niponica, i.e., Ath. niponicum,
Ath. biserrulatum, and Ath. yokoscense, are the type species
of ser. Niponica, ser. Biserrulata, and ser. Yokoscentia, re spe c-
tively. The species of Diplazium and Deparia were selected as
outgroups, because they are the closest relatives of Athyrium
and Anisocampium (Sa no & al., 200 0; Wang & al., 20 03; Adjie
& al., 2008). In addition to the rbcL and trnL-F sequence s avail-
able in GenBank (Appendix), we obtained new sequences of
Anisocampium cumingianum (rbcL, HM156338) and Athyrium
biserrulatum (rbcL, HM156336; trnL-F, HM156337). Voucher
specimens are deposited in the Herbarium of Taiwan Forestry
Research Institute (TAIF).
DNA extraction, PCR amplification and sequencing. —
Total genomic DNA of Anisocampium cumingianum and Athy-
rium biserrulatum was extracted from silica-gel-dried leaves
using the modified cetyl trimethyl ammonium bromide (CTAB)
procedure of Doyle & Doyle (1987). Two plastid genome regions
(rbcL, trnL-trnF) were amplified separately using the stan-
dard polymerase chain reaction (PCR) and published primer
sets (primer ESRBCL1F, ESRBCL1361R from Schuettpelz
& Pryer, 2007 for the rbcL gene; primer f from Taberlet &
al., 1991; Fern-1 from Trewick & al., 2002 for the trnL-trnF
region including the trnL intron and the trnL-trnF IGS). The
resulting PCR products were purified using GFX PCR DNA
and Gel Band Purification kits according to the manufacturer’s
instructions, then directly sequenced. Sequencing reactions
were conducted using the DYEnamic ET Dye Terminator Cycle
Sequencing Kit. Sequences were analyzed using MegaBACE
100 0 DNA Analysis Syst ems (Amersham Biosciences, Taipei,
Tai wan) following the manufacturer’s protocol.
Sequence alignment and phylogenetic analyses. —
ments of all sequences were generated using CLUSTAL X
(Thompson & al., 1997) and subsequently edited manually in
Bio Edit (Hall, 1999) for both intron and IGS of the trnL-F region.
Ambiguously aligned regions were excluded from all subse-
quent analyses. Phylogenetic analyses were conducted using
both maximum likelihood and Bayesian inference of phylogeny.
Maximum likelihood analyses were conducted using PHYML
v.2.4.3 (Guindon & Gascuel, 2003) under the optimal model of
sequence evolution. For each dataset, an Akaike information
criterion as implemented in jModeltest v.0.1.1 (Posada, 2008)
was us ed to sele ct the model of nucleotide subs tit ution , incl ud i ng
gamma shape and proportion of invariant sites. Maximum like-
lihood bootstrap support (MLBS) values were estimated from
1000 replicates in PHYML. Bayesian inference was performed
with MrBayes v.3.0b 4 (Ronqu ist & Huel senbe ck , 2003). Ma rkov
chains were run for 10,000,000 generations and sampled every
1000 generations, and the first 1000 trees were the “burn-in” of
the chain. An allcompat consensus tree was calculated to obtain
topology with average branch lengths as well as posterior prob-
abilities (PP) for all resolved nodes.
The phylogenetic relationships of Anisocampium we re de-
duced from the combined rbcL and trnL-F sequences (in total
2003 base pairs) (Fig. 2). Anisocampium was sister, with high
support (PP = 1.0, MLBS = 98%), to a clade comprising Athy-
rium and Cornopteris which showed high support (PP = 1.0,
MLBS = 99%). Monophyly of eac h of the th ree genera was also
robu st with hig h sup por t for each (PP = 1.0, MLBS = 100%). In
Anisocampium, a clade of An. sheareri and An. cumingianum
was sister to An. cuspidatum (≡ Ath. cuspidatum), and together
sister to An. niponicum (≡ Ath. niponicum), although support
for the first two clades was not very high.
60°E 90°E 120°E 150°E
Anisocampium niponicum
Anisocampium ×saitoanum
Anisocampium cumingianum
90°E 120°E 150°E
Anisocampium sheareri
Anisocampium cuspidatum
Fig. 1.
Distribution of Anisocampium.
60 (3) • June 2011: 824–830Liu & al. •
Phylogeny and taxonomy of Anisocampium
The Anisocampium clade is distinct from the Athyrium
clade, and similar to the topology of previous analyses (Sano
& al., 2000; Wang & al., 2003; Schuettpelz & Pryer, 2007;
Adjie & al., 2008). In the taxonomic definition below, we re-
define the genus Anisocampium to include An. cuspidatum
(≡ Kuniwatsukia cuspidata) and An. niponicum (≡ Athyrium
niponicum), and separate the genus from Athyrium and Cor-
nopteris. The results are not consistent with current taxo-
nomic treatments based on morphological comparisons (Kato,
1977; Ching, 1978; Kato & Kramer, 1990; Wang, 1997; Wang
& al., 1999). Kuniwatsukia and Anisocampium have been
treated as synonyms of Athyrium (Kato, 1977; Fraser-Jenkins,
2006), while Cornopteris has long been presumed to be re-
lated to Diplazium (Ching, 1945; Wang, 1997) or Athyrium
(Tagawa, 1959; Kato, 1979). Based on morphological similar-
ity, Fraser-Jenkins (2006) placed Ath. cumingianum (≡ An.
cumingianum), along with Ath. niponicum, in Athyrium sect.
Niponica, while recognizing Cornopteris. He also noted the
similarity of Ath. cumingianum with Ath. cuspidatum and Ath.
