Table Of Content720
Annals
of the
Garden
Missouri Botanical
Table Chromosome numbers Lycopodium based on
1. in sens. lat. 11.
Common
Base number Anomalous
numbers
G
enus 11 denominator Numbers
1 reported' reported genus^
in
Huperzia 6 X 11 66 132
67 67
134
68 68
136
204 X)
(3
Phlegmariurus 6 X 11 66 132 128 H.
ca. refiexa
{Huperzia) 68 136
275 x
68?)
ca. (4
3x11
Lycopodium
34 34 22 clavatum
L.
90-92
31 L. jussiaei
68
102(3 X
34)
2x11
Diphasiastrum
23 23 48 wightianum
L.
7x11
Lycopodiella 78 78
156
6x11
Pseudolycopodiella 35 35
{Lycopodiella) 68
68
70 70
5x11
Palhinhaea
52 36 cernua
104
L.
1
{Lycopodiella)
156(3 X
52)
54
108
55
110
165 X
ca. (3 55)
&
For
a discussion of the classification used here see Wagner
' Beitel (1992).
= For references to these numbers see Table 2. Chromosome numbers in Lycopodium sens, lat
&
Beitel, 1992). Aneuploid changes account Huperzia
The 128 chromosomes
for pairs of in
common
the denominators shown made
here, and count
poly- (Table an approximate
refiexa
1)
is
ploidy results in further changes shown of the
in the actual by Walker who suggested that is
966), it
1
(
numbers =
reported. 132,
same n
order of magnitude as a count of
The
anomalous numbers
listed in the last column which number other huperzias.
a reported in
is
of Table can be interpreted the
1 in several ways. The substantiating
photograph of a figure
Lycopodium =
clavatum
Wiih n 22 from cernua (Ku-
Bolivia count 136 Lycopodiella
of
pairs in
most Whe
likely a taxon different from
is the worldwide
=
name
species of that that has « 34. numbers of
Diphasias-
Lycopodium chromosomes
ing with in
trum wightianum = a
with « 48 was counted by photograph
this size, a drawing in addition to a
who
Ninan
(1958), wrote, *'The With
bivalents
at dia-
kiuesis exhibit very peculiar shapes and number for L)'
are
of to the hypothesis that the base
presenting
diflferenl sizes, unreasonab^
difficulties in interpre- copodium
not
sens, 11,
lat. is it is
One
templed
tation."
is to think that D. - .olyploKl
wightia- to assume many aneuploid and p
that
nam
a tctraploid based on n ?loo«
is
23,
the only
nees coidd have accumulated
be
based
--
w^"«"
cnanges
iiisiory oi tne genus. :>ucn
—
be
than
on 11 number other
assume
for to a
'
Lycopod
nurnbe^
greater
e.g., 7 or 17, would require an even
01Igaard
(1'
attempte^l
have
of
alterations. Earlier studies
numbe
Thi?
a
of
rs require the existence of exact multiples
.
pcrhap
number e.g..
pothesized base as a criterion,
^
The
n two
31.
species, however, Huper-^^
are placed
Lycopodium 136
in 68 and in
sens,
str.,
ent groups by
011gaard Takami.
(1987).
which 17
are exact multiples of (e.g.,
all
Wagner
Number 3 721
Volume
79,
Cytology Lycopodium
of
1
I
Table Chromosome numbers Lycopodium
2. in sens. lat.
Chromosome
Photo
or
Species Locality number drawing Reference*
Huperzia
&
chinensis (Christ) Japan n 68 Photo Takamiya Kurita
Ching
(1983)
&
herterana (Kumm) India n 132 Drawing Mehra Verma (1957)
&
Sen Sen
I
I
=
herterana 1 India 2n 405 Photo, Drawing Ninan (1958)
ca.
+
45
(180
II I)
= &
lucidula (Michx.) Canada 2n 264 Love Love (1958)
Trevisan
n^ &
Wagner
lucidula U.S.A. 67 Photo, Drawing Beitel F.
(1982)
&
miyoshiana (Makino) U.S.A. n 134 Soltis Soltis (1988a)
Ching
&
=
occidentalis (1988a)
(Clute) U.S.A. n 134 Soltis Soltis
ca.
