Sparganium
identification key and species descriptions
By Josh Sulman
University of Wisconsin-Madison
Department of Botany
Updated May, 2013
Identifying
bur-reeds in the field presents a special challenge to botanists, wetland
scientists, and anyone interested in aquatic plants. The group has its own set of
specialized terms. Often, at least until mid-summer, it is necessary to be
familiar with vegetative characteristics of the different species. Most fruits
are not ripe until late summer, so understanding the features of the flowers
and inflorescences is important for spring or summer field surveys.
Additionally, there is remarkable phenotypic plasticity within species. Plants
of some species are typically limp and floating, but erect and emergent during
periods of low water levels. Also, typically erect and emergent species are
occasionally found limp and floating in deep water or flood conditions.
Resources
The treatment of Sparganium by Robert Kaul in the Flora of North America (1997) is a thorough and reliable guide to the genus.
The worldwide monograph (Cook and Nicholls, 1986, 1987) is the most comprehensive taxonomic treatment, and is now available online:
Part 1: Subgenus Xanthosparganium
A key is presented below to the 8 species found in Wisconsin. Detailed descriptions of the plants will follow.
General description and ID tips:
The flowers are borne in globose heads, the female flowers borne in separate heads from the male flowers. The male heads are borne at the tips of the inflorescence axis and branches, and the whole male head is deciduous. After it has fallen, you can tell where it was attached by the scar that is left on the stem.
The female flowers are packed into spiky heads of their own. Each female flower will bear one or two stigmas, depending on the species. The stigma remains prominent in fruit, developing into a sharp-pointed beak. The beak may be straight, curved or hooked and may function in dispersing the fruits. The bur refers to the spiky appearance of the fruit heads.
AXILLARY vs. SUPRA-AXILLARY HEADS: POSITION
OF HEAD RELATIVE TO SUBTENDING LEAF
How the heads are borne in relation to the leaf tends to be one of the most reliable characters for ID, but is also a source of confusion! The terms axillary and supra-axillary are applied to the position of the female heads of Sparganium:
If the head is
borne in the axil of the leaf below it, it is termed axillary.
If the head is
attached to the stem somewhat above
the leaf, it is termed supra-axillary.
A supra-axillary head may be sessile or stalked. Such a stalk (or pedicel) is NOT considered a branch. A branch must bear at least two heads (male or female), and always terminates in a male (staminate) head.
In some individuals, the subtending leaf may be absent. This is often the case in branched inflorescences, on the upper female heads, as in Fig. 1. Also, male heads usually are not subtended by a leaf.
Even within a
single plant, some heads may be axillary while others are supra-axillary: in
this case, it should be keyed as supra-axillary.
|
Fig. 2: Sparganium angustifolium from Vilas County, WI July 2008. Lowest
head axillary and stalked; next head supra-axillary and stalked, 3rd
head slightly supra-axillary and sessile; scars of missing male heads remain
on upper stub of stem. |
2
KEY TO SPECIES OF SPARGANIUM OF WISCONSIN
A. stigmas 2, mature fruits
flat-topped or slightly domed.
Plants common in eutrophic habitats, throughout Wis.; up to 6 ft. tall. . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1.Sparganium
eurycarpum
AA. stigmas 1, fruits
tapered acutely to a pointed beak. Plants in meso- to oligotrophic habitats, mostly central and
northern, occasionally SE Wis.; under 3 ft. tall . . . . . B.
B. plants limp and floating . . . . . C.
C. Plants mostly under 30 cm,
often completely submerged; female heads under 1cm across.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . 2. Sparganium natans
CC. Plants longer, with long leaves
floating on the surface, female heads over 1 cm across. . . . H.
H. Leaves white
beneath, inflorescence branched. . . . . . . . . . . 3.Sparganium fluctuans
HH. Leaves green beneath, inflorescence not branched,
though heads often stalked . . . . . . . . . . . . . . . . . . . . . . . . . .
. . . . . . . . . . . . . . . . . . . . . . . . . . 4. Sparganium angustifolium
BB. Plants erect and emergent . . . . . D.
D. Leaves narrow, usually <8mm across,
some heads supra-axillary, in an unbranched inflorescence; forming loose colonies. . F.
F. heads clustered at top of stem; NW Wis.,
NE Minn, rare . . . . . . . . . 5.Sparganium glomeratum
FF. heads spaced along stem, common,
widespread. . . . . . . . . . . . . . . 6.Sparganium emersum
DD. Leaves up to 2cm wide, heads axillary in a simple or branched
inflorescence; forming dense colonies . . . . . G.
G. mature fruits glossy, with beak 5-6mm. . . . .
. . . . . . . . . . . . . . . . . . 7.Sparganium androcladum
GG. mature fruits dull, with beak 2-5mm. .