An alternative classification to ours is to recognize Athy-
rium s.l. as including Anisocampium and Cornopteris. Cor-
nopteris is characterized by exindusiate sori and fleshy pro-
jections (corni) at the base of costae and costules, by which it
Athyrium vidalii
Athyrium nakanoi
Athyrium strigillosum
Athyrium otophorum
Athyrium biserrulatum
Athyrium filix-femina
Athyrium rupestre
Athyrium yokoscense
Anisocampium sheareri
Anisocampium cumingianum
Anisocampium cuspidatum
Anisocampium niponicum
Diplazium nipponicum
Diplazium hachijoense
Deparia lobatocrenata
Deparia petersenii
Deparia conilii
Cornopteris decurrentialata
Cornopteris crenulatoserrulata
Cornopteris christenseniana
Fig. 2.
Phylogenetic tree of
Anisocampium and allied gen-
era based on rbcL and trnL-F
region sequences. Tree topology
is deduced from Bayesian infer-
ence analysis; numbers indicate
values of posterior probabilities
(above branches) and bootst rap
(below branches); numbers less
than 0.5 or 50 are not shown.
Three species of Deparia and
two of Diplazium are outgroups.
Liu & al. •
Phylogeny and taxonomy of Anisocampium
60 (3) • June 2011: 824–830
is distinguished from Athyrium. Kato (1977), Kato & Kramer
(1990) and Fraser-Jenkins (2006) treated Anisocampium and
Kuniwatsukia as synonyms of Athyrium, but separated Cornop-
teris. Our phylogenetic analysis indicates that such a circum-
scription results in the paraphyly of Athyrium.
Like Adjie & al. (2008), we did not find any unique diag-
nostic characters to define Anisocampium. The genus can only be
defined and distinguished from the closely related Athyrium and
Cornopteris by a combination of features of the stipe base, projec-
tions, spines and sori (Table 1). The base of the stipe is as broad
as the upper part of the stipe in Anisocampium, while the stipe is
broader near the base with lateral pneumatophores and narrowed
to the very base in Athyrium, Cornopteris and Deparia within
the athyrioid ferns (Liew, 1972; Kato, 1977, 1979, 1984). The
stipe base is persistent after the leaves wither. They are arranged
tightly around the short, erect rhizomes in most species of these
two genera, or are loosely arranged on the creeping rhizomes in,
e.g., Athyrium atkinsonii Bedd. and Cornopteris decurrentialata
Nakai. The persistent stipe base, called a trophopod (Johnson,
1986), is possibly a derived, adaptive trait in plants in the temper-
ate zone. The persistent stipe base is absent in Anisocampium and
Diplazium, even in species with erect rhizomes.
Fleshy projections (corni) in the groove on the adaxial sur-
face at the base of the costae and costules are characteristic of
Cornopteris, while they are absent from Anisocampium and
Athyrium. There are spines in many species of Athyrium and
Cornopteris, but none in Anisocampium. The spines are continu-
ations of the interrupted ends of ridges along the groove of the
costae and costules on the adaxial surface of the leaf. The sori
are rounded-reniform, shortly U- or J-shaped and indusiate in
most species of Anisocampium, but they are also oblong-linear
in An. niponicum. In Cornopteris, however, they are rounded
to linear and exindusiate and in Athyrium usually linear or
J-shaped and indusiate.
Diagnostic characters separating Anisocampium and related
genera are shown in Table 1. A key for their identification and
a brief taxonomic treatment of the revised genus Anisocampium
are given below.
1. Groove of costa interrupted at base and not confluent with
groove of rachis; multicellular hairs present on stipe, rachis
and costae, rarely few or absent ................... Deparia
1. Groove of costa continuous with groove of rachis; multicel-
lular hairs absent or sometimes present ................... 2
2. Sori exindusiate; fleshy projections present at base of costa
2. Sori indusiate; fleshy projections absent .................. 3
3. Sori linear or oblong, not J- or U-shaped; groove of ra-
chis and costa U-shaped with flat base in cross section
..................................................... Diplazium
3. Sori various in shape, linear, oblong, J- or U-shaped, or
round-reniform; groove of rachis and costa V-shaped in
cross section ................................................. 4
Rhizome ascending or shortly creeping; stipe base usually
broadened, with pneumatophores on sides, narrowed down-
ward; spines at base of costules and midvein present or
absent adaxially; lamina 2- or 3-pinnate, sometimes pinnate
or bipinnatifid, usually gradually narrowed to apex
...................................................... Athyrium
4. Rhizome creeping or ascending; stipe base not broadened,
devoid of pneumatophores; spines absent; lamina pinnate
or bipinnatifid, in A. niponicum bipinnate to tripinnatifid,
abruptly narrowed to apex or terminated by pinna conform
to lateral pinnae .............................Anisocampium
Anisocampium C. Presl, Epimel. Bot.: 52. 1849; Tardieu &
Christensen in Lecomte, Fl. Gén. l’Indo-Chine 7(7): 281.
1940; Copeland, Gen. Fil.: 151. 1947; Ching in Acta Phy-
totax. Sin. 16(3): 13. 1978 Type: Anisocampium cum-
ingianum C. Presl.
= Kuniwatsukia Pic. Serm. in Webbia 28: 455. 1973; Ching in
Acta Phytotax. Sin. 16(3): 13. 1978 Microchlaena Ching
in Bull. Fan Mem. Inst. Biol. 8: 322. 1938, non Micro-
chlaena Wall. ex Wight & Arn. 1834; Iwatsuki in Acta
Phytotax. Geobot. 24: 188. 1970 Type: Kuniwatsukia
cuspidata (Bedd.) Pic. Serm., syn. nov.
= Athyrium Roth sect. Niponica Ching & Y.T. Hsieh in Bull.
Bot. Res., Harbin 6(4): 131. 1986, pro parte – Type: Athy-
rium niponicum (Mett.) Hance, syn. nov.
