•\.

.•,

Literature re:view.:of�:t• he. Genus

•

Hydrocotyle L. (Apiaceae),
with particular emphasis on
Hydrocotyle ranunculoides L.f.

Swan River Trust

Report No 18
1994
SWAN RIVER TRUST
PO BOX 7248 Cloisters Square
PERTH
Western Australia 6850
Telephone: (09) 327 9700
Fax: (09) 327 9770

ISBN 07309 5987 2

ISSN 1033 9779
Literature review of the Genus
Hydroctyle L. (Apiaceae),
with particular emphasis on
Hydrocotyle ranunculoides L.f.

Swan River Trust

16th Floor 'London House'
216 St George's Tee
Perth 6000

Catherine McChesney
February 1994
Report No 18
------------•------------
=
=
FOREWORD
Hydrocotyle ranunculoides has become a significant ecological problem for the Canning River environment.
In an effort to gain a greater understanding of the ecology and biology of this aquatic weed, the Swan
River Trust sponsored a summer project at the University of Western Australia to;

1. To review literature relating to the biology of the genus Hydrocotyle and to investigate seed viability
and germination responses in the local weed species, Hydrocotyle ranunculoides.

2. Prepare a report on the results of the investigation as a basis for further studies into the control and/or
eradication of Hydrocotyle ranunculoides.

This publication covers the review of literature relating to the biology of the genus Hydrocotyle. A further
publication will contain information on seed germination and salinity tolerance.

Author's Acknowledgements
The project was undertaken as a Summer Scholarship provided to the Department of Botany, The University
of Western Australia, by the Swan River Trust. The author gratefully acknowledges the support of the
Swan River Trust in making this Scholarship possible. Greatly appreciated was the guidance of Ms.
Klemm, Ms. Ruiz-Avila, Dr Loneragan, Assoc/Prof Bell and Dr Dodd throughout the study, and the
assistance of Mr Waldie in setting up glasshouse experiments.

Maps in Appendices prepared by B Harrison

Cover Drawing by R Ruiz-Avila

Disclaimer
The recommendations contained in this report reflect the opinion of the author and may not necessarily reflect the opinion of
the Swan River Trust.

ii
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=
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SUMMARY AND RECOMMENDATIONS.

A revision of the literature relating to the genus Hydrocotyle revealed that:

1. it is a widespread genus characterised by a rhizomatous creeping growth form, with many species
occurring in aquatic or moist terrestrial habitats;

2. it is not known as a particularly weedy genus, but has been recorded as a weed in Chile (H.
ranunculoides), Georgia USA (H. ranunculoides), Argentina (H. bonariensis), Brazil (H. umbellata) and
Australia (H. ranunculoides, H. bonariensis, H. laxiflora and H. tripartita), although little information
appears to be available on the management of these species;

3. H. ranunculoides has been recorded as a component of the vegetation in a number of cool water
bodies occurring in either high altitude tropical lakes or low altitude coastal regions of the temperate zone.
Within the limitations of these cool water bodies, this species appears to be a 'generalist' in its ecological
responses;

3. H. ranunculoides is likely to proliferate in the Canning River due to its weedy plant characteristics
and modifications of this water way by human activity. Weedy characteristics include (i) a flexible growth
response, (ii) effective regeneration (e.g. vegetative propagation by fragmentation and possibly clonal
integration), (iii) possibly high resistance to herbivory and (iv) a co-occurrence with other weed species,
such as Eichhornia crassipes, Salvinia moles ta and Alternanthera aquatic um. Modifications to the Canning
River include the construction of the Kent Street Weir, which prevents upstream salt penetration during
summer, and an increase in nutrient-rich urban drainage, particularly upstream of the weir due to reduced
flushing to the ocean. The freshwater and high-nutrient conditions upstream of the weir, where H.
ranunculoides largely occurs, are likely to favour the growth of this species;

4. H. ranunculoides is utilized in aquatic wastewater treatment systems, particularly in Florida. It is the

weedy characteristics of H. ranunculoides (e.g. high productivity in high nutrient conditions, overwintering,
effective vegetative propagation) which makes this species so suitable for use in such systems.

