Aquatic Botany, 34 (1989) 367-373 367

Elsevier Science Publishers B.V., Amsterdam - - Printed in The Netherlands

WATER T E M P E R A T U R E A N D F R E S H W A T E R M A C R O P H Y T E
DISTRIBUTION

EVA PIP
Department of Biology, University of Winnipeg, Winnipeg, Manitoba, R3B 2E9 (Canada)
(Accepted for publication 10 January 1989)

ABSTRACT

Pip, E., 1989. Water temperature and freshwater macrophyte distribution. Aquat. Bot., 34: 367-
373.

Distributions of aquatic macrophytes and community species richness were examined in rela-
tion to water temperature at 345 sites in central North America. No significant interspecific dif-
ferences were found for the monthly temperature ranges of the various macrophytes found within
each of the ponds, lakes and lotic habitats during the growing season in the area studied. However,
some species showed significant differences when all habitat types were pooled. Species richness
was positively correlated with maximum seasonal water temperature in lakes. Temperature in
itself does not appear to be an important factor governing the distribution of macrophytes within
the study area, although it may operate through other site variables such as water body size and
type, and through indirect effects on water chemistry.

INTRODUCTION

Temperature is known to be an important factor in macrophyte growth. It

influences a variety of physiological responses such as dormancy and turion
formation (Haag, 1979; Winston and Gorham, 1979; Sastroutomo, 1980), seed
germination (Teltscherova and Hejn~, 1973), development (Setchell, 1946;
Barko and Smart, 1981; Barko et al., 1982), photosynthetic rate (Titus and
Adams, 1979; Barko and Smart, 1981) and oxygen consumption (Anderson,
1969). A number of workers have noted that growth of some species is more
robust at high temperatures, although the growth cycle is compressed (Ander-
son, 1969; Young, 1974; Grace and Tilly, 1976; Barko and Smart, 1981 ). Shel-
don and Boylen (1977) have pointed out that in many lakes the lower bound-
ary for rooted angiosperm growth coincides with the depth of the maximum
penetration of the summer thermocline. In deeper waters, low temperature
may limit the length of the growing season (Moeller, 1980). Changes in the
temperature regime of a water body have been reported to result in alterations
of macrophyte community composition (Allen and Gorham, 1973). However,

0304-3770/89/$03.50 © 1989 Elsevier Science Publishers B.V.

368

as has been pointed out by Barko and Smart (1981), almost nothing is known
regarding the influence of temperature on the distributions of individual ma-
crophyte species.
The objective of the present study was to determine whether significant in-
terspecific differences exist with respect to water temperature for macrophytes
in central North America, and whether temperature is related to community
species richness within similar types of water bodies.

MATERIALS AND METHODS

Sampling was carried out during the M a y - S e p t e m b e r 1972-1985 seasons

(except 1977 and 1979) at 345 sites located within the area bounded by 47-
54 °N and 94-106 ° W. All sites contained water year-round and represented a
wide spectrum of trophic states. The distribution of macrophytes in the study
area with respect to water chemistry has been reported elsewhere (Pip, 1979,
1987a, 1988). Each site was examined for macrophytes by dredging with a rake
or using SCUBA. Search time at each site was limited to 1 h. The species of
some taxa were grouped as genera because reproductive structures required for
specific identification were often immature or absent. Surface water temper-
atures were measured with a thermometer; subsurface measurements were made
with a Yellow Springs Instruments telethermometer. Temperatures were mea-
sured where the plants were growing; where plants were tall and vertically
spanned several thermal strata, the maximum temperature at the top of the
shoots was recorded.
While most locations were visited only once, 37 sites were sampled a number
of times during the season and in different years. Monthly values for these
sites were included in statistical analyses. A number of sites was also visited in
February 1985, in order to determine temperatures beneath the winter ice cover.
Temperatures for all sites were grouped according to month. All of the dif-
ferent regions of the study area were represented in each month, to avoid bias
that might arise from possible regional temperature differences. No significant
differences in water temperature were found for the different years encom-
passed by the study. Sites were classified as lakes ( > 10 ha), ponds ( < 10 ha)
and lotic habitats. Cochrans C and Barlett-Box F tests (Sokal and Rohlf, 1981 )
were applied to all data subjected to one-way analysis of variance in order to
ensure variances were sufficiently homogeneous. The critical significance level
for all statistical tests was P < 0.05.