. . . . . . . . . . . . . . . . . . . . . . 8.Sparganium americanum
Species descriptions
The
following are based on field work over two summers in Wisconsin and neighboring
regions of the U.P. of Michigan, and northern Minnesota. Associates lists were
compiled from a matrix of 134 species observed at 75 Sparganium habitat localities: associates for each species below
are listed in descending order of frequency, to a minimum of two observed
localities (Sulman, 2010).
1. Sparganium eurycarpum Engelm.
Leaves
cat-tail-like, up to 6 ft. tall, but keeled abaxially
along the middle, trigonous toward base. Leaves usually dark green, turning brown from the tips in late
summer. Female heads axillary or without subtending leaf; main axis and
lateral branches bear numerous male heads. Female flowers and immature fruits
conical, tapered to sharp beak. Mature fruits truncate above, with abruptly
pointed beak, bearing (1)-2 stigmas. This is the most common bur-reed
throughout the southern two-thirds of the state. This is the only bur-reed to
have 2-stigmas/flower; in vegetative material it can be identified by the
leaves that are flattened, but strongly keeled all the way to the tip. Grows in extensive colonies, in water 0 to 3 ft. deep.
Typical habitats include ponds, marshes, lakeshores, streambanks,
wet spots in sedge meadows, and stagnant to slow-moving water generally.
Flowering may be rare in some colonies, especially in deep water. Found in mesotrophic to hypertrophic conditions. It can be
aggressive in its native habitats, and is considered a nuisance in some
situations, as in waterfowl habitat along the Mississippi R. at Alma,
Wisconsin, where this species has invaded open water visited by tundra swans
during the migration (see http://www.nps.gov/miss/naturescience/tundraswan.htm).
S. eurycarpum can be displaced by
invasive Typha X glauca, as along the Minocqua Thoroughfare in Oneida County, and the
shore of University Bay, on the UW-Madison campus. Here, it is a minor
component of an 8ft. tall Typha stand.
Formerly, S. eurycarpum constituted a
major component of the marsh vegetation of University Bay (Heddle, 1910),
forming a 10ft. wide “Sparganium zone”
or a Sparganium-Typha
latifolia zone.
Frequent
associates in Wisconsin include Sagittaria latifolia, Typha latifolia, Lemna minor, Eleocharis palustris, Schoenoplectus tabernaemontani, Ceratophyllum demersum, Nymphaea, Phalaris arundinacea, Polygonum amphibium, Alisma, Carex lacustris, Elodea canadensis, Nuphar, Sagittaria rigida, Sparganium
americanum, Utricularia spp,
Zizania, Asclepias incarnata, Bidens sp., Bolboschoenus fluviatilis, Leersia oryzoides, Sium suave, Acorus calamus, Calamagrostis canadensis, Calla palustris, Carex comosa, Carex
utriculata, Cicuta bulbifera, Eleocharis sp., Glyceria grandis, Lycopus americanus, Lycopus uniflorus, Polygonum sp., Potamogeton richardsonii, Spirodella polyrrhiza, Wollfia.
2. Spaganium
natans L.
Leaves
flat, limp, often entirely submerged, or floating at the tips; a few small
upper leaves may extend above the surface. Leaves green on both sides, 2—5mm
diameter; often forming dense underwater lawns. Flowering stems leafy,
elongating to the water surface: in deep water, producing 6in. internodes, and
leaves up to 2 ft. long. Grows in clear water, 6in. to 3 ft.
deep, often on a soft, organic substrate. Native to lakes throughout the
glaciated portions of Wisconsin, more frequent northward, though not common.
Last southern Wisconsin collections 150 years old (S.H. Watson, 1850’s, WIS),
but found in Walworth Co., summer 2009, in a marl-bottomed, groundwater-fed
lake. This species may occur in other calcareous lakes in the Kettle Moraine
region, but is short in stature and easily overlooked, especially in vegetative
condition. Typical habitats in northern Wisconsin and Minnesota are marshy to
boggy shallow lakes, in protected bays, inlets, and wild rice beds: best
observed by canoe. The species has a wide range of conductivity tolerance,
varying from 686 μS in the calcareous WI lake to 42μS in an acidic thermal pond in Yellowstone
NP. It appears to thrive in areas of low competition, but may grow amongst much
taller vegetation. This is a circumboreal species, and is at the southern limit
of its range in Wisconsin. Frequent associates include Nuphar, Nymphaea, Utricularia
spp, Calla palustris, Chara, Myriophyllum sibiricum, Potamogeton natans,
Sparganium emersum, Typha latifolia,
Zizania palustris.
3. Sparganium fluctuans (Morong) Robinson
Leaves floating, conspicuously
parallel-veined on lower surface, with fainter cross-partitions, especially
visible in dead tissue.