RECOMMENDATIONS

Relatively little information appears to be available on the biology of H. ranunculoides or the genus as a
whole. Specifically, the review highlights the need for an understanding of aspects of the biology of H.
ranunculoides that includes the investigation of:

(a) successional status;

(b) the relative importance of asexual and sexual reproduction in contributing to population growth;

(c) the possible occurrence of ecotypes within this species;

(d) the effect of environmental parameters such as salinity, temperature and nutrients on the growth,
survival and reproduction of H. ranunculoides.

iii
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= TABLE OF CONTENTS

1. Overview 1
2. Introduction 1
3. Taxonomy 2
4. Geographic distribution 3
5. Ecology 4
5.1. Ecology of Hydrocotyle ranunculoides 4
5.1.1. Tropical Zone Waterbodies 4
5.1.2. Temperate Zone Waterbodies 8
5.1.3. Conclusion: Ecology of H. ranunculoides 9

5.2. Ecology of Other Hydrocotyle Species 10

5.2.1. Hydrocotyle verticillata 10
5.2.2 Hydrocotyle vu/garis 11
5.2.3. Hydrocotyle novae-zelandiae 12
5.2.4. Hydrocotyle bonariensis 13
5.2.5. Conclusion: Ecology of other Hydrocotyle Species 15

6. Genetics, uses and chemical properties 15

7. Herbicide applications 16
8. Herb ivory 16
9. Ecophysiology 17
10. Wastewater treatment and resouce recovery 19
11. Conclusion 22
12. References 25

lV
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1. OVERVIEW
= Eichhornia crassipes, Salvinia molesta and
Alternanthera aquaticum.
Hydrocotyle ranunculoides is a relatively recent Reasons for the proliferation of this species in the
weed of the Canning River of Western Australia. Canning River may relate to (i) environmental
Extensive rhizomatous mats of this species have factors, such as high nutrients, reduced salinity with
been known to reach from bank to bank in certain the construction of the Kent Street Weir, suitable
parts of the river, which may potentially undermine temperatures for rapid growth, and the absence of
the value of the water body by interfering with competitors and predators and diseases in this new
water use. This species is currently controlled in habitat, and (ii) 'weedy' plant characteristics such
the Canning River by a combination of chemical as effective dispersal and recolonization (by
and physical techniques, which are successful iil vegetative propagation and perhaps seed
the short term, but result in further spread in the germination), rapid growth, and the ability to
long term presumably due to reestablishment of tolerate a wide range of conditions (such as
fragmented segments after manual handling. nutrients, light and temperature). Further studies
Consequently, a further understanding of the of the biology and ecology of H. ranunculoides in
species is required for effective long term the Canning River are required to achieve
management. The objective of this paper is to successful long-term control or eradication of this
examine the literature relating to the genus species.
Hydrocotyle to assist in gaining an understanding
of H. ranunculoides.
The genus Hydrocotyle is a member of the large 2. INTRODUCTION
dicotyledon family, Apiaceae (Umbelliferae). It
Hydrocotyle ranunculoides is an aquatic
is a widespread genus of about 7 5 to 100 species
macrophyte that has invaded and proliferated to
characterised by a rhizomatous creeping growth
weed proportions in the Canning River Regional
form, with simple and usually inconspicuous
Park of Western Australia (Klemm et al., 1993).
umbels. Species mostly occur in moist habitats
The species was first observed in 1983, but its
ranging from aquatic (seasonally submerged,
distribution was not extensive until early 1991
floating or emergent) to moist terrrestrial
when it covered sections of the Canning River in
conditions. It is not regarded as a particularly
large mats, reaching from bank to bank in some
weedy genus, but some species are regarded as
places, and occupying an estimated volume of
weeds in Chile (H. ranunculoides), Argentina (H.
17,500 m 3• Physical removal of H. ranunculoides
bonariensis), Brazil (H. umbellata), and Australia
in November 1991 was successful in the short term.
(H. ranunculoides, H. bonariensis, H. laxiflora and
However, the physical impact of removal resulted
H. tripartita).
in a a further spread of the weed and, by September
Relatively little information is available on the 1992, the estimated volume within the Canning
autecology, synecology and management of H. River Park had increased to 40,000 m 3•
ranunculoides. However, several weedy
Spread of H. ranunculoides after its physical
characteristics are apparent from other members
removal was thought to be facilitated primarily by
of the genus and by the limited experimental work
vegetative propagation - small segments broken
with H. ranunculoides. For instance, H.
off from the large mats by manual handling are
ranunculoides is known to have (i) a flexible
able to generate into new mats after being
growth response (e.g. overwintering behaviour and
transported by the water to a new site. In the
possibly morphological plasticity with respect to
Canning River, the plant anchors to the bank at a
nutrients as shown by other creeping macrophytes);
depth of up to 15 cm, with the main bulk of the
(ii) effective vegetative propagation by
plant floating on the water. It has also been
fragmentation and possibly clonal integration (the
observed attached to overhanging branches and
latter being displayed by H. bonariensis in a
snags instead of to the river bank, although these
terrestrial sand dune environment); (iii) possibly
mats tend to remain small.
high resistance to herbivory and (iv) a co-
occurrence with other weed species, such as Klemm et al. (1993) claim that the current

