Table Of ContentNumber 3
Volume
79, 0llgaard 689
1992 Neotropical Lycopodlaceae
I
)
growth may
are the following: habit (e.g., erect, as- to appressed leaves, be intermediate to the
I
cending, recurved, pendulous); stem thickness; shoot group of H, saururus.
differentiation (homoblastic or heteroblastic); leaf Almost exclusively neotropical, with a single spe-
(homophyllous,
differentiation heterophyllous, or cies {H. dentata (Herter) Holub) in the Azores and
gradually heterophyllous); leaf arrangement (num- Reunion. The number of species uncertain, per-
is
ber of leaves in whorls, or of orthostichies in spi- haps higher than 20. a group of weakly dif-
It is
crowding
rals), (distance between whorls); leaf out- ferentiated, erect and homophyllous, terrestrial
hne and shape
solid (e.g., thickness, vein species growing in open habitats, especially pioneer
t
prominence, margin curvature), margin shape (e.g., vegetation, in the montane forest zone.
teeth, rugulate, smooth), leaf dimensions, direction,
color, texture, surface (smooth, stomate
papillate),
(Lam.) Trevisan (Wild
eflexa (//. bifida
distribution, shape of epidermal shape and &
cells;
Holub; H, densifc (Baker) Rolleri
color of decurrent sporangium shape &
leaf bases;
Deferrari; H. leptodon (Herter) Rolleri De-
and
spore &
size;
size.
De-
ferrari; H. parvifolia (Nessel) Rolleri
The
genus seems
be undergoing
to active evo- Lycopodium hrutum Herter; L. rev-
ferrari;
lution, especially in the Andes. Once the plants are non
ersum rigidum Sw., Gmelin;
C. Presl; L.
J.
adapted
to the general environments
at different squarrosum Lam., non G. Forster, nee
L.
levels of altitude, few selective forces seem to be SW.; squarrosum Sw., non G. Forster,
L.
operating on
the morphological
features that are Mor-
nee Lam.; i. stellae-polaris (Herter) C.
used
for recognition and The
identification. species
Urostachjs jergii Nessel)
ton;
therefore
are often
difficuh to define.
common
The most and widespread species of
the group, polymorphic, both with respect to size
THE
HUPERZIA
SELAGO GROUP
and shape presently poorly understood.
of leaves,
With
gemmiferous arranged whorls of
branchlets, and a distinct Leaves denticulate, usually in
spore
type with truncate corners and concave, 6 or more. Tropical America.
pitted proximal
faces. ^/
Mam
distribution in northern and southern tern-
ifi
perate
regions, and mexiae
in mountains of the Paleotrop- Deferrari (type from Peru), H. (Copel.)
ics. The group & pearcei
is represented in tropical America RoUeri Deferrari (type from Peru), H.
h
a
single species. Holub from H. spongiosa
(Baker) (type Bolivia),
&
from Colombia),
Deferrari (type
Rolleri
(Rolleri)
uperzia
//,
serrata (Thunb. ex Murray) Trevisan and Huperzia rigida F. Gmelin) Holub (His-
(J.
(synonyms
H. catharlnae may
Holub; Ly- Some names represent valid
(Christ)
of these
paniola).
copodium
sargassifolium Liebm.) and distributions are
taxa, but their delimitations
Leaves ^s yet uncertain.
coarsely
serrate; annual constrictions
as
morphological types of rejlexa,
the Different //.
J'ong shoots
arising connection with the
in
occur
formation other closely related species often
of w*^" ^^
whorls
of Mainly
bulbiferous branchlets.
and
«n the together on large road banks or land slides,
Paleotropics,
rare in the Americas, with only
few
growth con-
compared under uniform
here can be
stations in southern Mexico, Cuba
(Sierra
in
may
such populations the individuals
aestra), In
ditions.
in Hispaniola, and a single enigmatic
Neighboring
coUection & exhibit subtle or obvious differences.
in
Brazil (Santa Catarina). (Mickel
may
more populations exhibit similar
^'tel, 1988: and distant
5A-C).
fig.
may
be
fne variation and thus taxonomic recognition
hybrid H. x
serrata lucidula reported
is
from are not
However, often the differences
indicated.
Hispaniola,
where
the second parent species
same
IS unknown matched by other individuals, neither in the
(Mickel, 1984).
and therefore the plants
nor other populations,
in
Some
tUc A\arac\fr nf individual aberrants.
v,a,r<.
TIJE
HUPERZIA
REFLEXA GROUP
intermediate bet
Leaves
features
lanceolate may without exhibiting hybri<I
to subulate, usually with den- be hybrids,
.