Leaves white beneath, with layers of non-photosynthetic cells. Leaves flat, ribbonlike; max leaf width per plant 5-7mm. Inflorescence
consistently compound, each branch topped with one or more male heads,
typically several in tight clusters. The female flowers have curved stigmas. Fruits with strongly curved
beaks, the body turning reddish at maturity. Fruits may be uncommon; in
Shearer Lake, Taylor County in 2008, 1% of plants had flowers or fruit. Can be
confused with S. angustifolium; S. fluctuans leaves are truly flat, not
rounded on top as in S. angustifolium;
S. angustifolium leaves are green
beneath, and much narrower. Found in oligotrophic to mesotrophic
lakes in northern Wisconsin, in water 0.5-5ft. deep; often in a zone of
floating aquatics bordered on the shallow side with a zone of emergents. Often rooted in sandy
substrate. Many habitats have dark-stained water and relatively low (10
to 83μS, mean=63) conductivity; generally not found in the clearest,
lowest-conductivity waters. Frequent associates include Nuphar, Brasenia schreberi,
Nymphaea, Carex lacustris, Carex lasiocarpa, Dulichium arundinaceum, Eleocharis palustris, Eriocaulon aquaticum, Glyceria borealis, Potamogeton pectinatus, Potamogeton robbinsii, Potentilla palustris, Sparganium
americanum, Sparganium emersum, Utricularia spp., Zizania
palustris.
4. S. angustifolium Michx.
Leaves
floating, linear, the widest usually 2-4mm across; green beneath, slightly
rounded above. The inflorescences are simple, often with a lower pistillate head on a stalk; one or more pistillate
heads supra-axillary; female flowering heads often held up above surface;
fruits floating at or just below surface. Staminate heads few and clustered at
stem tip. Found in clear, oligotrophic lakes and occasionally, in streams;
frequent in lakes in the sandy regions of northern Wisconsin, south to Portage
County. Usually in sandy substrate, in water depths from 0 to 2ft. Prefers very
low to low (10 to 115μS, mean=40) conductivity water with low vegetation
density. Frequent associates include Isoetes, Lobelia dortmanna, Eriocaulon aquaticum, Carex utriculata, Nymphaea, Potamogeton natans,
Equisetum fluviatile, Littorella
uniflora, Nuphar, Potamogeton richardsonii, Sagittaria cuneata, Sagittaria rigida, Typha latifolia.
5. Sparganium glomeratum (Beurl. ex Laest.)
Neuman
Leaves
keeled or triangular in cross section, floating or emergent; pistillate heads supra-axillary, aggregated into a tight
cluster at top of stem, sometimes with 1 or 2 lower heads on peduncles. Mature
fruits shiny, football-shaped, narrowed to a short
beak. Staminate heads 1—2, adjacent to uppermost pistillate
head, soon deciduous, and then leaving no visible trace. Grows
in ditches, marshes and black ash swamps. Locally common near Superior,
Wis. and Duluth, Minn., where first reported by Olga Lakela
in the 1930’s; collected recently in other parts of northern Minnesota and
northwestern Wisconsin (Walton, 1999). This species is listed as Threatened in
Wisconsin. It was observed in a ditch along an old railroad grade, on peat
growing with Salix lucida,
S. exigua, Typha latifolia, Equisetum fluviatile, Sium suave, Alisma subcordatum, and Bidens sp.
6. Sparganium
emersum Rehmann
Leaves
usually emergent, triangular in cross-section, or flat
and keeled above. Leaves typically much exceed the inflorescence. Height
ranging from 0.5-2ft. Female heads (at least one) supra-axillary, inflorescence unbranched,
lower heads often stalked. The mature fruits have a curved beak, and are
somewhat constricted around the midsection; they tend to be red-dotted on the
lower surface. Male heads usually three or more, spread out along the upper
portion of the stem. This species is easily confused with S. americanum, (especially in flowering or early-fruiting
specimens!) S. emersum can usually be
identified based on the presence of supra-axillary heads, and a smooth, shiny
fruit surface. However, rare individuals have branched inflorescences (see U.
Müller-Doblies, 1969; Voss & Reznicek,
2012), or have all pistillate heads axillary. S. emersum may produce floating leaves
in deep or fast-moving water, where it rarely flowers. These plants can be differentiated from S. angustifolium by the keeled leaf.
Hybrids between S. emersum and S. angustifolium occur in the Rocky
Mts., and may occur in Wisconsin as well. S.
emersum occurs frequently across the northern half of Wisconsin, in shallow
water (up to ≈6in. deep) at lake margins, in bog moats, marshes, in
sluggish streams and ditches, and in beaver ponds. It occurs sporadically in
southern Wisconsin in marshes, lakes, bogs, fens, streams and river bottoms. It
occurs in low and high conductivity water (60 to 650, mean=176μS). Frequent associates include Nuphar variegata, Nymphaea odorata, Zizania palustris, Brasenia schreberi, Sparganium
americanum, Carex lacustris,
Ceratophyllum demersum,
Elodea canadensis, Potamogeton
natans, Sagittaria latifolia,
Calla palustris, Chara, Dulichium arundinaceum, Eleocharis sp., Leersia oryzoides, Lemna minor, Myriophyllum sibiricum, Scirpus cyperinus, Sparganium fluctuans, Sparganium natans, Spirodella polyrrhiza, Utricularia spp.