1
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=
infestation of H. ranunculoides in the Canning
River has the potential to undermine the value of
===
3. TAXONOMY

the regional park, and further concerns that it may Hydrocotyle spp. are members of the Apiaceae
spread to other fresh water bodies were also (Umbelliferae), a large dicotyledon family of about
expressed. Accordingly, this species was gazetted 300 genera and 3000 species (Townsend, 1989).
as a Class Pl (preventing importation, movement Flowers of the Apiaceae are always small and
and trade of the plant) and Class P2 pest (plant formed in a simple or compound umbel. The fruit
requiring control and eradication) under the consist of two symmetrical parts known as
Agriculture and Related Resources Protection Act. mericarps, each containing one seed, which
Since this species is a common aquarium plant separate at maturity.
readily available throughout Western Australia, it Drude (1897) recognised three subfamilies within
is thought that H. ranunculoides first invaded the the Apiaceae according to their fruit and pollen
Canning River through the release of aquaria characters: (i) Hydrocotyloideae, (ii) Saniculoideae
wastes into the system. and (iii) Apoideae. The subfamily
The distribution of H. ranunculoides appears to Hydrocotyloideae is raised in rank by some
be limited by water salinity, largely occupying the authorities and treated as a separate family (Hickey
freshwater region of the Canning River upstream and King, 1988). Hydrocotyle is the largest genus
of the Kent Street Weir, constructed to prevent the within the Hydrocotyloideae, consisting of about
upstream penetration of salt during summer for 75 (Muenscher, 1944; Cook, 1990) to 100 species
agricultural purposes. The reduced salinity of the (Ewart, 1930; Webb et al., 1988; Johnson and
water above the weir has changed the vegetation Brooke, 1989). On the basis of fruit characters,
in favour of freshwater fringing vegetation (Olsen Hydrocotyle is the most similar to the Platysace,
and Skitmore, 1991). Furthermore, the Centella, Xanthosia and Trachymene, also of the
combination of reduced flushing and nutrient rich Hydrocotyloideae (Tseng, 1967).
urban drainage has made this section of the river The name Hydrocotyle is derived from the Greek
more eutrophic. Earlier outbreaks of the freshwater words hydor (water) and kotyle (a dish or plate),
weed, Salvinia molesta, were controlled by apparently alluding to the shape of the leaves,
draining the weir to allow salt water to penetrate which are slightly depressed in the centre (Black,
further upstream and by manual removal. 1957). It is one of the few genera of the Apiaceae
Currently, H. ranunculoides is controlled by a with entire leaves, most others possessing much
combination of chemical and physical control divided pinnate leaves (Hickey and King, 1988).
techniques (Klemm et al., 1993). While this is A distinguishing character of the genus is the
successful in the short term, further understanding presence of only one ovule in each mericarp
of the biology and ecology of this species is (Tseng, 1967). In all other members of the
necessary to achieve long term management Apiaceae examined, each mericarp contains two
success. The objective of this review is to examine ovules, although the upper ovule always aborts.
the literature relating to the genus Hydrocotyle in Hydrocotyle is often regarded as a relatively
an attempt to increase an understanding of H. primitive member of the Apiaceae, although it also
ranunculoides, and thereby to assist in devising possesses a number of apparently advanced
effective management strategies for its control and characters (Rodriguez, 1971). For example, the
eradication in the Canning River of Western laterally compressed fruit is regarded as a primitive
Australia. feature while the creeping herbaceous habit and
the absence of ventral bundles in the vascular
system of the ovary and fruit are considered to be
advanced features. Hydrocotyle is one of the few
genera of the Apiaceae classified largely on the
basis of leaf features (Constance and Dillon, 1990).
A description of the genus by Cook (1990) is as
follows:

