Table Of ContentNEW
THE
CYPERUS SANGUINOLENTUS (CYPERACEAE) TO
AND
SOUTHEASTERN UNITED RELATION TO
STATES, ITS
THE SUPPOSED ENDEMIC CYPERUS LOU1SIANENSIS
Department
-n><t>i)y
Valdosta State University
698-001
ildosta,GA 31
U.S.A.
5,
m
C
and
show sanguinolentu iH 3<onomically ndistinguishable,
studies
C.
1
• <
ir
widesprea weed, sanguinolentus,
n 'iii Tfie C.
>
i l r ^i ^
^^
formerly thought to be restricted to the Eastern Hemisphere, reporte
is
^
^ d Stributl ° n and eCOl ° 9y in stern United States are
" '
'
"rented
RESUMEN
^
'^
comuny
Losestudiosdecampomuestran localmente una mala hierba
i,u- ,,
.
en este de Louisiana y sur de Mississippi que se extiende hasta Alabama y Georgia. Estudios
el el y
son taxonomicamente
morfometricosydeherbanomuestidii
is
^
> , •
|
como un sinonimo de C.sanguinolentus.la mala hierba
indistinguibles, C.louisianensis se trata
y
m
como nueva para Norte Ameiu a h-i uti u» n> istribuciony ecologia en
| ] i
sureste de Estados.
los
el
INTRODUCTION
where
Hemisphere,
Cyperus sanguinolentus Vahl widely distributed the Eastern
in it
is
weed Mingyuan & Dehu Reed 1977;Kuhn
been (Holm 1991; 970;
has cited as a et
al.
known from northeastern Africa, the Middle East, India Sri Lanka, central Asia,
982).
It is
1
and
southeastern China, Taiwan, Japan, Korea, the Phi ippines, donesia, Malaysia,
Asia,
li
Mingyuan &
Holm Kukenthal 935-1 hwi 1965;
Australia (Clarke 894; et 991 936;
al.
1 1 1
;
Dehu 970; Kern 974; Reed 977; Kunn 982; Haines & Lye 98: Wilson 1993) but has
1 1 1 1 1 ;
not been previously reported from the Western Hemisphere.
Cyperus sanguinolenlih highly vaiiable. Kukenthal (1935-1936) segregated
five
is
named
varieties and seven forms, including under the
six typical variety. Kukenthal's
(1935-1936) taxonomy
infraspecific of Csanguinolentus difficult to use, since he pro-
is
vi.ied nan has sevs nor paiallel oescaptions ol the axa. Kern (loop (loaned tour s^Pspe
l
eies, including ihe typical one, for Malaysia, and others Ohwi 1965; Haines & Lye
(e.g.,
have
1977) treated in hi m| taxono-
n ih inftaspecific
i I i I i i I
mies of Kukenthal (1935-1936) and Kern (1974). Further research on this widespread
and variable species throughout range needed more complete understand-
its for a
is
ing of its infraspecific variation; however, such beyond the scope of our study to deter-
is
mine the range,clMn
tat M
in
i ti
i i i I I , i , i
and achene achonn
Its bifid style lenticular with angle adjacent to rachilla clearly
place Csanguinolentus subgenus
into Pycreus. Clarke 894, 908) segregated Pycreus
as
(1
1
genus and
a treated the taxon as Pycreus songuinolentus Nees subgenus Reticulatae
in
section Vestitae. Kukenthal 935-1 936) adopted a broader definition of Cyperus, incor-
(1
ubgonu '>
porating this taxon into eetion oenus.The
a \(/A ./(/ of that floral scales
dium no
of Csanguinolcntt, nooves
n in n II (or sulci) typi-
I
I i I
ml
cal of section Sulco eait pith blood-red scale pigmenta-
floral
it
s i 'i
some a
tion. Although a^ .on n tm o loetghebeur 1986, 1989; Adams
i i I i
1994;Bruhl 1995) fragment Cyperus and segregate Pycreus the rank of genus, we have
at
more
followed the conservative generic taxonomy of Kukenthal (1935-1936) and
& Lye 977;Tucker
983, 987, 994).
1 1 1 1
hum
In 1977, Thieret described a ne\ pecies, /o /on. specimens he col-
C".
two
lected close Tangipahoa
at sites in Parish, Louisiana. Thieret (1977) placed
C.
m
subgc nu
louisianensis nuihni and
in a h lumolentus, pro-
t it
i
,
vided
several contrasting chaiacteristu separating Horn Csanguinolentus
(Table
s
2)
it
m
andadichotomous
K
I"
listinuuishii species subge-
an relate ith a in
in
it fi
i
nus was
Pycreus. Cyperus louisianensis listed by the Department of United States
Interior,
unn,
&
Fish Wildlife Seivic r in in ni 1 in nt d species (Anony-
, i i
mous
1993). Until Bryson and Carter (1994) showed was widespread and weedy
in
it
southern Mississippi th oiuht he narrow endemic
a restricted to
i,
, , , - ^ • t ^
1993,the autho
In first
to prepare a status survey ho iana ^hich provided supp
as th. initial financial
>
study.The ma|.n
for this i- it a ,<o .sea af a Iditional population:
i . i 1 1 i 't .r t, i ,
ao
and
Clouisianensis additional
hat tatus as potenti
ie a
. II r I it
rare plant, and to examine taxonomic relationship with the Old World weec
its
we
songuinolentus. In this report, provide a complete record of our and herbarh
field
unno
investigations intuth lta>unon
ti tin ir a P hi| of ' ea '
I i I i .