The
^'culate meioses.
margins, but sometimes Most spe- such as abortive spores or irregular
entire.
es With A example, of leaf shape and
spreading,
recurved differences, for
or reflexed leaves. slight
w
genus
species, adapted to the more exposed habitats direction, appear especially striking in the
repeated
are in
forest and Huperzia, which the features
liinit just above with ascending in
it
. rjT -*=* " -^a/-^ji
690 Annals of the
Garden
Missouri Botanical
\
Southern Ecuador, low para-
hundreds of leaves on a stem, creating regular and the fingerlike shoots.
mos i
1988: 13C).
recognizable patterns. At the same time these fea- (011gaard, fig.
tures, by which the individuals differ, appear to be
Huperzia Holub (Urostachys
christii (Silveira) or- \
without adaptive significance (Lellinger, 1989:
fig.
gaosanus
Nessel)
&
36; Mickel Beitel, 1988: IE, F; 0Ugaard,
fig.
1988: fig. 4A). Differing from H, reflexa mainly in the larger,
I
convex, more coriaceous leaves. South-
abaxially
Huperzia sintenisii (Herter) Holub eastern Brazil, on banks at high akitudes.
Tall and robust, very simQar to H, reflexa, but Huperzia Holub
rostrifolia (Silveira)
with a thick stem and broad West
relatively leaves.
A known perhaps
poorly high-altitude species,
Indies.
Southeastern
not from the preceding one.
distinct
'
Brazil.
Huperzia
Trevisan {H. blepharodes (Max-
affinis
on) Holub; Urostachys involutus Nessel)
0Ug. {Lyco-
Huperzia riobambensis (Herter) B.
podium Morton)
Leaves relatively large and wide, borne castoris (Herter) C.
in
al-
ternating whorls of with
5, long, slender, hairlike
montane
A
upper
robust representative of
large,
"PP^'
coriaceous
and low paramos with broad,
l^!!..Z'^l^^'^t^^}^^^ fo'-ests
montane
1988:
forest (011gaard, 4B).
fig.
whorls
arranged distant
reflexed, glossy leaves in
of 5 or Colombia.
6.
0U \
Lycopodium
(Herter) Holub; polycarpum
Huperzia 0Ug.
B.
beitelii
(Sod
m
U Coarse, erect or ascending to erect, to 1 tall,
Urostachys ringshausenii
Nessel; roland-
with re-
sometimes divisions
with constricted distal
ii-principes Nessel; dingesianus
Nessel)
[/.
somewhat clasping
duced, appressed ascending,
or
Forms amply branched Venezuelan
individuals with narrow
leaves, an unusual feature in the group.
usuaUy
shoots, with distant whorls of 5 short, Guayana. Low low open forest of high al-
scrub,
strongly recurved leaves. Central America, Colom-
152-155).
1989b:
titudes (011gaard,
On
bia to Bolivia. banks upper montane
in
forest
1988: Huperzia Trevisan
(011gaard, 4C). sieberiana (Spring)
fig.
A cov-
& densely
Huperzia friburgensis (Nessel) B. Wind. very robust high-altitude species,
011g.
leav
ered by narrow, subulate, entire ,
coriaceous,
Resembling common,
the small H.
of
reflexa AntiUes.
Lesser
arranged 10 more.
in whorls of or
Brazil, but taller, less branched, and with
entire
Southeastern
leaves.
Brazil.
Huperzia
unguiculata
B. 011g-
con£^
A with
Huperzia
hemleri (Nessel) B. robust species of mid-altitudes,
011g.
recurv
strongly
ceous, nearly usually
entire,
Superficially resembling H. sellowiana
(H.
9- and
'^^^'^^''^
leaves, arranged in whorls of 1 1
,
brongniartii group) and growth
in size habit, but
(0Ug
may
with denticulate leaf margins.
Its origin involve
2B).
fig.
this and a species in the H. reflexa group. The
com-
spores are normally developed. Southeastern
Bra- {Urostachys
Huperzia
intermedia Trevisan
^bel ^
mutatus
Herter)
,
itats in forest (Nessel, 1927,
t. 9).
nd
an
A branching
slender, often divaricately
# nn sand-
.v,„
ambling .
Venezuela*
Guadeloupe,
stone; leaf margins
entire.
A Espmto
and
large robust
local high-altitude
represen-
Minas Gerais,
southeastern
Bahia,
Brazil:
spe-
tative of the group, with broad,
distant leaves. Costa BraziUan
Santo. Perhaps to
as closely related
Panama
Rica, (Lellinger, 1989:
28). especially
fig.
cies of the H. brongniartii group,
ifc
Huperzia
austroecuadorica
B.