Rhizome creeping or ascending to shortly erect (in A. cus-
pidatum), radially symmetrical or dorsiventral (in A. cumingia-
num), apex scaly. Leaves more or less distant from each other,
stipe base scaly, not swollen, with 1 (in A. niponicum) or 2 leaf
traces in rhizome, deltoid-ovate or ovate-oblong, imparipinnate
to 3-pinnatifid, abruptly reduced to pinnatifid apex or bearing
terminal pinna conforming with lateral pinnae; pinnae lanceo-
late or narrowly elliptic, base of lower pinnae inequilateral;
veins free or anastomosing (in A. cumingianum); spines absent
on adaxial surface at base of costae, costules and midvein. Sori
oblong, J-shaped, horseshoe-shaped, round-reniform, indusiate;
indusia conforming with sori, margin ciliate, or erose, often fu-
gacious. Spores bilateral, with perispore. Chromosome number
x = 40. Four species.
Habitat. – Mesic forests from near sea level to 1900 m alt.
Distribution. – Japan, Korea, China, Taiwan, Vietnam,
Laos, Thailand, Myanmar, India, Bhutan, Nepal, Sri Lanka,
Philippines, Indonesia (Fig. 1).
KEy TO THE SpECIES Of AnisocAmpium
1. Lamina pinnate, apex of lamina pinnatifid, not conforming
with lateral pinnae .......................................... 2
1. Lamina terminated by a pinna conforming or subconform-
ing to lateral pinnae ......................................... 4
2. Lamina pinnate throughout; pinnae usually shallowly
lobed; sori round ..............................1. A. sheareri
2. Lamina bipinnatifid or bipinnate; pinnae lobed more than
2/3 to costa; sori oblong, J-shaped, or round ............. 3
3. Lamina bipinnate; pinnules free, lobed ...2. A. niponicum
60 (3) • June 2011: 824–830Liu & al. •
Phylogeny and taxonomy of Anisocampium
3. Lamina bipinnatifid; pinnules not free, at least adnate to
costa by broad base .....................2a. A. ×saitoanum
4. Pinnae to 7 pairs, lanceolate, base cuneate or round; veins
anastomosing ........................... 3. A. cumingianum
4. Pinnae 10 pairs or more, linear, base truncate; veins free
..............................................4. A. cuspidatum
1. Anisocampium sheareri (Baker) Ching in Acta Bot. Yunnan.
7(3): 314. 1985; Wang & al., Fl. Reipubl. Pop. Sin. 3(2):
75, pl. 15, f. 1–6. 1999 Nephrodium sheareri Baker in
J. Bot. 13: 200. 1875 ≡ Dryopteris sheareri (Baker) C. Chr.,
Index Filic.: 292. 1905 Athyrium sheareri (Baker) Ching
in Christensen, Index Filic., Suppl. 3: 44. 1934; Iwatsuki,
Ferns Fern Allies Jap.: 232, photo 151(1, 2). 1992; Kato in
Iwatsuki & al., Fl. Jap. 1: 203. 1995 – Type: China, Shearer
s.n. (K, photo!).
= Nephrodium polypodiforme Makino in Bot. Mag. (Tokyo) 13:
58. 1899 Dryopteris polypodiformis (Makino) C. Chr.,
Index Filic.: 285. 1905 – Syntypes: Japan, Z. Matsumura
s.n., Jul. 17, 1882 (TI), Y. Yoshinaga s.n., 1887, M. Makino
s.n., Aug. 1887, 1889, Nov. 1892 (K, photo!).
= Aspidium otarioides Christ in Bull. Acad. Int. Géogr. Bot. 11:
247. 1902 Dryopteris otarioides (Christ) C. Chr., Index
Filic.: 282. 1905 – Type: China, E. Bodinier 2050 (P, photo!).
= Nephrodium isolatum Baker in Gard. Chron., n.s., 14: 494.
1880 – Type: China, Maries s.n. (K, photo!).
Distribution. – Japan, South Korea (Cheju Is.), China
(North Central, Southeast, South Central), Taiwan (new record,
Hualien Co., Y.Y. Huang 620; deposited in HAST).
2. Anisocampium niponicum (Mett.) Y.C. Liu, W.L. Chiou
& M. Kato, comb. nov.Asplenium niponicum Mett. in
Ann. Mus. Bot. Lugduno-Batavi 2: 240. 1866 Athy-
rium niponicum (Mett.) Hance in J. Linn. Soc., Bot. 13:
92. 1872; Tardieu & Christensen in Lecomte, Fl. Gén.
l’Indo-Chine 7(7): 273. 1940; Dickason in Ohio J. Sci. 46:
123. 1946; Iwatsuki, Ferns Fern Allies Jap.: 239, photo
160(1, 2). 1992; Shieh & al. in Editorial Committee of
the Flora of Taiwan, Fl. Taiwan, ed. 2, 1: 424. 1994; Kato
in Iwatsuki & al., Fl. Jap. 1: 210. 1995; Wang & al., Fl.
Reipubl. Pop. Sin. 3(2): 132, pl. 25, f. 1–8. 1999; Liu &
al., Fern Fl. Taiwan, Athyrium: 34, fig. 14, map 14, photo
14. 2009 – Type: Japan, Keiske (K. Ito), Siebold, Buerger
s.n. (L, photo!).
= Asplenium uropteron Miq. in Ann. Mus. Bot. Lugduno-Batavi
3: 174. 1867 Athyrium uropteron (Miq.) C. Chr., Index
Filic.: 147. 1905 Type: Japan, Keiske (K. Ito) s.n. (L,
Table 1.
Comparison of Anisocampium and related genera.
