■ezi

tON BULLETIN 528

BIO SCI
LIBRARY

April, 1985

quatic Vascular Plants of New England:
Part 8. Lentibulariaceae

by
G. E. Crow and C. B. Hellquist

NEW HAMPSHIRE

AGRICULTURAL EXPERIMENT STATION

UNIVERSITY OF NEW HAMPSHIRE

DURHAM, NEW HAMPSHIRE 03824

UmVERSITY OF NEV/ MAMP.SHJM
LIBRARY

ISSN: 0077-8338

>

e.zi

[ON BULLETIN 528

BIO SCI

LIBRARY

April, 1985

quatic Vascular Plants of New England:
Part 8. Lentibulariaceae

by
G. E. Crow and C. B. Hellquist

NEW HAMPSHIRE

AGRICULTURAL EXPERIMENT STATION

UNIVERSITY OF NEW HAMPSHIRE

DURHAM, NEW HAMPSHIRE 03824

UNtVERSITY or NEVv' MAMP.SHI.Ht
LIBRARY

ISSN: 0077-8338

ACKNOWLEDGEMENTS

We wish to thank Drs. Robert K. Godfrey and George B. Rossbach
for their helpful comments on the manuscript. We are also grateful to
the curators of the following herbaria for use of their collections: BRU,
CONN, CUW, GH, NHN, KIRI, MASS, MAINE, NASC, NCBS,
NHA, NEBC, VT, YU. A special thanks is extended to Pamela Bruns
Brayton who prepared the illustrations. Gratitude is expressed to the
University of Wisconsin Press for permission to use or redraw several
figures from Fassett's Manual of Aquatic Plants.

This work is a result of research sponsored by the New Hampshire
Agricultural Experiment Station. The NHAES reserves the right to
reproduce, publish or otherwise use, and to authorize others to use, the
work for Government purposes notwithstanding notice of copyright.

Copyright 1985 by the University of New Hampshire. No part of this work may be
reproduced in any manner without permission from the authors and the University of
New Hampshire.

Programs of the New Hampshire Agricultural Experiment Station are open to all
persons without regard to race, color, national origin or sex. The University of New
Hampshire is an Affirmative Action 'Equal Opportunity Employer.

3 WbODODbia M7m

ABSTRACT

This paper is the eighth in a series of reports on the aquatic and
wetland flora of New England. It treats all species of the Lentibulari-
aceae occurring in New England and includes keys, comments on
taxonomy and nomenclature, habitat and distributional information,
water chemistry data, illustrations and dot maps. Those species
regarded as rare and endangered in the New England Region or in
one or more of the six New England States are also noted.

KEY WORDS: Aquatic Plants, New England Flora, Taxonomy, Lentibulariaceae,
Utricularia. Bladderwort.

i

m

no 52^

TABLE OF CONTENTS

Page

INTRODUCTION 1

LENTIBULARIACEAE 2

Utricularia 2

Key to Species 2

Utricularia fibrosa 4

Utricularia intermedia 4

Utricularia radiata 4

Utricularia purpurea 4

Utricularia minor 10

Utricularia geminiscapa 10

Utricularia vulgaris 10

Utricularia gibba 13

Utricularia biflora 13

Utricularia subulata 13

Utricularia cornuta 13

Utricularia resupinata 17

Literature Cited and Selected References 19

11

Aquatic Vascular Plants of New England:
Part 8. Lentibulariaceae

by
G. E. Crow and C.B. Hellquist

INTRODUCTION

This is the eighth in a series of reports on the aquatic and wetland
flora of New England. These reports are intended to aid conservation-
ists, fish and game personnel, consultants, botanists, and students in
the identification of aquatic plants. The coverage is strictly New
England but is of value throughout the northeast. Data have been
gathered from herbaria in New England and from personal field work.

Chemical data presented represent samples from many waters
throughout New England. The alkalinity readings are total alkalin-
ity, expressed as milligrams per liter (mg/1) CaCOa. The number of
observations is given in parentheses following alkalinity and pH
values. Since pH and alkalinity vary greatly during the day, the
values are only indicative of the water quality.