011g.
0Ug-
Huperzia
chamaeleon B.
(Herter)
Approaching
the growth
habit of the
sau-
//.
leaver
rurus group, with ascending-appressed
entire
leaves on With
very narrow, recurved, linear,
4
691
Number 3 0llgaard
Volume
79,
Lycopodiaceae
Neotropical
1992
prominently decurrent leaf bases on the and relatively distant whorls of sharply reflexed
without
&
1989: leaves (LeUinger, 1989: 27; Mickel Beitel,
somewhat tumid stem. Costa Rica (Lellinger, fig.
1988: 2E; 011gaard, 1988: lA, 2A).
figs.
fig.
18).
fig.
Huperziafirma Holub {H. coriacea (Rol-
(Mett.)
Huperzia arcuata
B. 011g.
&
from Colom-
Caldas,
Rolleri Deferrari,
leri)
by
Deviating from the preceding species mainly
synonym)
bia, perhaps a
is
and nodding shoot apices.
basally twisted leaves
its
Huperzia more
Resembling but robust,
reflexa, may become
entirely
Large epiphytic individuals
and with more coriaceous leaves. This and the Huperzia dicholoma,
and resemble
pendent, thus
members
foUowmg from
other
species are distinct Colombia
Southern
^^^ ecologically distinct.
j^^^
of the group because of thick-walled, nearly iso-
and Ecuador. Occurs sympatrically with the pre-
Andes
diametric epidermal of the leaves. of
cells
ceding species, but, in addition to the forest floor,
Venezuela and Colombia. Some Colombian plants 1988:
(011gaard,
occurs epiphytic habitats
also in
usually referred to this species, which are larger
ID).
fig.
m
may
in parts and up deserve
all at least to 1 tall,
1988:
recognition (0Ugaard,
as a distinct species
Holub {Vrostachys
Huperzia (Hieron.)
2D). lechleri
fig.
lehmannii Herter)
Huperzia
urhanii (Herter) Holub from Huperzia hippuridea,
Doubtfully distinct
only by the narrower leaves, usually ar-
differing
Closely
related to the preceding species; differing
number of orthostichies.
greater
ranged in a
hy
the smaller, sigmoid leaves, usually arranged in
southern
Overlaps the dis-
^^^ Southern Peru, Bolivia.
very
close whorls of 11-14. Ecuador (011gaar-d^
hippuridea.
of
tribution //.
1988:
2C).
fig.
&
Wind. {Urosta'
THE HUPERZIA BRONGNIARTII GROUP Huperzia nuda (Nessel) B. 011g.
nudiusculus Herter)
chys
|T
Usually
weakly
erect, differentiated, terrestrial
smaller than,
many but consistendy
plants, of them with a characteristicaUy hy- Very similar to,
Sao
Rio de Janeiro,
gromorphic, preceding species. Brazil:
growth with the
bottle -brush habit,
-like
and
cloud forests is
sparsely Occurs high-altitude
branched stems, and perpendicular, Paulo. in
long,
monomorphic,
and aspect believed
linear entire leaves; this
1927:
14).
inost apparent Huperzia hippuridea and relatives (Nessel, t.
in its
K
closest
relatives, H, arcuata, H, lechleri,
viz,
nuda,
H, and H. mexicana, Neo- Holub
recurvifolia, mexicana (Herter)
Huperzia
Many
^ropical, approximately 22 species
species.
pur
hip
from
distinguish //. i-
to
belonging Often difficult
in semishaded habitats in
forest floor
by the
epiphytic,
being
from this in
^Pper montane dea, differing
forest.
leaf
induced variation hi
seasonally
of
presence
along the shoots,
constrictions
appearing as
length,
Huperzia
prominently
hippuridea Holub holiviana and by
(Christ) leaves,
(//. more coriaceous
by
&
^Kosenstock) Rolleri Deferrari; H. montana reddish leaf bases. Leaf base char-
often
jg^^yrrent,
&
(L. Underw. poseidonis H. pithyoides (see
F. Lloyd) Holub; L. approaching those of
^^^^^^
"
'^'•"^
" "'
(Herter) C. Morton) *"
under '
"
be
Growth
habit with sparsely
bottle-brush-like,
group
branched present
mono- the
stems, and perpendicular,
long,
features of this spe-
xeromorphic
The
relatively
linea.
be
'espect
to stem and crowding
thickness, direction
durmg dry pe-
desiccation
more to
liable
^Heaves. which
Widely Central is
distributed, occurring in
America,
riods
the Greater Venezuela, including
Antilles,
observat
enezuelan Some inose oi ij. "^rr
Guayana, and south Bolivia. ^ ... ^^^^^
to .
conHrnung corre-
th.s
coexisting,
^al two species
from the
populations more
deviate consistently
habitat,
difTer.nces w.tli
morphological
Ine of the
aspect of form of the lation
the widespread, typical Amenca, Greater
Central
Northern
missing.