2
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=
Stems floating or creeping;
==
central USA (Steward, 1990). Fernandez et al.
(1990) compiled a survey of aquatic weeds in South
rooting at most nodes. Leaves America and noted thatH. bonariensis (Argentina),
distinctly stalked, simple or H. umbellata (Brazil) andH. ranunculoides (Chile)
peltate or with basal sinus, were only cited once or twice in the survey, and
orbicular to reniform, entire, that the importance of these species as weeds is
toothed or lobed, stipules difficult to assess and the extent of problems caused
present. Umbels simple, by them is unkown.
sometimes proliferous, rarely an In Australia, H. bonariensis, H. laxifl,ora and H.
interrupted spike, axillary, tripartita are regarded as weeds in certain habitats
usually inconspicuous. Fruit (Auld and Medd, 1987). H. bonariensis is a native
ovoid-ellipsoid to suborbicular, of South America and occurs mostly in maritime
strongly flattened dorsally; habitats along the New South Wales coast and
mericarps with three subequal around Adelaide. It is common on dunes and in
ribs; fruit wall a woody inner wasteland, gardens and lawns grown on sandy
layer. Seasonally submerged to soils. Both H. laxiflora and H. tripartita are native
floating, emergent or terrestrial; to Australia. These two species are also listed as
entomophilous or perhaps Australian weeds by Swarbrick (1973). H.
autogamous; diaspores globose laxiflora (stinking pennywort) occurs throughout
or somewhat flattened the Eastern States and is common in pastures and
mericarps; probably dispersed open grasslands, wasteland and woodland. In the
in mud by water or perhaps by dry sclerophyll forests of Victoria, it is a dominant
animals. understorey species, although it is not mentioned
as being a weed in this habitat (Foreman and Walsh,
1993). This species occasionally persists in lawns
Comprehensive monographs of the genus
and along the banks of streams (Auld and Medd,
Hydrocotyle have been published by Richard
1987). H. tripartita also occurs in eastern
(1820) and Eichler (1987a,b).
Australia, mostly in sheltered places on the banks
of streams, and sometimes as a weed in lawns. This
4. GEOGRAPHIC DISTRIBUTION species has become naturalised in New Zealand
(Webb et al., 1988; Johnson and Brooke, 1989).
Hydrocotyle is a widespread genus, occurring in Within Australia, Hydrocotyle ranunculoides
tropical and temperate regions of both hemispheres appears to occur only in the Canning River,
(Hickey and King, 1988), and being particularly Western Australia. H. ranunculoides is native to
well represented in North and South America Europe (Marchant et al. 1987) and is included in
(Mathius, 1971). Fifty five species of Hydrocotyle the flora of America (Mathius 1936; Cook et al.
occur in Australia, of which 24 occur in Western 1974; Mason, 1957; Aulbach-Smith et al. (1990;
Australia and 11 in the Perth region (Marchant et Wijninja, 1989), Africa (Lock, 1973; Thompson,
al. 1987; Klemm et al. 1993). At least ten native 1985; Agnew, 1974; Denny, 1973; Harper, 1992;
and one naturalised Hydrocotyle species occur in Townsend, 1989; Gaudet, 1977) and Chile
New Zealand (Johnson and Brooke, 1989). (Steubing et al., 1980), although these references
Summaries of information found on the occurrence do not indicate whether H. ranunculoides is native,
of Hydrocotyle species in Australasia and the rest naturalised or a weed in these regions. H.
of the world are contained in Appendices 1 and 2, ranunculoides is, however, stated as a weed of
respectively. unknown proportions in Chile (Fernandez et al.,
Hydrocotyle is not well known as a weed genus 1990) and a serious weed of northern South
throughout the world. However, H. umbellata is America (Mathius, 1936). A summary of the
regarded as an aquatic weed in south-western USA geographical distribution of H. ranunculoides is
(Anderson, 1990) and Hydrocotyle spp. are listed contained in Appendix 3.
amongst potential weeds in eastern and north-