The rare and endangered plant lists referred to are those prepared
for each of the six New England states by the New England Botanical
Club in cooperation with the United States Fish and Wildlife Service,
Office of Endangered Species, Newton Corner, MA (RI — Church and
Champlin, 1978; MA — Coddington and Field, 1978; VT — Country-
man, 1978; ME — Eastman, 1978; CT - Mehrhoff, 1978; NH - Storks
and Crow, 1978). Taxa indicated as rare, threatened or endangered for
the entire New England Region are also noted (Crow et ai, 1981,
Rhodora 83: 259-299.)

We invite comments and/or criticisms on this treatment. Infor-
mation on any species omitted or any additional localities will be
welcomed. If anyone is interested in specific localities of any of the
species indicated on the dot maps, please contact us.

Dr. Garrett E. Crow, Department of Botany and Plant Pathology, Nesmith Hall,
University of New Hampshire, Durham, NH 03824.

Dr. C. Barre Hellquist, Department of Biology, North Adams State College. North
Adams, MA 01247.

LENTIBULARIACEAE

Utricularia (Bladderwort)

Perennial aquatic herbs; plants consisting of stem systems, roots
absent; lateral leaf-like branches green, usually bearing bladder-like
carnivorous traps, bladders sometimes borne oh separate, non-
photosynthetic branches; flowers borne on scapes elevated above
water surface, solitary or in racemes; fruit a 2-valved capsule; peren-
nating by winter buds (turions).

Key to Species

1. Plants floating in the water or creeping over surface of substrate
(sometimes appearing anchored).

2. Plants with two distinct types of branches, green leaf-like
branches lacking bladders, and bladder-bearing branches lack-
ing chlorophyll, sometimes appearing anchored in substrate
(figs. 1A,2A).

3. Segments of leaf-like branches fine, becoming narrower with

each dichotomy, ultimate segments thread-like, not toothed

(fig. IB); corolla pale-yellow to greenish-yellow, lower and

upper lips nearly equal (fig. lA).

I. U. fibrosa

3. Segments of leaf-like branches distinctly flat, segments
remaining approximately the same width, divisions minutely
toothed (fig. 2B); corolla bright yellow, lower lip nearly twice
as long as upper lip (fig. 2A).

2. U. intermedia

2. Plants with branches all alike, leaf-like, bearing bladders.

4. Flowering scapes subtended by a whorl of inflated branches,
floats holding the flowers above the water surface (fig. 3A).

3. U. radiata

4. Flowering scapes not subtended by inflated branches (floats).
5. Leaf-like branches all whorled (fig. 4F); bladders borne at
tips of branches (fig. 4F); flowers purple (rarely albino).

A. U. purpurea

5. Leaf-like branches chiefly alternate; bladders borne scat-
tered on branches; flowers yellow.

6. Leaf-like segmens flat; spur of corolla very short, sac-
cate, almost absent.

^. U. minor

6. Leaf-like segments terete; spur of corolla.3 mm or more
long.

7. Leaf-like branches divided 6 or more times; free-
floating; plants large, stout, 30-100 cm long.

8. Scapes with 2-5 flowers; bracts subtending flowers
unlobed; submersed, bladder-bearing branches often
with small, apetalous cleistogamous flowers (fig.
4D); branches finer, segments entire, except for
small spicules at tips (magnification needed).
6. U. geminiscapa

8. Scapes with 6-12 flowers; bracts subtending flowers
with basal lobes; cleistogamous flowers absent;
branches coarser, segments with spicules along
margins (fig. 4B) (magnification needed).
7. U. vulgaris

7. Leaf-like branches divided 1-3 times; creeping over wet
mud or in shallow water, or in tangled masses floating
in shallow water; plants small, slender, less than
30 cm long.

9. Flower spur much shorter than lower lip of corolla,
thick, obtuse, 3.0-3.5 mm long (fig. IF); lower lip of
corolla 5-6 mm long; leaf-like branches with mostly
2 segments.
8.U. gibba

9. Flower spur somewhat shorter to slightly longer
than lower lip of corolla, narrow, conic, usually
curved, more than 4 mm long; lower lip of corolla
8-10(-15) mm long; leaf-like branches with 3 or more
segments (usually forked twice).

9. U. biflora

1. Plants anchored in substrate, apparent only when flowering,
appearing terrestrial (vegetative grass-like branches sometimes
emergent, or sometimes floating in densely tangled mats due to
wave action (fig. 51).

10. Flowers yellow (rarely purplish or white); bracts along scape
alternate (fig. 5A,D); flowers (l-)2-12.

11. Flowers clearly pedicellate; bract at base of pedicel peltate
(fig. 5B); spur of corolla appressed to lower lip (fig. 5C).