"Pecies, are
pop-
e.g., Ecuador. Here both typical
in
Ant (Fig. 1).
•^ations stems
occur, and populations with thick
692
Annals
of the
Missouri Botanical Garden
Figures
2. HupcTzia
1,
1- mexicana,
a. Habit. of leaf-
^- Detail of sporangiate Cross section
Huperzia division.-
l, c.
pithyoides.—^i.
Habit. ,
ot
°^ ^P^'-angiate division.-c. -d. Detail
proximal Cross section of leaf.
division, with
decurVenTlerf
bases
Huperzia
recurvifolia RoUeri
Venezuelan
leaves
with
acute, concolorous apices.
Deviates from H. ^"^y^na
hippuridea
(0Ugaard,
989b).
maiiily
in the 1
soft-
ly recurving
rather
than
sharply
reflexed
leaves
-^^^«"
^^'^ '^'^^"- ^"/^--«
°^ Holub
(Herter)
^h
ish lleeallftappiitc^e^s^.^^V^e^nezuela,
Guyana,
Brazil: Bahia.
nd
Mainly ^ a
in open, rocky stems
sandstone '^^'"^ ""^bust species with thick
habitats
of the
Roraima
varying
formation densely
crowded,
(RoUeri, 1989: 209-215) lanceolate, thick leaves,
Ex-
from
reflexed ascending-appressed position-
to
Huperzia
huberi in-
B. hibiting highly morphological features,
011g. variable
spe
termediate between forest
of the
those typical
Uo.eIy
Closelv rreellfaltiefdd t« *U^
to the
precedii- ng
species, and
of th-
cies mentioned .bov« .nd
those tvuical
oc
saururus
paramo
species mentioned under the H.
Number
Volume 3 Ollgaard 693
79,
Lycopodiaceae
1992 Neotropical
Ecuador
group. to Bolivia. Habitats transitional
between the upper montane
forests (ceja andina).
(011gaard
4D).
fig.
\
Huperzia
loxensis B.
! 011g.
Resembling Huperzia more
hippuridea, but
ro-
bust, with thicker stems and densely crowded, mi-
nutely
denticulate leaves. Southern Ecuador. Like
the preceding
species in habitats transitional be-
tween the upper montane and low paramos
forests
in a restricted area on the mountain crests east of
Loja
(Ollgaard, 1988:
3B).
fig.
Huperzia
binervia
(Herter) B.
011g.
As
H.
in weddellii,
exhibiting variable features
intermediate between
those of the forest and para-
* mo
species. In exposed may
habitats old individuals
exhibit the gradual
transformation from
the
bottle-
brush
aspect
of the proximal
divisions, to fingerlike
distal
divisions with closely appressed leaves.
Northernmost
Peru. This narrowly endemic
species
occurs
in habitats at or just above the forest limit
(01Igaard, 1988:
5D).
fig.
Ine
following seven endemic
species are to
southern and
eastern Most them
Brazil. of are rare
and
have
very
restricted distributions rupestral
in
habitats on
isolated mountaintops.
{Ly-
011g.
ifi
copodium
ouropretanum
Urostachys
Christ;
hennebergorum
Nessel)
Resembling
//. recurvifolia, but with shorter,
"iore
coriaceous
leaves, and more
often divari-
ound
'/
entire-margined,
coriaceous and more
leaves, a
open
branching Habit.—
Huperzia treilubensis.—&. b.
pattern. BrazU: Minas Gerais, Rio Figure 3.
e
Janeiro. The plant.— Detail of sporangiate division.
most Juvenile c.
widespread and morpholog-
'^aUy
least
specialized of the BrazQian
species.
shoot base. Brazil:
gated to a thickened bulblike
//
^Perzia & Minas Gerais, a very local species (Fig. 3).
regnelUi (Maxon) Wind.
B.
011g.
ery
{Huperzia
similar
to the preceding species, but with Huperzia deminuens (Herter) B. 011g.
^educed
and
appressed Holub)
leaves in the distal divisions, luederwaldtii (Nessel)
/azil: Minas
Gerais. Very no
rare, recent collec-
which
With divisions, in
erect fingerlike distal
•ons
being
known.