Character Anisocampium Cornopteris AthyriumaDiplaziumbDepariac
Rhizome Creeping or erect Creeping or erect Erect or ascending,
few creeping
Creeping, ascending
or erect
Creeping, ascending
or erect
Stipe base Not persistent Persistent Persistent, rarely not Not persistent Persistent or not
Scale margin Entire Entire Entire Toothed, rarely entire Entire
Multicellular hairs Absent Present or absent Absent, rarely present Absent, rarely present Present, rarely absent
Unicellular hairs Present Present or absent Present or absent Absent Absent
Pinna- or pinnule base Inequilateral Equilateral Inequilateral Equilateral Equilateral
SpinedAbsent Present Present or absent Absent Absent
Continuation of grooveeConfluent Confluent Confluent Confluent Not confluent
Shape of groovefV-shaped V-shaped V-shaped Widely U-shaped,
rarely V-shaped
Shallowly V-shaped
ProjectionsgAbsent Present Absent Absent Absent
Veins Free or anastomosing Free Free Free or anastomosing Free or anastomosing
Sori Round-reniform, U-
or J-shaped, or oblong
Round or oblong Round-reniform,
U- or J-shaped, linear,
or oblong
Linear or oblong Round-reniform,
U- or J-shaped, linear,
or oblong
Indusium Present Absent Present, rarely absent Present Present
Indusium margin Ciliate, lacerate, erose
or subentire
Entire, erose, rarely
Entire or erose Entire, erose, lacerate
or ciliate
Chromosome number 40 40 or 41 40 41 40 or 41
a Including Pseudocystopteris.
b Including Allantodia.
c Including Athyriopsis, Dictyodroma, Dryoathyrium, Lunathyrium, and Triblemma.
d Spines are continuations from ends of interrupted ridges embanking grooves at insertions of costae, costules and midveins on adaxial surface of lamina.
e Grooves on adaxial surface of costae and costules are confluent with grooves on costules and midveins or not confluent because of a separating ridge.
f Grooves are V-shaped or U-shaped with flat bottom in cross section.
g Fleshy projections (corni) in grooves at base of pinnae and/or pinnules differ from spines, which are continuous with ridges.
Liu & al. •
Phylogeny and taxonomy of Anisocampium
60 (3) • June 2011: 824–830
= Athyrium biondii Christ in Nuovo Giorn. Bot. Ital., n.s., 4: 91.
1897 – Type: China, G. Giraldi s.n. (B, photo!).
= Athyrium yunnanense Christ in Bull. Geogr. Bot. Mans. 17:
134. 1907 – Type: China, Cavalerie 3864 (K, photo!).
= Athyrium fissum Christ in Notul. Syst. (Paris) 1: 47. 1909 –
Type: China, Henr y 1310 7 (P, photo!).
Distribution. – Japan, Korea, China (Northeast, North
Central, Southeast, South Central), Taiwan, southern Vietnam,
Myanmar (central), northeast India (Arunachal Pradesh). An-
isocampium niponicum is also in north east India (Man ipur and
Tripura) (C.R. Fraser-Jenkins, pers. comm.).
Note. – Ogura (1921) and Kato (1977) found single-leaf
trace vasculature in An. niponicum, Ath. yokoscense (Franch.
& Sav.) Christ, and Ath. vidalii (Franch. & Sav.) Nakai, which
is contrasted with the bina r y leaf trace in most athy rioid ferns.
The leaf trace is divided into two and united to a single U-
shaped bundle in cross section in the upper part of the stipe.
2a. Anisocampium ×saitoanum (Sugim.) M. Kato, comb. nov.
Athyrium sheareri f. saitoanum Sugim., Keys Herb. Pl.
Jap. 3: 404. 1966 ≡ Athyrium ×saitoanum (Sugim.) Seriz.
in J. Phytogeogr. Taxon. 34: 68, f. 1–3. 1986 – Type: Japan,
Y. Saito s.n. (Herb. Sugimoto, photo!).
Distribution. – Western Japan; the distribution of Aniso-
campium ×saitoanum ext end s over most of the range of overlap
of A. niponicum and A. sheareri.
Note. – Anisocampium ×saitoanum is a putat ive hybr id de-
rived from crosses between An. niponicum and An. sheareri. It
has been recorded from 18 prefectures in western Japan (Fig. 1;
Serizawa 1986; Kurata & Nakaike, 1997). The parental spe-
cies are phylogenetically the farthest from each other in the
genus in Japan. Anisocampium ×saitoanum can reproduce to
a certain degree in culture, indicating incomplete reproductive
isolation between the species (T. Oka, pers. comm.). The hybrid
may therefore be reproducing in nature or may be the result
of multiple origins. Either case suggests a close affinity of the
two parental species.
3. Anisocampium cumingianum C. Presl, Epimel. Bot.: 59.
1849; Tardieu & Christensen in Lecomte, Fl. Gén. l’Indo-
Chine 7(7): 281. 1940; Copeland, Fern Fl. Philipp. 3: 418.
1960; Sledge in Bull. Brit. Mus. (Nat. Hist.) Bot. 2(11):
281. 1962; Tagawa & Iwatsuki, Fl. Thailand 3(3): 444,
f. 47(5–7). 1988; Manickam & Irudayaraj, Pterid. Fl. W.
Ghats: 231, pl. 178. 1991; Wang & al., Fl. Reipubl. Pop.
Sin. 3(2): 77, pl. 15, f. 7–8. 1999 ≡ Athyrium cumingia-
num (C. Presl) Ching in Christensen, Index Filic., Suppl.
3: 40. 1934; Fraser-Jenkins in Shaffer-Fehre, Rev. Handb.
Fl. Ceylon 15(B): 548. 2006 ≡ Aspidium otaria Kunze ex
Mett., Abh. Sencken berg. Naturf. Ges. 2: 318. 1858Neph-
rodium otaria (Kun ze ex Met t.) Baker in Hoo ke r & Baker,
Syn. Fil.: 288. 1867 – Type: Philippines, Cuming 239 (K!).