vith mguinolentus.
i <
(1935-1936)
Kiikenthal
melanocepholus]
).
eysmannii (Boeck.) K
AND METHODS
MATERIALS
populations south-
studies.— Systematic intensive surveys for louisianensis in
field C.
m made
Louisiana and southern Mississippi were by the authors during the peri-
5-21 September 1993 and 14-1 8 October 1993. Subsequently, the authors have
when
taxon time and circumstances have
arch sporadically for this al-
published account and more recent collections from the type
lowed. Thieret's (1977)
documentation, provided by Nelwyn Gilmore, Louisiana
with additional kindly
locality
Tangipahoa
Natural Heritage Program, were used to relocate the holotype locality in
were which
Attempts paratype unsuccessful,
Louisiana. to re-locate Thieret's locality
Parish,
not surprising since habitat in the vicinity of this site was substantially altered by high-
is
showed
way and commercial construction activities.Observations at the holotype locality
pond and
be abundant along the margin of shallow
the species to locally a artificial
by humans. Searches of potential
nearby ditches the flatwoods, habitat greatly altered
in
habitat began outward from the holotype
locality. Habitat descriptions, estimates of
population size,and voucher specimens were made when populations were found.
Greenhouse
studies.—Transplants and plants of Clouisianensis from seeds
started
were maintained under greenhouse
m litmus in a at the United States De-
ill. it
. r r I
partment
of Agriculture, Agricultural Research Service, Jamte Whitten Delta States Re-
m J
search Center
it Slot n m ih nt by the second au-
l. )\ u
i i i i
,
made
were
thor in order to understand better the history and phenology of
life C.
determine whether
louisianensis, especially to annual
or perennial.
it is
greenhouse
In experiments, seeds of Clouisianensis were sewn on top of 6
in flats a
cm-deep
mixture of a Bosket loam (Mollic Hapludaf) and sphagnum (50%
soil v/v)
silt in
week
the of March, June, September, and December and
first 1994, 1995, 1996. Trays
in
were watered from beneath
prevent seed and
to disturbance. Individual
soil
C.
m
mnsplantmi
(inciiM (idliiHi ml diameter and
into pots
'<< in
i ' t«
v .
watered from beneath.The greenhouse was maintained 30 35° C day and 25 30°
at to to
C 75%
night, at 60 to relative humidity, and without supplemental 994 and
lighting.
In
1
hum m
1995, flowering ami
transplanted from
several
sites
i 1 1 i I ,
mil
unty, inmi
in in n
I ii ^ ii n irn U
I I i ( | l i I I i I i I I i
m
pots and maintained under the same greenhouse corn inns [Tints grown from seeds.
it
I
Data were taken on date of seedling emergence, date of flowering and and
fruiting,
plant longevity.
Herbarium and morphometric
studies.— Specimens and
louisianensis
C.
I
sanguinolentus were borrowed from NY
selected herbaria (BRIT,GH,MO, and US) order
in
sanguinolentus. Quantitative data on the achene characteristics used by Thieret (1977)
m
m
in
W) distinct lorn an, Ik >an itiinoh nlu k fiom se pnt
ii ' ' 1 lln iiiipii
, <
own
and from our collections. Five achenes from each of specimen
3
1
C
and 20 specimens of sanguinolentus were examined. A Bausch &
L
(6X-30X) dissecting microscope with ocular micrometer was used measure
to various
achene dimensions and
(Table Fig. ratios (Table derived from these measure-
3, 1), 3)
ments were
used to analyze differences in the plane shapes of achenes used by Thieret
(Table to distinguish Clouisianensis and C smm,, n muni mloase (Anony-
2)
1.21
-
I 1
mous
was
1996) used to analyze these data statistically and to construct scatter dia-
grams comparing
for variation ianensi ind n!u Representative
in lot
r I
spikelets from selected specimens were photographed using an Olympus™ SZ40
stereozoom dissecting microscope eguipped with Kodak™ DC1 20 zoom cam-
a digital
Our more numerous
era. recent collections were used with
Thieret's (1977) original de-
and
ription type materials to prepare revised description and an
sc a illustration.
RESULTS
and
Distribution ecology of Cyperus louisianensis—
Since 1993, our surveys have
field
9^
^n^
V
-o-
<
...»
frontal view
g Cyperus louisianensis ami
i.