(Maxon,
1914:
23).
t. convex
and strongly
appressed,
the leaves are
Rio
//, Perzia
treitubensis
(Lyco-
(Silveira) B. 01Ig. rare rupestral species.
podium
hoehnei
(Nessel) RoUeri; inflexum
L.
&
Wind.
SUveira;
Urostachys Huperzia itambensis B. 011g.
capri Herter)
L
eaves A species with short, closely
of proximal crowded, highly specialized
divisions densely
leaves
^sely somewhat carinate
appressed, and abaxially
at the stem base often aggre- imbricate
Annals
694 of the
Missouri Botanical Garden
5mm
mm
5
5a
a
Habit.-b. rubra
Huperzia
T^^^""'^-!- Detail of sporangiate
hii-h division. 5.
of'snorW
T
iDii. D. Dueettadiill ot sporangiat're *
division.—
n „f
n.>f=i;i j._
proxi; mali •
on
the
erect
Stiffly stems.
Brazil: Minas
Gerais distinctive
other highly specialized species of very
known
only from
the type &
locality (0Ueaard
ra
<v appearance
g (Fig.
4).
U'indisch, 1987:
fig. IB).
Huperzia
rubra (Cham.) Trevisan
Huperzia
mooreana
(Baker) Holub
{H. sydo
a^
ram &
stems
(Herter) Rolleri Readily by deep red
identified
Deferrari; Lycopo its
dium hig
Another
luetzelhurgii
leaves. Minas Bahia?.
Rosenstock) ' Brazil: Gerais,
5).
iKuL ^y distinctive high-altitude species (Fig-
TV,»
he ^t;m..
thick, stiffly erect. stems ,
1
densely
ccoovveerreead
^ ^v..ioc;ij
by
rigid, strongly and
uniformly
recurved,
nioi
upper
hooklike The
belong to
following three species
leaves. Brazil: Bahia. This
narrow
"
endemic
the
is an- tane America,
forests southern Central
in -
-
J ' _ ^ - 1
Number
3
Volume 79, Ollgaard 695
1992 Neotropical Lycopodiaceae
They
from
and southeastern Brazil. differ the schlechtendalii Lycopodium
des, (Nessel) Holub;
by papery barbatum
preceding species their large, to sub- non chamae-
Christ, Kaulf.; L.
coriaceous, lanceolate leaves and relatively narrow peuce Herter)
leaf bases. They appear to bridge the gap between
Plants usually rather short and recurved to pen-
the H. brongniartii group and species in the H.
dulous, with linear, densely crowded, usually ba-
taxifolia group, especially to H. rosenstockiana.
sally twisted, falcately ascending leaves. Florida,
)
In the latter group the juvenile sporophytes of some
West
Indies, Central America, northern South
may
species, e.g., H, taxifolia, be initially erect
America, Ecuador Galapagos
south to with the
and resemble plants of H. brongniartii and H.
and Mato
Islands, Brazil (Acre, Crosso, Ceara).
sellowiana.
The
most widespread and variable species of the
jt croup. Closer study of the material referred to this
/o
lluperzi. a bT rongnia* rtii•• (SIprt•m?g)\ Trr.revi- san ^
: .
may s,how , presence more t,han one
species the oi
A
heterogeneous species with slightly distinct taxon, especially in the northern part of the range,
populations in Ecuador-Colombia and Peru-^Boliv- because there considerable variation in the thick
is
com
la, the northern population often with finely den- ness of stems, leaf length and direction, and
However, of
pactness of the plants. the correlation
fi
Colombia The unknown
to Bolivia. minutely rugulate leaf the variation with growth conditions at
is
margins, characteristic of this species, occur also this point (Lellinger, 1989: fig. 23;011gaard, 1988:
19D).
ifolia
fig.
(011
&
Cham.) Holul
Huperzia pithyoides
(Schldl.
Huperzia
foliacea (Maxon) Holub
{Lycopodium gigas Herter)
Smaller
in all parts than the preceding species.
Huperzia
One most impressive species
of the
with
oblong-lanceolate to oblanceolate leaves. Costa
^.^^ ^^^ ^^p^^^ ^f ^ g^^,, ^-^^ t^ee, with thick,
Kica, Panama. A
narrowly endemic, epiphytic spe-
prominent,
^^^^^ j^^ to the color of the
^.^j
^^^-^^^^
cies, probably
closely related to the preceding spe-
and spreading, coriaceous,
Recurrent leaf bases,
cies. The
presence of red dots on the leaf bases cm
up 3.5 long,
bisulcate leaves, often to
j^^^^.^
may
be
evidence
of relationship to species of the q^
may become pendulous, with
fal-
individuals
(Lellinger, 1989: 24;
.^?"P fig. Mexico Costa Rica,
io,T ^^^^j^ ^g^ending leaves. to
ulJll
Maxon,
1912:
t. 1). Colombia: Huila, Venezuela: Fal-
Greater
Antilles,
&
Mickel
1989: 34;
c6n Lellinger, fig.