= Phegopteris paucijuga Al der w. in Bu ll. Jard. Bo t. Buit enzorg ,
se r. 2, 16: 26. 1914 Anisocampium paucijugum (Alderw.)
Copel., Gen. Fil.: 151. 1947 – Type: Java, C.A. Backer 8290
(BO, photo!), syn. nov.
Distribution. – China (southern Yunnan), Taiwan (Liu &
al., 2000), Laos, Thailand, southern Myanmar, southern and
northeast India, Philippines (Luzon), Indonesia (Bali, East
Java, newly recorded to South Sulawesi [Bantimurung], T. Oka
100608011, TNS), Sri Lanka. Anisocampium cumingianum also
occurs in the Lesser Sunda Islands (Sumbawa, Timor).
Note. – The lamina of An. paucijugum, described from
eastern Java, is relatively small, i.e., 12 cm long and has two or
three pinnae on each side of the rachis. The size of the leaf and
the number of pinnae are within the range of variation of An.
cumingianum. The other cha r acter ist ics, e.g., the impa r ipinnate
leaf, anastomosing veins, and round sori, are shared with An.
paucijugum and An. cumingianum.
Anisocampium cumingianum is the most de rived spe cies in
the genus in its dorsiventral rhizome (vs. radially symmetrical),
the imparipinnate lamina (vs. paripinnate lamina with gradu-
ally less dissected apical part), and the anastomosing veins
(vs. free veins).
4. Anisocampium cuspidatum (Bed d.) Y.C. Liu, W.L. Chiou &
M. Kato, comb. nov. Lastrea cuspidata Be dd., Fer ns Br it .
India: t. 118. 1870 Aspidium cuspidatum Mett. in Abh.
Senckenberg. Naturf. Ges. 2: 376. 1858, non Desv. 1827 ≡
Nephrodium cuspidatum Baker in Hooker & Baker, Syn.
Fil.: 260. 1867, non C. Presl 1825 ≡ Dryopteris cuspidata
(Bedd.) Christ in Philipp. J. Sci., C 2: 205. 1907 ≡ Micro-
chlaena cuspidata (Bedd.) Ching in Acta Phy totax. Sin. 9:
99. 1964; Iwatsuki in Acta Phytotax. Geobot. 24: 188. 1970
Kuniwatsukia cuspidata (Bedd.) Pic. Serm. in Webbia 28:
455. 1973; Tagawa & Iwatsuki, Fl. Thailand 3(3): 442, f.
47. 1988; Wang & al., Fl. Reipubl. Pop. Sin. 3(2): 78, pl. 16.
1999 Athyrium cuspidatum (Bedd .) M. Kato in Bot. Mag.
(Tokyo) 90: 27. 1977 ≡ Dryopteris khasiana C. Ch r., Index
Filic.: 272. 1905 ≡ Polypodium elongatum Wall. ex Hook.,
Sp. Fil. 4: 234. 1862, non Aiton 1789 ≡ Phegopteris elongata
J. Sm., Hist. Fil.: 233. 1875, non E. Fourn. 1872 ≡ Lastrea
elongata Bedd . ex Cla rke, Tra ns. Lin n. Soc. Lon don, Bot. 1:
513. 1880, non C. Presl 1836 ≡ Dryopteris elongata (Bedd.
ex Clarke) Kuntze, Revis. Gen. Pl. 2: 811. 1891 – Type:
Nepal, Wallich Cat. No. 309 (NY, photo!).
= Aspidium yunnanense Christ in Bull. Herb. Boissier 6: 965.
1898 ≡ Microchlaena yunnanensis (Christ) Ching in Bull.
Fan Mem. Inst. Biol. Bot. 8: 325, t. 6. f. 1. 1938; Dickason
in Ohio J. Sci. 46: 120. 1946 Dryopteris yunnanensis
(Christ) Copel., Gen. Fil.: 122. 1947 Aspidium stenolepis
Christ, Bull. Herb. Boissier 7: 22. 1899 – Type: China,
A. Henry 9038A (US, NY, photo!).
Distribution. – China (South Central, Southwest Central),
northeast Myanmar, Northern Thailand, Bhutan, northeast
India, central and east Nepal. According to C.R. Fraser-Jenkins
(pers. comm.), Anisocampium cuspidatum also occurs in the
western Himalayas (Pithoragash, India). Hooker (1862: 234)
thought that the type specimen of Polypodium elongatum, col-
lected in Kew Gardens, originated in Sri Lanka, but Clarke
(1880: 512) doubted it (Fraser-Jenkins, 2006).
Note. – Anisocampium cuspidatum is distinct in having
shortly creeping rhizomes with an ascending apex and radially
arranged leaves with linear pinnae.
60 (3) • June 2011: 824–830Liu & al. •
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We th an k C.R. Fra ser-Jen kin s for pr ovidin g di st r ib ut ion da ta for
this group of ferns, T. Oka for useful infor mation on A. cumingianum
and A. ×saitoanum, Titien Ng Pr aptos uw ir yo for pr ovidi ng a phot o of
the type specimen of A. paucijugum, X. Cheng for providing infor-
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correcting the English of the manuscript. We thank the curators of
BO, HAST, K, KYO, PE, PNH, PYU, SYSU, and TAIF for allowing
us to examine specimens in their herbar ia. T his study was suppor ted
by a Grant-in-Aid for Scientific Study from the Japan Society of the
Promotion of Science to M.K. and from the Taiwan Nat ional Science
Council ( NSC-97-2313-B-054-005-MY3) to W.L.C.
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Species, sources and GenBan k accession s (rbcL, trnL-F) used in this st udy.