I
&
Beitel Mickel
&
I selago (L.) C. Martius Canada 2n = 264 Love Love (1958)
&
Schrank
=
selago
2n 90 Sorsa(1962)
Finland
ca.
=
45 Sorsa (1963b)
n
ca.
selago Manton(1950)
Great 113 37 Photo, Drawing
Britain ca.
II, I
&
=
selago 2n 264 Love Love (1958)
Iceland
&
selago 2« = 264 Love Love (1966)
U.S.A.
selago Wagner
U.S.A. n 134 Photo, Drawing F. (this paper)
&
selago Tak Kur
var. acumina- Jap an n 136 in Mitui
tum
Sugimoto (1980)
serrata (Thunb. ex India n 264 Photo, Drawing Ghatak (1965)
Murray)
Trevisan
&
serrata = Takamiya Kurita
68 Photo
Japan n
= (1983)
136 Photo
n
serrata - Takamiya (1984)
Japan 2n 204 Photo
vernicosa & - Drawing Ninan (1958)
(Grev. India n 136 Photo,
Hook.)
Trevisan
Huperzia
{Phlegmariurus)
&
Takamiya Kurita
Drawing
<^^yptomerina (Max- Japan n 136 Photo,
(1983)
im.)
Dixit
dichotoma = Sorsa Fabbri (1965)
in
(Jacq.) Puerto Rico n 132
ca.
Trevisan
&
Drawing Takamiya Kurita
fordii (Baker) 136 Photo,
Dixit Japan
(1983)
Ninan
Drawing (1958)
^miltonii Photo,
n 136
(Spreng.)
India
Trevisan
= 30- Sorsa in Fabbri (1965)
^^nifolia
(L.) Treviisan Puerto Rico ca. 1
rt
140
Drawing Ninan (1958)
Photo,
"^^crostachys 136
n
(Spring) India
Holub^
Ninan
Drawing (1958)
Phlegmaria 136 Photo,
n
(L.) India
Rothm.
&
Takamiya Kurita
^ Drawing
phlegmana 275 Photo,
« ca
Japan
.
(1983)
Drawing Ghatak (1965)
Photo,
Ph'Uantha & 170
n
(Hook.
India
Am.)
Holub
«= Drawing Ninan (1958)
Photo,
136
Ph'llantha'
India
n- Drawing Ninan (1958)
Photo,
& 136
P^icherrima
(Hook.
India
722 Annals of the
Garden
Missouri Botanical
Table
Continued-
2.
Chromosome
Photo
or
number
Species Locality drawing Reference
=
pulcherrima^ India 2n 330-340 Mehra & Verma
(1957)
=
reflexa (Lam.) Trevi- Jamaica n ca. 128 Walker (1966)
san
saururus (Lam.) Trevi- Bolivia n 132 Drawing RoUeri (1982b)
san
&
sieholdii (Miq.) Holub Japan n 136 Photo, Drawing Takamiya Kurita
(1983)
squarrosa (G. Forster) India n 136 Photo Ninan (1958)
Trevisan n 138 Drawing
Lycopodium
= &
annotinum L Canada 2n 68 Love L5ve (1958)
=
annotinum
Finland n 34 Drawing Sorsa (1958)
=
68 (1963b)
2/1 Sorsa
= &
annotinum
Japan « 34 Photo Takamiya Kurita
(1983)
=
Sweden
aiinoiinutn 2n 58
Ehrenberg (1945)
ca.
annotinum Sweden
n 34 Photo, Drawing Manton (1950)
annotinum
U.S.A. In 50
Dunlop (1949)
ca.
annotinum &
var. acri- Japan n 34 Photo Takamiya Kurita
foViuni Fern.