„ coil v(rFiitgi.. ^2,; i-.<=iiiii5<^i X
L7
,
tiuperzia
sellowiana
(Herter) B. {H. ulei
011g.
jggS: IC, D)
g^ij^j^ fig.
(Herter) Holub; Lycopodium brasilianum
Herter)
{Lyco-
Trevisan
mandiocana (Raddi)
Huperzia
omall
differences of phyllotaxis and a more lax, podium pseudo-mandiocanum Herter)
recumbent
to ascending growth habit separates this
preceding spe-
of the
diminutive replica
Like a
species from
H.
brongniartii. Southeastern Brazil-
except
being almost identical,
characters
the
cies,
G
toRi
Minas
and
Bahia
THE for the size. Brazil:
HUPERZIA
DICHCXIDMA GROUP
Argentina
northernmost
Paraguay,
Grande do
Sul,
Mainly
epiphytes of low- and mid-altitude for- (0Ugaard, 1988: IB).
fig.
ests,
several them
of
with soreadinff
initiallv erect,
to ^
perpendicular, some &
filiform or linear leaves,
^^.^^n Underw. F. Lloyd) B.
•„ (L.
«n
conspicuous
red coloration of the leaf bases, Lyco-
Holub;
Undaviana (Herter)
^^
^jj^
«specially
Huperzia mandiocana
pithyoides, H.
non Rosenstock; L.
podium andinum Herter,
and
H.
wihonii.
Maxon;
stamineum
arcanum Maxon; L.
L.
Th
Herter)
trichodendron
0U
preceding
gaard an epiphyte, but like the
Usually erect
(1987), but excluding H. homocarpa, which
pendulous when very large; often
becoming
belong species,
in the H. taxifolia
prominent leaf
not
usually
but
with bright red,
Cen-
above.
canaliculate
Hiuperzia th« ranillary leaves
dichotoma gra- K=.e..«-
Trevisan
(Jacq.) (//.
Andean South
wwea West
America, Indies,
m
(Spring) Trevisan; H. lindeneri (NesscI) Iral „
•, Mato
Guayana. .
Brazil:
Venezuelan
Peru.
Holub; H. south to
Holub;
mortonii (Herter)
//.
Annals the
696 of
Garden
Missouri Botanical
&
Grosso (Lellinger, 1989: fig. 44; Mickel Beitel, ranges of Venezuela it usually remains short, com
1988: 4B; 01Igaard, 1988: IC). (0U
fig. fig.
1988: 14A).
gaard,
fig.
(
Huperzia polycarpos (Kunze) {Urosta-
B. 01Ig.
Huperzia picardae (Krug) Holub
chys cuatrecasasii Herter)
form
Like the preceding species, but usually pendu- Perhaps representing a slender of the pre-
lous, and smaller in all parts, with basally twisted, ceding species, to which it corresponds rather close-
Panama, Greater Seems connected to robust
falcately ascending leaves. Costa Rica, ly. Antilles.
Colombia Peru 1989:
to (Lellinger, 35; 011-
fig. ifc
gaard, 1988: 19A). forms. Status uncertain.
fig.
&¥.Uoyo)
Huperzia bradeorum Holub Huperzia cuernavacensis Undev'W.
(Christ) brau- (L.
(77.
&
seana Holub
(Herter) RoUeri Deferrari)
A from Hu-
Erect epiphytes with patent, linear-lanceolate very robust species, distinguished
homophyllous
leaves with a slightly narrowed, petiolelike lamina perzia hartwegiana by the thick,
due
base, and usually prominently decurrent leaf bases. shoots, often exhibiting regular constrictions
and
Guatemala to Costa Rica, Venezuela: Isla Marga- to seasonally induced variation of leaf length,
whorls
rita, Ecuador. Probably related to the pendulous linear-lanceolate leaves borne in alternating
&
Guatemala (Mickel
5-6. Mexico
of Southern to
ifi
intermediate between H. bradeorum and H. 1988: 3C).
Beitel,
lini- fig.
1989:
folia (Lellinger, 16; 011
fig.