Anisocampium cumingianum C. Presl, J.M. Chen s.n., Taiwan (TAIF), HM156338, FJ807659. Anisocampium cuspidatum (Bedd.) Y.C. Liu, W.L. Chiou &
M. Kato, EU329029, EU329073 (Adjie & al., 2008). Anisocampium niponicum (Mett.) Y.C. Liu, W.L. Chiou & M. Kato, EU329042, EU329085 (Adjie & al.,
200 8). Anisocampium sheareri (Bake r) Ching in Y.T. Hsieh, EU329047, EU32909 0 (Adjie & al., 20 08). Athyrium biserrulatum H. Christ, YCLiu 9370, China
(TAIF), HM156337, HM156336. Athyrium filix-femina (L.) Roth, EU329032, EU32 90 76 (Adji e & al ., 2008). Athyrium nakanoi Mak ino, EU329039, EU32 90 82
(Adjie & al., 2008). Athyrium otophorum (Miq.) Koid z., EU329 04 4, EU329087 (Adjie & al., 2008). Athyrium rupestre Kodama, EU 3290 45, EU329088 (Adjie
& al., 2008). Athyrium strigillosum (Moore ex Lowe) Moore ex Salom, EU329049, EU329092 (Adjie & al., 2008). Athyrium vidalii (Franch. & Sav.) Nakai,
EU329052, EU329095 (Adjie & al., 2008). Athyrium yokoscense (Franch. & Sav.) H. Christ, EU329055, EU329098 (Adjie & al., 2008). Cornopteris christense-
niana (Koidz.) Tagawa, EU329061, EU329104 (Adjie & al., 2008). Cornopteris crenulatoserrulata Nakai, EU329062, EU329105 (Adjie & al., 2008). Cornopteris
decurrentialata ( Ho ok.) Naka i, EU329063, EU329106 (Adji e & al ., 200 8). Deparia conilii (Fr an ch . & Sav.) M. Ka to , EU 329 065, EU 32910 8 (Ad jie & al. , 20 08) .
Deparia lobatocrenata (Tagawa) M. Kato, EU 329 06 6, EU329109 (Adjie & al., 20 08). Deparia petersenii (Ku nze) M. Kato, EU3290 64 , EU 32910 7 (Adjie & al.,
200 8). Diplazium hachijoense Nakai, EU329068, EU329111 (Adjie & al., 2008). Diplazium nipponicum Tagawa, EU329067, EU329110 (Adjie & al., 2008).
... From the evolutionary emergence of primitive organisms to today's broad variety of organisms, people have been constantly exploring how many species there are on the earth and what kind of evolutionary relationship among species. With the development of open science and technological innovation, methods of species identification range from using morphological characteristics to the integration of various methods (e.g., molecular biology, bioinformatics, bionomics) [1][2][3][4][5], which help us gain a more in-depth understanding of the evolutionary process between organisms and their accurate position in the tree of life. Due to the multi-disciplines combination and the improvement of sharing databases, many misclassifications hidden in the past have been gradually discovered, and their key morphological boundaries have also been rewritten. ...
... Ferns, an ancient group, also have similar examples, one of which is Athyrium niponicum (Mett.) Hance 1873. A. niponicum had been treated as a member of Athyrium Roth 1875, but was later confirmed to be within Anisocampium C. Presl 1851 based on rbcL and trnL-F region sequences [2]. At the end of the paper, the author revised the morphological boundaries of Athyrium and Anisocampium according to the results of systematics [2]. ...
... Hance 1873. A. niponicum had been treated as a member of Athyrium Roth 1875, but was later confirmed to be within Anisocampium C. Presl 1851 based on rbcL and trnL-F region sequences [2]. At the end of the paper, the author revised the morphological boundaries of Athyrium and Anisocampium according to the results of systematics [2]. ...
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With the development of open science and technological innovation, using sharing data and molecular biology techniques in the study of taxonomy and systematics have become a crucial component of plants, which undoubtedly helps us discover more hidden outliers or deal with difficult taxa. In this paper, we take Dennstaedtia smithii as an example, based on sharing molecular database, virtual herbarium and plant photo bank, to clarify the outliers that have been hidden in Dennstaedtia and find the key morphological traits with consistent of molecular systematics. In molecular phylogenetic analyses, we used rbcL, rps4, psbA-trnH and trnL-F sequences from 5 new and 49 shared data; the results showed that Dennstaedtia smithii is nested within Microlepia rather than Dennstaedtia. We further studied the morphological characters based on the phylogeny result and found that D. smithii is distinguished from other species of Dennstaedtia by spore ornamentation and the unconnected of grooves between rachis and pinna rachis. According to morphological and molecular phylogenetic studies, our results supported that D. smithii should be a new member of Microlepia and renamed Microlepia smithii (Hook.) Y.H. Yan. Finding hidden outliers can promote the consistency of morphological and molecular phylogenetic results, and making the systematic classification more natural.
... 1). На основании результатов морфологических и генетических анализов вид был включен в род Anisocampium C. Presl (Liu et al., 2011). Ниже приводится морфологическое описание нового для России вида. ...
... Следует учесть факт гибридизации между этими видами, о чем было сказано выше. Главные отличия A. niponicum от A. sheareri заключаются в строении вай и форме сорусов (Liu et al., 2011). ...
On the Krabbe peninsula (Khasansky District, Primorsky Krai, Russian Federation), a population of a new genus and species of fern for Russian Federation, Anisocampium niponicum (Mett.) Y. C. Liu, W. L. Chiou et M. Kato (Athyriaceae), was discovered. The description of plants and locality is given. A high vitality of the population was noted, which is associated with the absence of anthropogenic influence, as well as with the growth of A. niponicumunder conditions close to its ecological optimum.