(1983)
annotinum ah = &
snbsp. Iceland 2n 68 Love Love (1958)
&
Love
pestre
Love
= &
casuarinoides Spring Japan 2n 68 Takamiya Tanaka
Photo
(1983)
clavatum L =
Bolivi « 22 (1982a)
la RoUeri
clavatum Canada = &
2« 68 L5ve L5ve (1958);
Love (1976)
clavatum
Ecuador =
n 34 987)
gfllgaard 1
(
clavatum
Finland n = 34
Drawing Sorsa (1958)
=
2r 68 (1963b)
Sorsa
clavatum = &
India Verma (1957)
n 34 Drawing Mehra
clavatum =
sens. lat. India 68
n Drawing Ghatak (1965)
vatum
=
cla Great
Brilain n 34 Manton (1950)
Photo, Drawing
clavatum n=
Jamaica
34 Walker 966)
Photo
1
(
clavatum Japan = &
2n 68 Takamiya
Tanaka
Photo
(1981)
&
Tanaka
102 Takamiya
2rt Photo
(1982)
2n 136 Takamiya (1989)
Photo
clavatum
Sweden
=
2n
66 (1945)
ca. Ehrenberg
do
vat urn Taiwan = &
n 34 Shieh (1983)
Photo Tsai
clavatum
U.S.S.R. =
n 14
Drawing Baranov (1925)
X
clavatum
vcstitum Ecuador =
n 34
011gaard(1987)
clavatum
var.?. U.S.A, =
2n 60
ca. Dunlop (1949)
clavatum
^ub^p. Canada = &
(1958)
2a 68
Love
L5ve
megastackyon
&
(Fern. Biss.)
&
Love Love
clavatum
var. nippon-
Japan
Kuriu
n 34
Photo
icum Nakai
(1983)
contiguum
Klottsch
Ecuador =
n 34
011ga
dendroideum
Mirht.
Canada =
2n 68
Love
))
^ ^'V^iU-- ~ jr-^ ^
Wagner 723
Number
3
Volume
79,
Lycopodium
Cytology
of
1992
Table
Continued.
2.
Chromosome
Photo
or
Species Locality number drawing Reference*
=
jussiaei Desv. 90-92
in Poi- Ecuador n Photo, Drawing 011gaard (1987)
ret
=
jussiaei 34-36
n Wilce (1972)
= &
lagopus (Laest. ex 2n 68 Love Love (1958)
L
Hartm.)
Zinzerl.
I
ex Kuzen.-Proch*
magellanicum
Ecuador n 31 011gaard (1987)
(Beauv.) Sw.
mageUanicum =
Argentina n 31 Photo 01]gaard(1987)
= &
ohscurum
L. Canada 68 Love Love (1958)
2/1
&
ohscurum
Japan n 34 Photo Takamiya Kurita
(1983)
ohscurum &
Wagner Wagner
U.S.A. 34 Drawing
n Photo,
(1966)
I
ohscurum =
Dunbp
U.S.A. 2n 50 (1949)
ca.
vestiturn Poiret Ecuador n 34 01lgaard 987)
»
1
(
Diphasiaslrum
I
alpinum = &
(L.) Holub Canada 2n 48 Love L5ve (1958)
nlpinum =
22-24
Finland n Drawing Sorsa (1963a, b)
=
2n 44
alpinum =
Manton
Great Britain n 24 25 Photo, Drawing (1950)
alpinum & = &
Scandinavia 2n 46 Love Love (1961)
Canada
&
complanatam Ho =
(L.) Canada 46 Hersey Britton (1981)
2rt
lub
&
complanatum & =
Canada 2n 48 Love Love 958,
ca
1
(
=
1961)
Scandinavia 2n 46
complanatum =
Finland n 22-24 Drawing Sorsa (1963a)
complanatum
= Kukkonen
(1967)
Finland n 24
ca.
&
complanatum Tak Kur MHui
Japan n 23 in
980)
( I
complanatum
= Wilce 965)
Drawing
Labrador n 23 1
(
&
complanatum 98
x = Heraey Britton
Canada 2n 46 Photo
tris- ( 1 1
tachyum?
complanatum
Dunlop 949)
var. U.S.A. n 40 Drawing ( 1
^longatum
&
<iigitatum = Ilersey Britton (1981)
(A. Braun) Canada 2n 46
Holub
&
Love Love (1958)
digitatum^
48
Canada 2n ca
.
digitatum 2n- Love (1976)
Canada 46
WUce
^igitatum = (1965)
Drawing
46
2n
U.S.A.
& n- Walker (1966)
fowcettii (Doyd Photo
maica 23
Ja
Underwood)
Holub
&
9«
^ habereri (House) Canada 2n = 46 Photo Hersey Britton ( J J
Holub
Wagner
^ • 980)
habereri F.
23 ( 1
U.S.A.
DatnboMt 962)
=
^isileri (Rouy) Holub Germany 46 Drawing ( 1
2/1
Love (1976)
=
^Mobinifolium
46
Canada
(Willd.)