RESULTS

Temperature differences between the three classes of water bodies (Table

1 ) were examined using one-way analysis of variance. Data for sites sampled
once and those sampled a number of times were pooled. A total seasonal range
 369

TABLE 1

Mean monthly surface water temperatures at the 3 water body types

May June July August September

2 SE n Z SE n 2 SE n 2 SE n x SE n

Ponds 16.9 0.9 14 19.3 0.7 40 23.0 0.6 22 20.9 0.9 27 14.7 0.4 37
Lakes 17.1 0.9 16 19.0 0.8 25 22.9 0.2 91 21.1 0.5 33 15.1 0.4 33
Loticsites 15.5 1.3 6 18.8 1.4 13 21.7 0.4 26 19.7 0.8 22 12.9 0.6 17

of 0-30 ° C was encompassed by the study. The results indicated no significant

differences between site classes in May ( n = 3 6 ) , June ( n = 7 8 ) or August
(n -- 82 ). However, in July (n = 139), Student-Newman-Keuls multiple range
tests (Winer, 1971 ) indicated that lakes showed significantly higher temper-
atures than lotic habitats. In September (n-- 87), both ponds and lakes showed
significantly higher temperatures than lotic sites. Multivariate analysis of var-
iance (Tatsuoka, 1971) showed that time of season (month) was the most
important factor affecting water temperature (F--- 80.6, P < 0.001, n = 422),
while water body type was less important, but still significant (F= 4.3, P = 0.014,
n = 422 ). There was no significant tendency for any water body class to be over-
or under-represented in the sampling program in any particular month
(F=0.25, P=0.98, n=422).
Winter water and bottom sediment temperatures below the ice cover of 0.75-
> 1 m thickness ranged from 0 to 5 ° C at different site types, with the warmest
temperatures generally located in the sediments. It was not unusual to find
green, living, submerged macrophyte shoots at these temperatures under the
ice cover in late winter in some lakes.
The mean temperatures at which the more common species or genera were
found in each summer month are given in Table 2. Differences among species
were examined for each month and water-body class individually using 1-way
analysis of variance and Student-Newman-Keuls multiple range tests. None
of the interspecific differences was significant for any month within any of the
three water body classes. However, when all water body classes were pooled,
significant interspecific differences were found for May (F ratio=l.66,
P=0.018, n=182), June (F ratio---1.69, P--0.003, n--482), July (F ra-
tio=1.43, P=0.025, n=1230) and September (F ratio=l.46, P=0.029,
n= 551 ). In May, a significant difference was identified only between Pota-
mogeton foliosus Raf. and P. pectinatus L. In June, significant differences were
identified between P. foliosus and each of P. pectinatus and P. richardsonii
(Benn.) Rydb. and between Sium suave Walt. and each of P. pectinatus and
P. richardsonii. In July, P. praelongus Wulfen contrasted with each of Hippuris
vulgaris L., Callitriche spp., P. pusillus L. and P. vaginatus Turcz. August was
 370

TABLE2

Mean m o n t h l y w a t e r t e m p e r a t u r e s observed for each species in the study area