19B). Huperzia Holub
sotae (Rolleri)
both
Resembling preceding species closely,
the
THE HUPERZIA GROUP
TAXIFOLIA
and
shape ar-
with respect stem thickness, leaf
to
constric-
Pendulous or recurved, slender to very robust rangement, and the presence of regular
more
sol -
epiphytes or rupestral plants, with whorled, but a more delicate plant, with
as- tions,
The
Argentina.
cending appressed, Northernmost
to linear-lanceolate to lanceo- textured leaves.
Morton (t}pe
more
late, rarely linear to subulate, often subcoria- names Lycopodium buesii (Herter) C.
RoUen
ceous to coriaceous leaves. Some species entirely from Peru) and Huperzia aristei (Nessel)
m
&
Colombia,
homophyllous, from
several gradually heterophyllous, Deferrari (type indicated as
or
Argentina
showing at least potentially strong reduction and perhaps northern
possibly mislabeled,
H. sotae,
of
modification of the leaves of distal divisions. Neo- Peru), appear represent close allies
to
known
10-14
They only
tropics, species lower upper montane are
in to or perhaps are conspecific.
i^'
1970:
forests. from fig. 1
the type collections (Rolleri,
Under
certain conditions abruptly dimorphic
C), 2, 3(A)).
t, t.
may
The
leaves be induced.
modification usually
depends on age and
light conditions, juvenile and
ifi
Holub;
shaded plants often being entirely homophyllous. Holub; ff.cu6afia (Herter)
. o r-j -' sis (Herter)
,
passerinoide^^
Leaf texture is variable according to exposure of H. haitensis{UeTtQv)Uo\nh; H,
the habitat within most of the subtubulosa (^^^^^^!
species. (Kunth) TrevLsan; H.
"
Lycopodium herminieri Spring; i>-
Holub;
Her-
Huperzia hartwegiana & schtoendeneri
(Spring) Trevisan Cham.;
ca- tens Schldl. L.
(//.
rubig'-
U.
Nessel;
Urostachys hruelkei
ter;
fi-
Herter; Urostachys maxonii Herter)
Nessel) nosus spongiosus
Nessel; U.
in
The arranged
most
robust species of the group, usually The typical form with the leaves
imal
with strongly coriaceous,
nitid leaves
in alternating
whorls of Northern
four. Central America, Andes subcoriaceous,
sions with ascending,
'^"^^^'"T
sho
from Venezuela with •
northern
to Peru. Apparently divisions,
leaves, in the distal, fertUe
adapted to a relatively drier range of habitats than appressed, leaves. Central
apically involute
—
— ~ ^ o
the other species of the group,
O including d«Arjy rocky Wweesstt *
r» "^ '»-.**A^ i.^-^r^y iilUICd, IlUIlllCIli ^iJUi-*.* *-
tr *»^ci, -I
wi
most
habitats. In Ecuador and Colombia
it varies from to Peru and BrazU; Parana. This is the
g^^
the
mitia spread and species in
variable of the •
and recurved in lava rocks. In the northern and Qoser
coastal problematic.
delimitation
its
is
697
Ollgaard
Number 3
Volume 79,
Lycopodiaceae
Neotropical
1992
para-
more than one taxon timberline, and often on solitary trees in the
of
recognition
may lead to the
may
The be in similar habitats
Some approach H. mo. closest relatives
forms
lini-
the species.
within
New 1988: 17C).
Guinea (0Ugaard,
stems and fig.
homocarpa, with thin flac- in
and H.
folia
and
homophyllous shoots relatively
hanging,
cidly
Holub (Lyco-
(Herter)
Huperzia rosenstockiana
approach H. hartwegia-
Others
whorls
distant leaf
.
Morton)
podium cassandrae (Herter) C.
and crowded
stems,
erect
na, with thick, initially
A
Costa
large
whorls of coriaceous leaves. some
4-leaved but
pendulous epiphyte,
Usually a flaccidly
Rican form has very large sporangia and spores. ^
Ecuador erect.
p^^^j^^s southern terrestrial,
poorly
At present the basis for the variation is Leaves a deep translucent
^.^j^ nodding shoot
tips.
&
Mickel
1989: 39;
understood (LeUinger, fig. yeDowish green, broadly lanceolate with
g^^^ald
to
1983: 8b;
4
Beitel, 1988: fig. A; 011; fig. margins, a feature shared with
minutely rugulose
0Ugaard, 1988: 15A). may
be
fig. which related.
Huperzia brongniartii, to
it
Ecuador. In sheltered sites in
Southern Colombia,
1988:
Huperzia tubulosa (Maxon) comb. nov. (011gaard,
B. 011g., verv humid forest at the timberline
Basionym: Lycopodium tuhulosum Maxon,
^^
14D).