2/1
Holub
Wagner
980)
- F.
^bifufolium 23 ( 1
Canada
Love 976)
^chense =
(Rupr.) Ho- Canada 2n 46 { 1
lub
Wi!.r (1965)
=
^uchense
23
n
Labrador
&
Love Lovr 966)
uich -
( 1
rri4e 46
U.S.A. 2/1
Annals of the
724
Garden
Missouri Botanical
*
Table Continued
2.
Chromosome Photo
or
number drawing Reference*
Locality
Species
=
2n 46 Love
Japan (1976)
sitchense subsp.
ni-
&
koense
L. L.
&
an n 23 Photo Takamiya Kurita
Jap
sitchense var. ni-
(1983)
Takeda
koense
=
Ho- Ecuador n 23 011gaard(1987)
thjoides (Willd.)
lub
&
=
Canada 2n 46 Photo Hersey Britten (1981)
tristachyum (Pursh)
Holub
&
=
Canada 48 L5ve Love (1958)
tristachyum 2/1 ca.
=
Canada 2n 46 L5ve (1976)
tristachyum
tristachyum U.S.A. n 23 Drawing Wilce(1965)
&
Holub Taiwan n 68 Photo Tsai Shieh (1983)
veitchii (Christ)
&
48 Drawing Ninan (1958)
wightianum (Grev. India n Photo,
Hook.) Holub
=
X (Rouy) Holub Germany 2n 46 Drawing Damboldt (1962)
zeilleri
=
Wagner
X U.S.A. n 23 (1980)
F.
zeilleri
Lycopodiella
alopecuroides U.S.A. n 78 Photo, Drawing Bruce (1975)
(L.)
Cran.
alopecuroides x ap- U.S.A. n 78 Drawing Bruce (1975)
Photo,
pressa
=
x
alopecuroides pros- U.S.A. 78 Drawing Bruce (1975)
Photo,
71
trata
appressa (Chapman) U.S.A. n 78 Photo, Drawing Bruce (1975)
Cranfill
x
appressa prostrata U.S.A. n 78 Photo, Drawing Bruce (1975)
&
inundata Holub Canada 2n 156 Love Love (1958)
(L.)
inundata Canada 2n 156 Love (1976)
inundata
Finland n 78 Drawing Sorsa (1961)
mun
data Great 78 Manton (1950)
Britain n Drawing
Photo,
inundata
U.S.A. n 78 Drawing Bruce (1975)
Photo,
margueritae Bruce, U.S.A. n 156 Drawing Bruce (1975)
Photo,
&
Wagner
Beiter
prostrata (Harper) U.S.A n 78 Drawing Bruce (1975)
Photo,
Cranf,
subappressa Bruce, U.S.A
n 156 Drawing Bruce (1975)
Photo,
&
Wagner
Beitel**
I*^eudolycopodiella (Lycopodiella)
&
caroliniana (L.) Holub Japan - 68 Takamiya Kurita
Photo
rt
(1983)
&
caroliniana Japan Kurita
68 Takamiya
n Photo
(1983)
caroliniana =
U.S.A
« 35 Photo, Drawing Bruce (1976)
=
« 70
2n=
115«
meridionalis Un-
(L. Jamaica n 69 Walker 966)
«^
ca. 1
(
derw. &* F. Lloyd)
Holub
Palhinhaea
{Lycopodiella)
&
cernua
Carv. Kurita
(L.) Japan
n 108 Takamiya
Photo
&
V«»c. Franco
(1983)
cernua
India
n 104 (1958)
Ninan
Photo, Drawing
Number Wagner
3 725
Volume
79,
Lycopodium
Cytology
1992 of
Table Continued.
2.
Chromosome
Photo
or
Species Locality number drawing Reference
cernua n 104 Photos Ghatak(1965)
Indi
la
n 156 Drawings
208
n
cernua
Indila n 104 Photo Kuriachen (1965)
n 110
n 136
=
n
ca. 160II,
201
=
cernua
Jamaica n ca. 165 Photo, Drawing Walker (1966)
=
Trinidad n 165
ca.
cernua &
Taiwan
n 102 Photo Tsai Shieh (1983)
For
references, see Literature Cited. The following references were not seen and therefore not included in this
&
Hadac,
table: E. V. Haskova. 1956. Taxonomicke poznamky o tatranskych rostlinach ve vztahu k jejich Bratislava/
&
cytologii. Biol6gia Brat. 11: 717-723. Love A. D. Love. 1948. Chromosome numbers of northern plant species.