10. U. subulata

11. Flowers subsessile; bract at base of flower attached basally
(fig. 5E); spur of corolla divergent (fig. 5D).

11. U. cornuta

10. Flowers purple; bracts along scape paired, fused (fig. 5F,G);
flowers 1.
12. U. resupinata

1. Utricularia fibrosa Walt. Fig. 1, Map 1

Rare in pools, wet peat, and long sandy shores of southeastern
Massachusetts. This species is easily confused with U. biflora, but is
more robust and has two distinct branch types, one with bladders and
one without. Range extends from southeastern Massachusetts and
central Connecticut south along the Coastal Plain to Florida and
along the Gulf Coast to eastern Oklahoma and eastern Texas;
California.

Rare and endangered plant lists: New England, Massachusetts,
Connecticut

2. Utricularia intermedia Hayne Fig. 2, Map 2

Common in shallow water, peaty soils, and damp sands through-
out New England. Although more abundant in acidic waters this
species is often found in moderately alkaline waters. The green, leaf-
like branches above the substrate lack bladders, hence the plant may
not be readily recognized as a bladderwort until the plant is removed
from the substrate and the bladders observed. Range extends from
Greenland, Newfoundland, and Nova Scotia west to Alaska, south to
New England, Long Island, New York, northern Delaware, Pennsyl-
vania, Ohio, northern Indiana, northern Illinois, northwestern Iowa,
and California.

alkalinity: mean 9.7 mg/1; range 3.5-52.5 mg/1 (17)

pH: mean 6.7; range 5.9-7.9 (17)

3. Utricularia radiata Small Fig. 3, Map 3

Common in acidic waters of southern New England, widely scat-
tered in northeastern New England. This taxon is treated by some
authors as Utricularia inflata Walt. var. minor Chapm., but Reinert
and Godfrey (1962) regard it as a distinct species from the more robust
Utricularia inflata Walt, of the southeast. This taxon is easily identi-
fied in the flowering state, but sterile, submersed material is
occasionally difficult to identify. Range extends from Nova Scotia
and New England south along the Coastal Plain to southern Florida
and westward along the Gulf Coast to eastern Texas; westward
inland to Pennsylvania, northwestern Indiana, West Virginia, and
Tennessee.
Rare and endangered plant list: Vermont

alkalinity: mean 7.8 mg/1; range 1.0-33.0 mg/1 (26)

pH: mean 6.6; range 5.6-7.3 (27)

4. Utricularia purpurea Walt. Fig. 4, Map 4

Common, free-floating in acidic waters of eastern and southern
New England, rarely in moderately alkaline waters. This species
often flowers sporadically in many populations while flowering regu-

Figure 1
Utricularia fibrosa: A. habit x 1. B. leaf-like branch x 4.
C. spur of corolla x 2, redrawn from Fassett.
Utricularia biflora: D. habit ^ 1. E. spur of corolla x 2,
redrawn from Fassett.
Utricularia gibba: F. habit x 1.

Fi^re 2

Utricularia intermedia: A. habit x 1. B. portion of leaf-
like branch lacking bladders x 8. C. bract, side and
back views, " 4. D. winter bud x 1. All from Fassett.
Utricularia minor: E. habit x 1. F. portion of leaf-like
branch bearing bladder x 10. G. winter bud x 1, from
Fassett.

8

Figure 3
Utricularia radiata: A. habit >* 1 , redrawn from Fassett.

9

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larly and abundantly in others. The flower color ranges from purple to
deep pink and rarely white (forma alba Hellquist). Range extends
from Newfoundland, and Nova Scotia, west to southern Ontario,
northern Indiana, Michigan, and Wisconsin, south along the Coastal
Plain to Florida, Louisiana, and Texas.
Rare and endangered plant list: Vermont

alkalinity: mean 7.8 mg/1; range 2.5-33.0 mg/1 (39)

ph: mean 6.6; range 5.6-7.3 (36)