... In the Malesian region, based on published literatures and initial examination of specimens, there are more than 20 recognized species of Athyrium s.s. (Wardani & Adjie, 2018;Hassler, 2004Hassler, -2022, one A nisocam-pium (Liu et al., 2011) and five Cornopteris (Kato, 1979(Kato, & 1986. There has no A thyrium s.s. ...
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WARDANI, W., ADJIE, B., YULITA, K. S. & SALAMAH, A. 2022. Two new records of Athyrium for Bali. Reinwardtia 21(2): 43‒47. — Here we reported two species of Athyrium from Bali for the first time based on herbarium study and field work. We provided morphological description, voucher specimens, and taxonomic note for both species. A key to all the four known species of Athyrium s.l. in Bali was also provided.
... The genus Athyrium is a medium sized terrestrial fern, mainly distributed in the northern hemisphere that in the tropics grow only in humid high altitude (Rothfles 2012, PPG I 2016, Wei et al 2018. This genus can be differentiated from other Athyriaceae genera by a combination of characters, i.e., confluent groove with projections, inequilateral pinna or pinnule base, entire scale margin, linear to horseshoe-shaped indusium (Liu et al 2011). Athyrium nigripes (Blume) T. Moore firstly published in Blume's Enumeratio Plantarum Javae (1828) as Aspidium nigripes, based on specimen collected from the wetland at a valley between Mount Burangrang and Mount Tangkuban Parahu in West Java. ...
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Athyrium nigripes is a highly variable species described and recorded mostly from Java and few from surrounding islands. There are different arguments on its delimitation and hence its extent of distribution. We attempted to elucidate disagreement on delimitation of four related species described from Java: Athyrium nigripes, Aspidium costale, At. pulcherrimum and At. nitidulum with phenetic approach. The analysis formed four clusters; cluster I consist of the large sized and include type specimen of At. pulcherrimum and Asp. costale, cluster II for plants with standard form and include type specimen of At. nigripes, cluster III only for serrated plants and cluster IV contain the At. nitidulum form. Since Asp. costale clustered in group I, this species is more closely related to At. pulcherrimum than to At. nigripes nested in group II. Large size and degree of division indicate major clustering within the complex species.
... Beside these treachery elements is used for taxonomic purpose in this family (Xu & Wang, 2009) stomata types (Yan-Ping & Jian-Meng, 2012) vascular bundles (Karafit, Rothwell, Stockey, & Nishida, 2006) and the petiole anatomy (Hernandez-Hernandez, Terrazas, Mehltreter, & Angeles, 2012) have been used for the taxonomic purpose in Athyriaceae. However, on the basis of morphological and few earlier anatomical studies the grouping of the taxa looks very artificial some of the phylogenetic studies were performed which can possibly help in the delimitation of genera and species within Athyriaceae (Chunxiang, Shugang, Xiaoyan, & Qun, 2011;Liu et al., 2011;Sano et al., 2000;Wang, Xie, & Zhao, 2004;Wei et al., 2013Wei et al., , 2018. ...
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For the robust identification of taxonomically complex fern family like Athyriaceae, light and scanning electron microscopy is significance implications. This article present first microscopic investigation of foliar micromorphology of 3 genera and 10 species belonging to Athyriaceae namely, Athyrium, Deparia, and Diplazium were collected from different localities in Malakand Division , Northern Pakistan. In present study we compare foliar micromorphology of all 10 species using standard protocols of light microcopy (LM) and scanning electron microscopy. Qualitative micromorphological variations in shape of epidermal cells, anticlinal wall pattern, stomatal type and shape, stomatal pore shape, guard cells shape, and trichomes types were studied. In addition, some quantitative characters were also studied and data were statistically analyzed in epidermal cell size, stomatal size, stomatal pore size, stomatal density, and stomatal index. The pivotal result of study include; shape of epidermal cell in all species is irregular on both abaxial and adaxial surfaces. The anticlinal walls are sinuous in most of the species but some species have irregular lobed and broadly lobed wall. Leaves are hypostomatic in all studied species. Two main categories of stomatal type were found: polocytic and anomocytic. Unicellular nonglandular trichomes were observed in only one species Athyrium mackinnoni. The variation in foliar micromorphological characters between the genera and within the species was useful in identification and classification and have potential taxonomic significance for species differentiation. An identification key using micromor-phological characters are provided to distinguish genera and species.
... In the Malesian region, based on published literatures and initial examination of specimens, there are more than 20 recognized species of Athyrium s.s. (Wardani & Adjie, 2018;Hassler, 2004Hassler, -2022, one A nisocam-pium (Liu et al., 2011) and five Cornopteris (Kato, 1979(Kato, & 1986. There has no A thyrium s.s. ...
Full-text available
WARDANI, W., ADJIE, B., YULITA, K. S. & SALAMAH, A. 2022. Two new records of Athyrium for Bali. Reinwardtia 21(2): 43‒47. — Here we reported two species of Athyrium from Bali for the first time based on herbarium study and field work. We provided morphological description, voucher specimens, and taxonomic note for both species. A key to all the four known species of Athyrium s.l. in Bali was also provided.
... A preliminary BLAST search (Johnson et al., 2008) of the sequences of the new species in GenBank showed that it was most genetically similar to A. epirachis Christ (1905: 51) Ching (1934, which belongs to the A. otophorum clade (Wei et al. 2018b), thus we included most species of that clade. A species of Cornopteris Nakai (1930: 7) was used as outgroup based on recent phylogenetic studies (Liu et al. 2011, Wei et al. 2018b. ...
A new fern species, Athyrium aberrans (Athyriaceae), currently only known from southeastern Xizang, China is described. Morphologically, A. aberrans is superficially similar to A. falcatum in the habit and pinna shape, but the new species can easily be distinguished from the latter by having purplish petioles and rachises, a bulbil on the rachis below the lamina apex, and finely toothed pinna margins. A phylogenetic analysis based on five plastid markers demonstrates that A. aberrans is most closely related to but obviously distinct from A. epirachis.