I Icel. Univ.
Inst. Appl. Sci., Dept, Agric. Rep. B. 3: 1-131.
As Lycopodium
lucidulum.
'
I
macrostachys
and
phyllantha are treated as synonyms by Ninan.
As Lycopodium
' setaceum,
As &
clavatum
* subsp. nionostachyum (Grev. Hook.) Selander.
*A9
complanatum
var. me,
Jlabellifor
As |Wthern
* appressa'' See Bruce
et ah (1991).
As
"appressed
inundata"
See Bruce
et al. (1991).
I
•Somatic
count
of a presumed hybrid— 105?
triploid possibly
^
AB
Kurita, 1983). Such and
suggestions do not take formation of a sterile diploid, (2)
is
aneuploidy
into consideration. [AABB], which formation of a allopoly-
fertile
is
chromosome
doubling.
through endomitosis or
ploid
found
Lycopodium another pattern
In sens. lat., is
Allohomoploid
Nothospeciation
in
formed
which sexual nothospecies
in a fertile is
IrCOPODlLM
SENS.
LAT.
homo-
parents and hybrid are
without doubling, an<]
Even AA BB
though x
same
the potential for of the ploidal level,
self-fertilization ploid, i.e.,
exists, Lycopodium
in
[AB]
sens. lat. as in the ho-
all
mosporous
and
pteridophytes with gametes of both sex- (The use of brackets here be!
** produced oposcd
the same competent hybrid,
in gametophyte, evidence reproductively
for high & Wagner
frequencies of intergametophytic matings Werth (1990).)
has been & DiphasiasUum
found. Soltis Soltis (1988b) studied a In Figure 3 three species of
total of 22 complanatum, and silchense—-are
widely
scattered populations of L, cla^ digitatum,
^^fum,
L, annotinum, and Huperzia miyoshiana, shown with D. tri~stachyum, a .species that liybrid-
•"n, The from
using resulting tlu^se
electrophoretic liybrjrl^
analyses of polymorphic ^^^^
[xj
[x]
[x] and
oci,
calculated low hahereri, zeilleri,
estimates of intragametophytic crosses, />
have been found
^f-ferlilization. They of which in llie
concluded, sahinifolium (all
therefore, that
genomes
*n^se
species predominantly are fertile to the extent that their
Be- wild),
cross-fertilize.
chromosomes, and
<^ause /.v/^^r.^^;..™ ^„„„ show complete pairing of their
with the exception
lat".,
norma Tl M
^» ^ycopodiella 5).
sens, has under-
lat., entirely
ground
{n
gamelophytes, lumber
had been presumed the
in
it
parents involved
same the
as that for all
imming 23) the
is
&
98
^tl^rground Wagner, 1980; Hersey Britton,
1 1),
(F.
Lycopodium
of
germination
Unfortunately,
bridization
vnxh
On ran onlv be carried on\ diflfirulty
a^«/^rf>^
I
&
98 ;_Whi.Ucr Websler, 986)
^^'ing
and 977, 1
have turned Whittier, 1
interspecific hybridization 1 1
common (W
**"» to be
in the Lyci podiaceae
**<
be mnde. Som*- indiratimi
have yet to
normal spores
1985).
however, to hy thr fart
ntt^-^t.-d
ypicaUy of ihnr fertility, i«
in plants, nothos
D
X
hnhrrrri.
populations of
Uon) we
isolated
find
which that
'\B,
726 Annals of the
Garden
Missouri Botanical
complanacum
sitchense
D. D.
tristachyum
D-
digitatum
D.
3
Figure Diphasiastrum. Diagram showing complanatum to form D.
3.
hybridization of D. tristachyum with D.
n n .. ^ ^ ^
...n.,;. .;..!,„....„ f„.™ ^- •.,- have
,.:.u .