May June July August September

2 SE n x SE n 2 SE n 2 SE n x SE n

Potamogeton ampli[olius T u c k e r m . - - 17.0 2.5 3 23.8 0.6 19 21.3 1.4 7 13.3 0.8 9
P. epihydrus Raf. 17.0 - 1 19.7 1.6 7 22.6 0.7 16 20.6 0.7 20 17.5 0.5 2
P. foli[ormis Pets. - 22.7 1.9 3 23.3 0.4 16 - 12.0 - 1
P. filiosus Raf. 12.5 0.5 9 15.8 0.5 13 22.5 0.7 23 20.4 0.9 23 14.2 0.9 13
P. friesii Rupr. 18.0 - 1 23.8 3.8 2 23.7 0.5 22 22.5 1.2 4 12.3 1.5 3
P. gramineus L. - - 19.0 0.9 17 23.5 0.4 38 20.8 1.4 13 14.5 0.5 13
P. natans L. 14.8 2.7 3 18.3 1.6 8 22.5 0.4 32 22.2 1.0 17 13.4 0.7 15
P. obtusifolius Mert. & K o c h - - 23.1 0.8 7 20.3 2.6 3 -
P. pectinatus L. 18.2 1.0 13 22.9 0.8 18 23.0 0.4 32 21.6 0.7 22 15.1 0.4 29
P. praelongus Wulfen 20.5 0.5 2 17.4 1.6 5 24.3 0.5 22 20.5 0.9 4 14.0 0.8 10
P. pusillus L. 15.0 - 1 19.6 1.4 10 21.7 0.4 12 23.0 1.2 9 15.1 0.6 10
P. richardsonii ( B e n n . ) Rydb. 17.5 1.6 6 22.4 1.1 18 22.9 0.3 74 20.6 0.7 28 15,1 0.4 34
P. robbinsii Oakes 13.0 - 1 20.1 2.7 4 25.0 1.5 5 18.0 0.6 3 15.8 0.9 4
P. spiriUusTuckerm. 17.0 - 1 19.0 1.0 2 23.8 1.2 6 23.3 2.7 3 -
P. vaginatusTurcz. 22.0 1.0 2 21.8 2.6 3 21.8 0.5 16 - 14,8 0.9 7
P. zosteriformisFern. 14.3 2.1 4 19.2 1.8 9 22.9 0.4 39 19.2 0.8 16 14,3 0.6 20
Charaspp. 17.0 1.2 7 19.9 0.8 19 23.2 0.4 39 22.2 1.0 15 14,0 0.5 27
Sparganiumspp. 15.9 1.1 8 18.2 0.8 28 22.4 0.4 41 20.4 0.9 24 13,9 0.6 18
Ruppia maritima L. 19.5 1.1 5 23.3 0.3 2 23.0 0.6 9 25.0 - 1 12.3 2.3 3
Najasflexilis (Willd.) Rostk. & 14.3 2.1 4 18.2 1.5 6 23.4 0.5 34 20.6 1.2 10 14.1 0.7 12
Schmidt
Alisma triviale P u r s h 15.0 0.5 4 19.6 0.9 16 22.1 0.6 20 21.1 1.1 14 13.5 0.6 12
Sagittaria spp. 15.5 2.0 4 20.9 1.3 15 22.5 0.3 49 19.8 0.8 22 13,6 0.5 20
Elodea canadensis Michx. 12.0 1.0 3 17.7 1.6 8 22.7 0.6 28 18.3 0.8 12 14.7 0.4 32
VaUisneria americana Michx. 17.0 - 1 20.1 1.5 7 23.0 0.5 30 21.2 1.0 11 14,5 0.5 21
Zizania aquatica L. 14.8 2.7 3 17.8 1.6 7 23.5 0.7 12 20.5 0.9 12 12,2 1.0 8
Eleocharis spp. 16.1 1.6 6 18.7 0.8 19 22.6 0.4 30 19.6 0.6 23 13.5 1.1 8
CaUapalustris L. 18.5 1.5 2 21.0 2.0 7 22.7 0.7 13 20.5 1.2 6 -
Lemna minor L. 16.0 1.0 7 19.1 1.2 18 22.8 0.3 37 20.6 0.9 22 13.6 0.5 18
L. trisulca L. 16.6 1.2 11 20.0 1.6 14 22.4 0.5 27 20.1 1.1 18 14.0 0.6 17
Spirodelapolyrhiza (L.) Schleid. - - 19.2 2.8 5 23.1 0.5 26 20.2 0.8 17 12.2 1.5 3
Polygonum amphibium L. 17.3 1.5 9 22.3 0.4 37 18.5 0.6 19 13.9 0.7 11
CeratophyUum demersum L. 15.5 0.9 16 18.8 1.1 21 22.8 0.3 57 20.2 0.7 37 14.6 0.5 26
Nymphaea odorata Ait. 17.0 - 1 15.7 1.2 3 23.6 0.8 11 20.2 1.6 5 14.8 1.0 4
N tetragona Georgi - - - - 24.0 0.6 14 18.7 1.3 3 14.5 0.5 2
Nuphar microphyUum (Pers.) Fern. - - 18.0 4.0 2 23.5 0.6 12 25.5 0.5 2 -
N. variegatum Engelm. 15.4 2.0 4 20.8 1.6 11 23.1 0.4 39 21.2 1.0 19 12.5 0.9 11
Ranunculus aquatilis L. 13.5 1.4 3 16.7 1.0 5 23.3 0.7 14 19.8 2.0 6 13.2 1.0 9
CaUitriche spp. 15.0 - 1 18.3 0.3 2 21.3 0.7 13 21.3 1.2 12 -
Myriophyllum exalbescens Fern. 17.3 0.8 24 20.9 0.8 35 22.5 0.3 83 20.7 0.7 33 15.1 0.4 42
Hippuris vulgaris L. 21.0 - 1 17.8 1.0 12 21.0 0.5 15 18.7 1.9 7 13.4 0.8 7
Slum suave Walt. 17.5 1.2 4 16.7 0.9 22 22.2 0.5 23 18.5 0.8 11 13.3 0.6 8
Mentha arvensis L. 18.0 1.6 3 17.9 1.2 8 22.2 0.5 14 20.2 1.1 5 -
Utricularia intermedia H a y n e 15.8 1.1 5 22.5 0.6 18 20.1 0.9 14 13.3 0.7 5
U. minor L. 14.0 - 1 18.1 2.1 3 22.7 0.7 9 18.9 1.5 7 16.5 0.5 2
U. vulgaris L. 15.6 0.9 9 18.6 0.8 32 22.6 0.3 51 20.3 0.8 27 13.4 0.7 16
Megalodonta beckii ( T o r r . ) G r e e n e 11.9 0.8 4 16.6 1.2 6 22.6 0.7 19 19.4 1.4 11 13.4 0.8 15
ZostereUa dubia Jacq. 10.0 - 1 14.0 - 1 21.3 2.4 4 19.8 1.0 8 15.8 0.7 10
 371