7. Sparganium androcladum (Engelm.) Morong
Robust,
emergent plants to 3 ft. tall, leaves keeled or triangular in cross section,
inflorescences branched, or simple with axillary heads. The lateral
inflorescence branches usually bear
staminate heads only. The pistillate heads are large
and few; the mature fruits are glossy,
with beaks 4-7mm, straight and often hooked at the tip. This species is often
confused with S. americanum, especially
in flower or immature fruit. Their ranges overlap in central and northwestern
Wisconsin. In Wisconsin this species is sporadic across southern and western
Wisconsin. It is locally abundant in the bed of Glacial Lake Wisconsin, in
Jackson, Monroe, Juneau and Wood counties. Typical
habitats are margins of flowages, in mesotrophic,
moderate to low conductivity water (from 27 to 82, mean=52μS), 0 to 3 feet
deep, on silty, clayey or sandy substrates. It was
observed at four localities in Wisconsin. Frequent associates include Brasenia schreberi, Dulichium arundinaceum, Nymphaea, Sagittaria latifolia, Bidens sp., Eleocharis palustris, Phalaris arundinacea, Scirpus cyperinus.
8. Sparganium americanum Nutt.
Robust,
emergent plants to 3 ft. tall, leaves triangular in cross section below and
keeled toward the tip, light green. In deeper water this species may produce
floating leaves, which are keeled. The inflorescence may be branched or simple,
with axillary female heads (except where no leaf is present). The tepals of the
flowers may or may not have a dark spot. The fruits have a dull, non-glossy surface, beaks 2-5mm long, straight or slightly
curved. Typical habitats are peaty to sandy lakeshores, ditches (in central
WI), beaver ponds, in sluggish streams, and floating vegetation of boggy lakes,
in water 0 to 3 ft. deep. The plants may form large colonies. This species is very
common in northern and central Wisconsin, in oligotrophic to mesotrophic water, from low to moderately high conductivity
(15 to 200, mean=67μS). Frequent associates in Wisconsin include Eleocharis palustris, Nymphaea, Brasenia schreberi, Nuphar, Sagittaria latifolia, Utricularia spp, Typha latifolia, Dulichium arundinaceum, Elodea canadensis, Zizania, Glyceria grandis, Potamogeton epihydrus, Potamogeton natans, Schoenoplectus
tabernaemontani, Scirpus cyperinus, Sparganium emersum, Sparganium eurycarpum, Calamagrostis canadensis, Calla palustris, Carex lacustris, Sagittaria rigida, Acorus calamus, Carex comosa, Glyceria borealis, Glyceria canadensis, Leersia oryzoides, Lemna minor, Potamogeton amplifolius, Potamogeton sp., Puccinellia, Sparganium fluctuans.
References
Cook, CDK and MS Nicholls. 1986. A Monographic Study of the Genus Sparganium. Part 1: Subgenus Xanthosparganium. Bot. Helv. 96:2
Cook, CDK and MS Nicholls. 1987. A Monographic Study of the Genus Sparganium. Part 2: Subgenus Sparganium. Bot. Helv. 97:1
Heddle JR 1910. The plant geography of the University Bay region. Univ. of Wisconsin: B. A. Thesis, 78pp.
Kaul, RB 1972. Adaptive leaf architecture in emergent and floating Sparganium. Amer. J. Bot. 59(3): 270-278.
Kaul, RB 1997. Sparganiaceae. In: Flora of North America Editorial Committee, eds. 1993+. Flora of North America North of Mexico. 12+ vols. New York and Oxford. Vol. 22.
Müller-Doblies U. 1969. Über die Blütenstände und Blüten sowie zur Embryologie von Sparganium. Bot. Jahrb. Syst. 89: 359-450.
Nichols SA 1999. Distribution and habitat descriptions of Wisconsin Lake Plants. Madison: Wisconsin Geological and Natural History Survey Bulletin 96.
Sulman, JD 2010. A systematic study of the genus Sparganium. M. S. Dissertation, University of Wisconsin, Madison, Wisconsin, USA.
Voss EG and AA Reznicek 2012. Field Manual of Michigan Flora. Ann Arbor: University of Michigan Press.
Walton GB 1999. Report for the 1994-1995 status survey for Sparganium glomeratum in Minnesota. Prepared for the Minnesota Department of Natural Resources Natural Heritage Program. Submitted Dec. 29, 1995.