... There has been no revision of the Philippine Athyrium after the work of Copeland (1958). However, comprehensive revision of the family Athyriaceae in Taiwan has been published by Liu et al. (2009) Several infrageneric classifications of Athyrium have been proposed primarily based on gross morphology and phylogenetic studies (Liu et al. 2011, Rothfels et al. 2012, PPG1 2016, Wardani & Adjie 2018, Wei et al. 2018. ...
Full-text available
Athyrium nakanoi Makino is a new fern species record for the Philippines, discovered in Mt. Dulang-Dulang, Kitanglad Range, Mindanao, Philippines extending its original distribution from India, Nepal, Bhutan, China, Taiwan and Japan, Indonesia and Malaysia. Diagnostic description of the species is provided together with its distribution, conservation status and a dichotomous key to the Malesian Athyrium Sect. Polystichoides.
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A new record of the fern genus Cornopteris Nakai from Peninsular Malaysia is presented. The species, C. opaca (D. Don) Tagawa was encountered during recent fieldwork in Mount Berinchang, Cameron Highlands. Previously, it has only been recorded from Mount Kinabalu in Sabah, East Malaysia. A description, photographs, and a distribution map of known occurences in Malaysia have been provided.
The following new families of ferns are validly established: Taenitidaceae, Nephrolepidaceae and Loxogrammaceae. Information on the nomenclature and taxonomy of them are supplied.The Taenitidaceae are regarded as a group to be classified at the side of the families which form the order of the Pteridales, particularly of the Hemionitidaceae, as one of the branches independently originated from the same schizaeoid stock. The genera Taenitis, Schizolepton, Platytaenia and Holttumiella are ascribed to the family.The Nephrolepidaceae are considered as closely related to the Oleandraceae in the order of Davalliales. Nephrolepis, Arthropteris and Psammiosorus are treated as members of the family, although apparently very different from each other.The Loxogrammaceae, whose name had been already established but not validly published by Ching, are interpreted as the remnant of a synthetic group from which both the Grammitaceae and the Polypodiaceae could be derived.The following new combinations for species and varieties belonging to one or another of the three families are established. Platytaenia diversifolia (Holttum) Pic. Ser., Taenitis brachysora (Bak.) Pic. Ser. with its var. brachysora and var. polypodioides (Bak.) Pic. Ser., and Anarthropteris lanceolata (J. Smith ex J. D. Hook.) Pic. Ser.In addition the new generic name Tryonella is established to replace the illegitimate name Heteropteris Fée non H.B.K. Two species are ascribed to it: Tryonella lonchopbora (Mett.) Pic. Ser. and Tryonella crispatula (Bak.) Pic. Ser.
The Athyriaceae are a large and complex family. The phylogenetic relationships among the genera in the family are not well understood, and the systematic positions of Neoathyrium crenulatoserrulatum, Kuniwatsukia, Pseudocystopteris, Diplaziopsis, Allantodia and Callipteris have long been controversial. Nucleotide sequences of the chloroplast trnL-F region from 34 species of the Athyriaceae and three species in two other families were analyzed to gain insights into the intergeneric relationships. The phylogenetic relationships of the genera were inferred using the neighbor-joining and maximum parsimony methods, and both methods produced largely congruent trees. These trees reveal that: (1) the results of the trnL-F region sequence analyses and those of the rbc L gene sequence analyses are largely congruent; (2) Neoathyrium crenulatoserrulatum (Makino.) Ching & Z. R. Wang should be transferred to Cornopteris Nakai; (3) Pseudocystopteris Ching and Kuniwatsukia Pic. Serm. should be placed in Athyrium Roth; (4) Diplaziopsis C. Chr. belongs to the Diplazium clade; (5) Allantodia R. Br. emend. Ching, Callipteris Bory should be included in Diplazium Sw.; (6) Athyriaceae may better be divided into five subfamilies: Cystopterioideae, Athyrioideae, Deparioideae, Diplazioideae and Rhachidosorioideae.
Trophopods, persistent petiole bases containing starch, are found m all members of the fern genus Athynum (Aspleniaceae) in North America north of Mexico. Trophopods are produced on both cataphylls and foliage leaves, remain on the plant for several years, and contribute significantly to the starch storage capacity of the plant. The characteristic trophopods of A. pycnocarpon, A. thelypterioides, and A. filix-femina can be distinguished in the absence of leaves and on dried specimens. Those of A. alpestre are, however, very similar to those of A. filix-femina. Trophopods may have wider comparative application in the genus as a whole.
The first part of this paper deals with the nomenclature of some genera of ferns. The following new generic names are proposed to replace the illegitimate names now in use. Lecanolepis for Lecanium Presl non Lecanium Reinw.; Kuniwatsukia for Microchlaena Ching non Microchlaena Wall, ex Wight et Arn.; Phymatopteris for Phymatopsis J. Smith non Phymatopsis Tul. ex Trevis. Phymatodes Presl must be regarded as a nomenclatural synonym of Dipteris, and the genus now named Phymatodes remains without a name. Phymatosorus is established as the new name for it. Dryopteris subgen. Steiropteris is raised to the rank of genus. The new combinations for most of the species of the above-mentioned genera are made under the new generic names. Some genera are studied as far as their typification is concerned, and their lectotypes are designated as follows. Trichomanes: T. crispum; Micropodium: M. longifolium; Eremopodium: E. sundense; Ragiopteris: R. onocleoides; Blechnopsis: B. orientalis; Phymatodes: Ph. conjugata; Goniophlebium. G. articulatum. The place of pubblication of Neuromanes and the type (N. pinnatum) are established. Additional remarks on the nomenclature of the above-mentioned genera are made, and on the implications derived from their typification are pointed out.