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=
5. ECOLOGY drawdown in Lake Naivasha in November 1973.
Initially, .three zones could be identified: a
5.1. Ecology of Hydrocotyle composite zone (dominated by Conyza spp., closest
ranunculoides to the dry land), sedge zone (dominated by Cyperus
papyrus, C. digitatus and C. immensus), and a
It appears that there is little information available seedling zone (closest to the lake water). After
on the autecology of H. ranunculoides. However, reflooding, Sphaeranthus dominated the seedling
an examination of the abiotic and biotic conditions zone and, by March 1975, the drawdown
occurring in habitats in which this species has been succession had progressed to a papyrus (Cyperus
recorded may provide possible reasons for its papyrus) fringe swamp.
proliferation in the Canning River environment.
The mature papyrus swamp community consisted
Hydrocotyle ranunculoides occurs in three
of plant species occupying the swamp-land and
intensively-studied high altitude tropical lakes:
swamp-lake interfaces, with very few species
Lake Naivasha of Kenya, Lake Bunyonyi of
occurring within the papyrus stand. H.
southwest Uganda and Laguna de la Herrera of
ranunculoides commonly occurred at the swamp-
Colombia. It is also recorded as a dominant species
lake interface, occupying the swamp edge in
of a Mehuin Brook of the temperate humid zone
shallow water (Gaudet, 1977). It is therefore
in Chile, and coastal USA.
apparent that H. ranunculoides formed part of the
sub-climax community along the edges of Lake
Naivasha. However, this does not necessarily
5.1.1. Tropical Zone Waterbodies
indicate that H. ranunculoides is itself a late
Lake Naivasha, Kenya successional species, particularly since it was also
common on recently exposed shore. An
understanding of the position of H. ranunculoides
Lake Naivasha is a freshwater closed basin lake in plant succession has implications for effective
which occurs at an altitude of 1890 m above sea weed management in the Canning River of Western
level (Harper, 1992). The lake occupies an area of Australia. Many weed species are pioneer or
approximately 140 km2 (Harper, 1992) and has an opportunist species that are difficult to control
average and maximum depth of 4 m and 7.6 m, because short-term programs provide the
respectively (Gaudet, 1977). The shoreline is environment these plants are best able to colonise
gently sloping, which is ideal for the development and exploit (Mitchell, 1980). For these species,
of a wet mud flora (Gaudet, 1977), and consists of allowing the habitat to develop naturally may be a
silts, clays and diatomites (Thompson and Dodson, more effective weed control strategy than weed
1963). Details of the lake's chemical properties removal (Wade, 1990). Provided that H.
are listed by Denny (1985), including a ranunculoides is shown to be consistently a late
conductivity of 363 us cm- 1, pH of 8.8, alkalinity successional species in the Canning River
of 3.3 meq dm· 3, and a sodium and chlorine environment, short-term weed control programs
concentration of 45 and 14 mg dm-3, respectively. may possibly be more successful than for
Although no measurements were made, opportunistic weed species.
eutrophication of the lake may be occurring due to
Lake Naivasha has been observed to undergo
high agricultural production as the lake's edge
considerable changes in aquatic plant composition
(Gaudet, 1977).
during the period of 1930 to 1988 (Harper, 1992).
The lake is characterised by a natural, unpredictable During the so-called 'normal-phase' of 1930 to
fluctuation in the water level, given the term 1975, species dominant outside the papyrus clumps
'drawdown'. A combination of the characteristic were Nymphaea caerulea (water lilies) and H.
drawdown and the freshness of the water is thought ranunculoides together with the submerged species
to give rise to a high species richness of aquatic Ceratophyllum demersum and Potamogeton spp.
plants associated with succession on wet mud at (Beadle, 1932; Gaudet, 1977). By 1982 the
the lake's edge (Harper, 1992). Gaudet (1977) submerged vegetation was completely eliminated
documented the succession following a natural from the lake (Harper, 1984). However, the