5. Utricularia minor L. Fig. 2, Map 5

Widely scattered in acidic, alkaline, and slightly saline waters,
wet meadows, bogs, and shores throughout New England. This spe-
cies is rarely found in flower, but is easily identified by its flattened
stems. Range extends from Greenland, Labrador, and Newfoundland
west to British Columbia, south to Nova Scotia, New England, Long
Island, New York, northern New Jersey, Pennsylvania, northern
Ohio, northern Indiana, northern Illinois, northwestern Iowa,
Nebraska, Colorado, and California.
Rare and endangered plant list: Vermont

alkalinity: mean 9.6 mg/1; range 3.0-33.5 mg/1 (7)

pH: mean 6.9; range 6.0-8.1 (7)a

6. Utricularia geminiscapa Benj. Fig. 4, Map 6

Widely scattered in bog waters, acidic or moderately alkaline
waters throughout New England. This species may be mistaken for a
diminutive Utricularia vulgaris, thus flowering specimens are needed
for accurate identification. Range extends from Newfoundland and
New Brunswick west to Michigan and Wisconsin, south to New En-
gland, New York, eastern Pennsylvania, Delaware, and eastern
Virginia.
Rare and endangered plant list: Vermont

alkalinity: mean 24.6 mg/1; range 5.4-69.5 mg/1 (5)

pH: mean 6.7; range 3.5-8.6 (5)

7. Utricularia vulgaris L. Fig. 4, Map 7

Common and abundant throughout New England in acidic and
alkaline waters. This is the most robust and common bladderwort in
New England. Some stems range up to one meter in length. Range
extends from southern Labrador west to Alaska, south to Virginia,
North Carolina, Ohio, Indiana, Missouri, Oklahoma, and northern
Texas; California and Mexico.

alkalinity: mean 18.6 mg/1; range 2.5-109.0 njg/1 (75)

pH: mean 6.7; range 5.7-8.6 (70)

Figure 4
Utricularia vulgaris: A. habit x 1. B portion of leaf-
like branch bearing spicules x 6. C. winter bud ^ 1,
from Fassett.

Utricularia geminiscapa: D. habit (cleistogamous
flower at arrow) x 1. E. winter bud x 1, from Fassett.
Utricularia purpurea: F. habit x 1. G. winter bud x 1.

12

13

8. Utricularia gibha L. Fig. 1, Map 8

Common in shallow water, bogs, and along sandy shores of
southern New England, widely scattered in northern New England.
In New England this species is much more abundant in acidic sites,
while westward it is often found in wet marly sites of higher alkalin-
ity. Range extends from Nova Scotia, southern Quebec, and New
England west to New York, southern Ontario, Michigan, Wisconsin,
and Minnesota, south to Florida, Oklahoma, and Texas; California
and Mexico.

alkalinity: mean 15.6 mg/1; range 3.0-69.5 mg/1 (16)

pH: mean 6.7; range 5.8-8.6 (14)

9. Utricularia biflora Lam. Fig. 1, Map 9

Rare in shallow pools, wet peat, and along sandy shores of south-
ern New England. This species is easily confused with the more
common U. gibba and the rarer U. fibrosa, and is somewhat interme-
diate between them. Range extends from the Coastal Plain of eastern
Massachusetts, southern Rhode Island, and southern Connecticut
south to Florida and west along the Gulf Coast to eastern Oklahoma
and southeastern Texas.

Rare and endangered plant list: New England, Massachusetts,
Connecticut

10. Utricularia subulata L. Fig. 5, Map 10

Rare in wet peats and sands in southeastern New England. Pres-
ently this species is known in New England from only two extant
sites, one in southeastern Massachusetts and one on Nantucket
Island. Range extends along the Coastal Plain from southwestern
Nova Scotia, southeastern Massachusetts, Rhode Island, and Long
Island, New York, to Florida and along the Gulf Coast to Texas;
inland to Tennessee, Arkansas, and eastern Oklahoma.
Rare and endangered plant list: New England

11. Utricularia cornuta Michx. Fig. 5, Map 11

Common in sphagnum bogs, wet sands, and along sandy or
peaty shores in New England. More abundant in acidic regions of
New England, but also found in the alkaline regions of northeastern
Maine and northern Vermont. This species is inconspicuous, except
when in flower. The bladder bearing, subterranean vegetative mate-
rial may be found floating in densely tangled mats after being washed
free by wave action. Range extends from Newfoundland and Quebec
west to northern Ontario, Wisconsin, and Minnesota, south to Dela-

Figure 5

Utricularia subulata: A. habit showing subterranean
bladder-bearing branches and vegetative grass-like
branches x 1. B. peltate bract x 8. C. corolla showing
appressed spur x 2.