associated with no significant differences, but in September Zosterella dubia

Jacq. was found at significantly higher temperatures than Zizania aquatica L.
Macrophyte species richness was examined by m o n t h in each of the water-
body classes. Species richness was significantly positively correlated with tem-
perature only in July in lakes (r--0.22, P= 0.015, n--95 ), when water temper-
atures reached the seasonal maxima. Thus, the warmest lakes also tended to
show the greatest numbers of macrophyte species.

DISCUSSION

It has been suggested (e.g. Barko and Smart, 1981 ) that different species of
macrophytes may differ with respect to their ranges of thermal tolerance, which
may in turn affect the geographical ranges of various macrophytes. Although
interspecific differences could not be demonstrated within any given water-
body type in the present study, some differences emerged when all sites were
prefpooled. These differences were due in some cases to different water body
tolerances of certain species (Pip, 1979, 1987a,b) and the temperature differ-
ences seen for the respective water body types in July and September. How-
ever, in other cases, differences were seen as a result of the increased numbers
of samples when sites were pooled.
The results of the present study showed that, while at some times of the year
some species occupied habitats that were distinctly colder than those of other
species, these differences did not remain consistent throughout the growing
season. Because of the wide ecological temperature tolerances of most of the
species examined, none of the taxa could be classed as either cold- or warm-
water types within the area studied. While temperature may have operated
indirectly through its effects on variables such as pH and other chemical vari-
ables, since many of the macrophytes in the area show well-defined ecological
ranges with respect to water chemistry (e.g. Pip, 1979, 1987a,b) temperature
did not in itself appear to influence the geographical distributions of individual
species, in terms of limiting the sites where any given species could occur. Of
course, it was still possible that some of the rare species occurring in the region
may have had thermal restrictions, but these could not be examined statisti-
cally owing to low sampling numbers.
A number of species, e.g. Potamogeton foliosus, P. gramineus L., P. natans
L., P. pectinatus, P. pusiUus, P. richardsonii, Chara spp., Alisma triviale Pursh,
Lemna minor L., L. trisulca L., CeratophyUum demersum L., Callitriche spp.
and MyriophyUum exalbescens Fern., were observed in shallow ponds where
the surface temperature was as high as 30 ° C. However, ponds were also the
most strongly vertically stratified, where a difference of as much as 11.5°C
could be observed in the first 0.5 m below the surface on a calm hot day. Thus
larger rooted macrophytes reaching the surface in such habitats experienced a
wide range of temperature throughout the length of their shoots, although the
372

r e p r o d u c t i v e o r g a n s were e x p o s e d to t h e highest t e m p e r a t u r e s . A n u m b e r of
w o r k e r s (e.g. A n d e r s o n , 1969; S t a n l e y a n d N a y l o r , 1972; T i t u s a n d Adams,
1979 ) have r e p o r t e d t h a t some m a c r o p h y t e s can w i t h s t a n d w a t e r t e m p e r a t u r e s
as high as 35 °C a n d m a i n t a i n high rates of p h o t o s y n t h e s i s .
Since differences b e t w e e n t h e t e m p e r a t u r e ranges o f t h e various species were
m i n o r , species richness was also relatively u n a f f e c t e d b y t e m p e r a t u r e . T h e cor-
relation between m a x i m u m s u m m e r t e m p e r a t u r e s a n d species richness in lakes
m a y have b e e n r e l a t e d to t h e fact t h a t t h e w a r m e s t lakes were also t h e m o s t
shallow a n d n u t r i e n t - r i c h , a n d t h e r e f o r e p r o v i d e d the greatest n u m b e r of niches
a n d r o o t i n g areas for m a c r o p h y t e s .
T h u s , t e m p e r a t u r e in itself a p p e a r s to be relatively u n i m p o r t a n t in deter-
m i n i n g species d i s t r i b u t i o n o f m a c r o p h y t e s w i t h i n the s t u d y area. It is likely
t h a t o t h e r variables, such as depth, light availability, b o t t o m s u b s t r a t e type,
t u r b u l e n c e a n d w a t e r c h e m i s t r y (e.g. K a d o n o , 1982; Pip, 1987a,b, 1988) are
the p r i m a r y g o v e r n i n g factors for m a c r o p h y t e d i s t r i b u t i o n in the s t u d y area.

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