10
Contr. U. Herb. 17 178,
S. Natl.
(2): t.
(1913)
Trevisan
Huperzia funiformis (Spring)
much
Typical plants smaller than the preceding ^ pendulous epiphyte with a distinctive,
large,
species in all parts, and with more distant leaves. due to the very regularly ap-
aspect,
j-opelike
Costa Rica, Panama. This species difficult to convex, subulate
is strongly
coriaceous,
pressed,
W
delimit from the Huperzia taxifolia, due to the
presence of intermediate forms (Lellinger, 1989: has no apparent close rel-
Venezuela Peru.
to It
Maxon,
fig- 41; 1913: 10. 1913). recognition as an in-
t. g^d perhaps deserves
^^-^gg
1989: 25; 011-
dependent group (Lellinger, fig.
Huperzia 1988: 17D).
0Ug. gaard,
killipii (Herter) B. fig.
A
delicate species resembling the preceding spe-
GROUP
HETEROCARPON
THE HUPERZIA
cies, but often completely homophyllous, with
el-
but differmg
group,
preceding
10-mm-long Resembling the
liptic to oblanceolate, usually up to
and
bases,
proximal claspmg and usually twisted leaf
leaves. Venezuela, Colombia. the
in
most of
reduction in
leaf
the strong potential distal
inly
Huperzia
homocarpa Holub
(Herter)
occurring
rupestral plants,
epiphytes or
pendulous
and southern
Homophyllous, lax, pendulous, with rather dis- montane of the eastern
forests
j„
tant
whorls
of relatively short, linear-lanceolate, country,
^f the
p^^jg
usually
basally twisted leaves. Greater Antilles, Cos-
ta Rica, Panama,
Northern Andes, total distribu- Holub longear-
Huperzia heterocarpon (Fee) (//.
tion uncertain. Difficult to deHmit in relation to H. Urostachys portoanus
Holub;
istata (Christ)
taxifolia, H. and H. dichotoma (011-
linifolia, spegazzinii Nessel)
Nessel; U.
gaard, 1988:
16
fig. A).
sporophylls usu-
the
Leaves canaliculate,
long,
and
length
mainly leaf
in
variable
aUy cuspidate,
Huperzia
lancifolia (Maxon) Holub Minas Gerais
reduction. Brazil:
degree of distal leaf
Argentina:
Grande do Sul,
Rio
Resembling more and Rio de Janeiro to
H. acute,
tubulosa, but with
common and widespread of
The most
elliptic proximal Panama, known Missiones.
with cer-
leaves.
tainty only from the type 1989: 29). the species (Fig. 7).
(Lellinger,
fig.
&
Wind.
(SUveira) B. 011g.
Huperzia loefgreniana
Huperzia
sodiroana
lindenii (Spring) Trevisan {H.
Holub)
(Nessel)
{Huperzia
leitzii
(Herter) Holub;
Holub; H. (Herter)
stuehelii
W.
wohlberedtii Holub) assemblage of plants,
(Nessel) heterogenous
A somewhat
intermediate to.
species,
A preceding
the
related to
distinct, flaccidly pendulous species, variable
mbers
^ith
respect dimorphism.
to the degree of leaf
Juvemie
and
plants sometimes erect re- rift
initially
leaves
and short
shoots,
distal
the s.juarish
Curved.
leaves in
Southern the
Colombia, Ecuador, Forest at
Annals the
of
698
Garden
Missouri Botanical
I
*
t
Figures
Huperzia
6 7 martii. -a. Habit.-b.
6. division.
Detail of sporangiate division.-c. DetaU of proximal
Huperzta
heterocarpon.-a.
Hab,t.-b.
7.
Detail of sporangiate
division.-c. proximal division.
Detail of
the proximal
in divisions of the stems. & Wind.
Minas
Brazil: Huperzia
sllveirae (Nessel) B. 011g-
Gerais and Rio de
Janeiro Santa
to
Catarina.
With
Huperzia
martii (Wawra)
Holub
(Lycopodium
ifc
]
ci-
poense
Damazio) Rio
Distinguished by
the densely
crowded,
rather
™^
short and narrow proximal HUPERZIA LimFOLlA GROUP
leaves and
the very
strongly reduced
leaves
ot
.he dU.al, fer.Ue ed by
divi-
characterize
Flaccidly hanging
epiphytes,
^"-
'""°-
"'""' stei.UV"i.h
-7 '^^^- alten..^;
.eaves
ITa^e!;";.-
'r"°
^
whoriea
at least in the proximal not
divisions,