Utricularia cornuta: D. habit showing subterranean
bladder-bearing branches and vegetative grass-like
branches * i. E. bract x 4. Both from Fassett.
Utricularia resupinata: F. habit showing subterranean
bladder-bearing branches and vegetative grass-like
branches x i. G. paired bracts x 4, from Fassett.
H. corolla showing divergent spur x 2, from Fassett.
I. mat of floating, densely tangled branches.

15

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ware, Pennsylvania, Ohio, northern Indiana, and northern IlHnois;
along the Coastal Plain to Florida and eastern Texas.

alkalinity: mean 3.7 mg/1; range 1.0-7.0 mg/1 (6)

pH: mean 6.4; range 5.6-6.9 (6)

12. Utricularia resupinata B. D. Greene Fig. 5, Map 12

Uncommon in acidic regions of southern and central New En-
gland. This species, like U. subulata and U. cornuta, is difficult to
identify in the sterile state and is usually overlooked. Many of the
historical sites have been revisited to search for this species, but to no
avail. This species seems to flower more often when the water recedes
during dry seasons. Range extends from Nova Scotia west to Quebec,
northern Michigan, and northwestern Wisconsin south to New York,
northern Pennsylvania, northern Indiana, and northeastern Illinois;
along the Coastal Plain to Georgia, Florida, and Alabama.
Rare and endangered plant list: Vermont

18

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19

Literature Cited and
Selected References

Hellquist, C. B. 1974. A white-flowered form of Utricularia purpurea
from New Hampshire. Rhodora 76: 805.

Lloyd, F. E. 1933. The structure and behavior of Utricularia purpurea.
Canad J. Res. 8: 234-252.

Lloyd, F. E. 1935. The traps of Utricularia. Proc. 6th Bot. Congr. 1: 54-73.

Lloyd, F. E. 1935a. Utricularia. Biol. Rev. 10: 72-100.

Reinert, G. W. and R. K. Godfrey. 1962. Reappraisal of Utricularia
inflata and U. radiata (Lentibulariaceae). Amer. J. Bot. 49: 213-220.

Rossbach, G. B. 1939. Aquatic Utricularias. Rhodora 41: 114-128.

Uttal, L. J. 1960. Notes on Utricularia hiflora and U. fibrosa.
Rhodora 58: 41.

22
Station Bulletins of Botanical Interest

Grasses of New Hampshire. I. Tribes Poeae (Festuceae) and Triticeae
(Hordeae). A. R. Hodgdon, G. E. Crow, and F. L. Steele. Bull. No.
512. 1979.

The Flora of Plum Island, Essex County, Massachusetts. M. J.
McDonnell. Bull. No. 513. 1979.

Aquatic Vascular Plants of New England: Part 1. Zosteraceae,
Potamogetonaceae, Zannichelliaceae, Najadaceae. C. B. Hellquist
and G. E. Crow. Bull. No. 515. 1980.

Aquatic Vascular Plants of New England: Part 2. Typhaceae and
Sparganiaceae. G. E. Crow and C. B. Hellquist. Bull. No. 517 1981.

Aquatic Vascular Plants of New England: Part 3. Alismataceae.
C. B. Hellquist and G. E. Crow. Bull. No. 518. 1981.

Aquatic Vascular Plants of New England: Part 4. Juncaginaceae,
Scheuchzeriaceae, Butomaceae, Hydrocharitaceae. G. E. Crow
and C. B. Hellquist. Bull. No. 520. 1982.

Aquatic Vascular Plants of New England: Part 5. Araceae,
Lemnaceae, Xyridaceae, Eriocaulaceae, and Pontederiaceae.
C. B. Hellquist and G. E. Crow. Bull. No. 523. 1982.

Aquatic Vascular Plants of New England: Part 6. Trapaceae, Halora-
gaceae, Hippuridaceae. G. E. Crow and C. B. Hellquist. Bull. No.
524. 1983.

Aquatic Vascular Plants of New England: Part 7 Cambombaceae,
Nymphaeaceae, Nelumbonaceae, and Ceratophyllaceae. C. B.
Hellquist and G. E. Crow. Bull. No. 527

Hiker Traffic On and Near the Habitat of Bobbins Cinquefoil, an
Endangered Plant Species. R. E. Graber and G. E. Crow. Bull. No.
522. 1982.

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