ACHTH
Achene
thick -rv- hr-n;
ACHTH/ACHL
Ratio used b\
Tl
ACHL/ACHW
approxi
Ratio
,,
ACHL/ACHMX
southern and
Mississippi isolated populat
Georgia. Cyperus appears
louisianensis to
I:
Tammany
and
adjacent
sippi,
St. Parish,
often abunc and
iedy, locally ant, restricted to periodically disturbed
specially those maintained by it owing, such as ditches and edges of
artificial
lere appears to be aggressive and freguently forms dense
stands often
it
in
weeds
ve
(Table
4).
Greenhouse observations —In greenhouse experiments, C.louisian plant
from
emerged from mid- and May mid-September each year; plants flowered
late until
September Octo-
August mid-December, with peak flowering early to early
in
late until
emerging the
and subsequently Iniimd mvns/s plants later in
plants j><
ber;
',
> , .
emerging suggesting that
were flower than those earlier, C.
year shorter at initiation
made September
were from
photoperiodic. Most of our field collections
louisianensis
is
peak ob-
December specimen the with fruiting
16 through 9 (see citations lot U.S.A.),
were
served and mature plants collected in mid-October; however, a few fruiting plants
(1993-1994).
and Ma\ mild
observed and/or collected in April foll> in >i '.-.inter If
i
may account the few collections of
photoperiodic, for
indeed louisianensis this it
C. is
from May mid-September by us and by other botanists. All C. louisianensis plants
until
greenhouse from the and those grown from seeds in green-
transferred into the field
showed no evidence of
house experiments died shortly after fruiting, c
sanguinolentus— was by small
with Cyperus Jhlerei 977) limited a
Relationship
(1
two than miles apart.Our
number specimens collected from only sites located less
of
1
more
shows variable
sample from wider geographical area that louisianensis
larger a C. is
than previously thought the diagnostic achene characters used byThieret (1977) to
in
shows some of the spikelet variation ob-
from sanguinolentus. Figure 2
separate
C.
it
C Of numer-
and the
herbarium specimens of sanguinolentus louisianensis.
served C.
in
we examined from throughout the Old World,
have
ous specimens of C.sanguinolentus
Oct 960
ones from Japan M. Furuse 30 Sep 959 (GH); M. Furuse
certain [e.g., s.n„ s.n., 1 1
1 1
NSM from
(GH); Hutoh 1517 (NY); Okamoto 584 (BRIT, NY)] are indistinguishable C.
1
and and achene characteristics used byThieret
general spikelet
louisianensis in in (Fig. 2)
degree
Our and herbarium observations show considerable variation in the
977). field
(1
some
of development of pigmentation and sulcus in floral scales. At least of the variation
photopenod and temperature. ob-
pigmentation appears related to Field
scale
in floral
show mid-September the scales
servations the southeastern United States that in floral
in
pigmentation, and by mid-October they are deeply
are typically pale with only faint
may
vary
pigmented 2A and Opposite sides of an individual spikelet also
Figs. 2B).
(cf.
show
pigmentation 2G and 2H). Scatter diagrams (Figs. 3-6) C.
greatly Figs.
in (cf.
achene
C.sanguinolentus are indistinguishable with regard to the critical
USMON
i)IS(
widely distributed the coastal plain
Cyperus louisianensis in
is <
States and often locally abundant in habitats subject to periodic artificial disturbat
is
and maintenance. Distribution, frequency, and habitat of C.louisianensis are indicative
narrow endemic Furthermore, appears
semi-aggressive weed, not of a species. t
a it
expandin lah
the range of this taxon apparently in in >uth< rn nil
is I
j
and maintenai
from road construction
has possibly resulted part
that dispersal in
its
.
Andropogon
L
virginicus
Axonopusfissifolius Kuhlm.
(FUldi'l
Bacopacaroliniana
(Walt.) Robins.
Boltonia diffusa
Ell.
G'eere
nodiflora
/./pp/o
(L.)
Magnolia
virginiana
L.
Mitreola sessilifolia (GniH.) G. )on
I
Oxypolis
filiformis (W.-iltei)
it
I r 1
Phyllanthus urinaria
L.
Polygonum
hydropiperoides
Polypremum procumbent
1
Rhynchospora
corniculata
(I
Spoi()i)olu^i
Stenotaphru
(yperussanguinolenl iandf.tou«/flnens/5.-A.U.S.A.,16September1993,fflrte/-77i«.-B.
Fig. 2. Spikelet variation in
U.S.A., 18 October 1993, Carter 11 562. -C. U.S./
= mm.
7577.-F. China, 7sang20665.-G&H. Opposite ides of same spikelet, Japan, Okamoto NSM 584. Scale bar
K. 1
7
c louis
.' sang
•
•
.
J
X:V:-:'
•.
»',v
•
;d*»
oa
Hs.
\
' *
° &°
§ >o
„
•
".
.--.
-
.05 1.15 1.25 1.35 1.45 1.55 1.65 1.75 1.85
ACHL/ACHW
regard to critical ratios defining
ACHL/ACHMX
ACHL/ACHMX
,ou
1S
ang
s
+
| [
ACHTH/ACHL
c
ACHL/ACHMX
o th ?f iitk ".Tios defining achene shape; character abbreviations keyed
ir
il