,. taxa
. . , . x All
Aofterm*.
ifoliu
~ WUcc
n 23 chromosomes. from
pairs of Branchlet drawings show drawings are
relative sizes of leaves. Habit
(1965).
the g^*^
initia clavatum x vestitum, in
species,
L.
no
spore with parental species or only one
parent Lycopodium
i^uputiturn sseennss.,
str.
isir, .,
&
(Wagner Wagne
in
FertUe homoploid nothospeciation P|^"v^*
Homoploid
nothospecies Lycopodium
in
sens. phytes was reported in a classic
first
^^^^5
confined
Walke
although most reported examples quadriauriic
are Two
in that genus. and
P.
species, P. multiaurita
Bruce (1975) found
hybrids
in Lycopodiella interme i
swarm
be- in Ceylon, formed a hybrid of
tween alopecuroides and
L.
i, appressa, and parents.
be- occupying an between the
ecotone
tween i. alopecuroides and o^^
chromosome
L. prostrata,
with
hybrids were but without
fertile
^
genomes, same
pairing of the chromosome number ^«™^ number a^
Ur.A *k« ^i^^^^i^cmnft
Kli'no- oil
and
as the parents, morphologically normal
spores,
parental
species.
011gaard also
have
Homoploid Ceratopteris
hybrids of
Number 3 Wagner 727
Volume
79,
Lycopodium
Cytology
1992 of
ti^^r^j.
4
i V.
^*
»1
^ t
^
«
<
#
w
#
l»
I
»
#
i
^
^
j
'CURES
4, 5. 4. Photomicrograph of chromosomes of Diphasiastrum x sabinifolium at diakinesis with 23 pairs
—
chromosomes.
01 Scanning
5. electron photomicrograph of spores of D. xhabereri.
produced
miliar in the ferns resulting in either sterile allo-
i
and
(1974), homoploid
hybrids in the Cyatheaceae, diploids or fertUe allopolyploids (Beitel, 1 986, 1 988)
&
first reported by Conant No
Cooper-Driver (1980), allohomoploid hybrids have been reported in
are found
in Puerto The
Rico.
cyatheoid hybrids the genus,
backcross
and form
hybrid swarms,
but recombi-
nant
second
generation may become
hybrids
sta-
Discussion
bUized and
maintain
by means
their genetic
integrity
A number can now be made
of generalizations
selfing
In
the North American Lycopodium regarding the cytology of Lycopodiaceae. The basic
sens,
lat.,
the
morphological x
variation seen Diphasias- chromosome numbers are high, the lowest being
in the
trum =
hybrids seems clubmosses are other
clearly to be environmentally 23. In this respect the like
produced,
i.e., sun and shade forms 1979a, homosporous pteridophytes and unlike the hetero-
(Beitel,
b; Beitel et al., 1982). However, although we have sporous Selaginellaceae and Isoetaceae, which have
searched
for years, we have x numbers seed plants. Also, like other ho-
not found backcrosses like
"1 these Lycopodiaceae bear both
hybrids. This seems mosporous pteridophytes,
surprising since Di-
phasiastrum
same gametophyte and
species have been found be sex organs on the poten-
to pri-
niarily & The
outcrossers undergo intragametophytic mating.
(see above and can
Soltis Soltis, tially
1988b). from homosporous ferns
Hybridization produces the hybrid Lycopodiaceae differ in
original
and
if such hybrids apparent absence of apogamy and in a greater
retain this capacity, then con- the
tinued
outcrossing should ultimately lead to back- tendency for allohomoploidy, as illustrated pri-
crossing, Diphasiastrum
by
introgression, and hybrid swarms. Ap- marfly
nm To explain the curious **step-wise" increases
3nd may hTomosom'e base num-
it be may
that rarity a factor; there nc
is
^ot be enough 31-34, 52-55, 66-70, and 78,
23,
individuals of associated parental hers, i.e., I
we
species hypothesis that are dealing
to cross can only a
with. Related perhaps, the fact offer
is
some
»fiat species of Lycopodium here with a polyploid series, involving aneu-
are great
sens.
lat.
X
—clone formers and changes as a minor element, i.e., 2 11, 3
rely heavily on vegetative re- ploid
' -
X 11,5 X 11,6 X and 7 X The graded
1
be that there in realitv verv 1 1 1
is , .
mbe
oduction
clubmosses had
high
Unlike the original
that
Diphasiastrum and Lycopodiella, hy- possibiUty
mbe
idizat
I
1
728
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Missouri Botanical
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