4
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emergent H. ranunculoides was still common
= Vegetation surveys were conducted on four
during this 'reduced phase' and was often seen representative swamp sites: Majera Bay, Sites A
colonising large floating Salvinia molesta mats, an and B, Bugoroba Swamp and Gweru Bay (Denny,
introduced species that dominated the floating 1973). H. ranunculoides was listed as present at
vegetation at this time. This floating community all sites excepting Majera Bay. The absence of H.
moved in rafts or islands of various sizes around ranunculoides from this site may be due to the
the lake as a result of wind action, and covered conditions (abiotic and biotic) or to the chance
about a quarter of the lake surface. Submerged event that dispersal did not occur to this area.
vegetation has been progressively returning since However, a determination of causes is uncertain
1984 and this return is coincident with a decline in since this objective was not directly addressed in
crayfish population density (Harper, 1992). The the study and no differences in conditions between
persistence of H. ranunculoides during the Majera Bay and the other sites were identified.
'reduced phase' of 1975 to 1983 suggests that, At Sites A and B, H. ranunculoides was common
unlike the submerged species, it is not severely on floating mats dominated by Cladium
predated, if at all, by the introduced crayfish, jamaicense. Other 'island' species included
Procambarus clarkii. Polygonum salicifolium, Pycnostachys coerulea
and the fem, Thelypteris squamigera. Excepting
Lake Bunyonyi, Uganda the presence of these floating mats at Sites A and
B, the vegetation pattern was similar to Majera Bay.
Lake Bunyonyi is another high altitude (1950 m
above sea level) lake of Tropical East Africa Bugoroba Swamp was different in physical
(Denny, 1973). It has an open water surface of structure to the previously mentioned swamps of
approximately 60 km2 and an additional area of Lake Bunyonyi (Denny, 1973). It occupied a
21.7 km2 that is covered with swamp vegetation. shallow bay (with the depth less than 3.5 m) that
Lake Bunyonyi is a dammed valley lake and, unlike was considerably silted, with the entire bottom
Lake Naivasha, has a steep-sloping shoreline and consisting of very soft anoxic mud. The dominant
is comparatively deep, with a maximum depth of species of the swamp was Cladium jamaicense
40 m. Permanently anoxic conditions occur below which formed large masses of floating vegetation
7-15 m, depending upon local weather conditions, as it did at swamp SitesAandB. A 100 m2 quadrat
and the epilimnion is nutrient-poor and low in taken within the swamp indicated that H.
phytoplankton (Denny, 1972). No obvious changes ranunculoides formed a part of this community,
in the bottom substrate occur around the lake although its cover was regarded as insignificant.
shores. The lake is relatively undisturbed, and Present in slightly higher proportions was the
artificial fertilizers have not been used in the related species, Hydrocotyle mannii.
cultivation of nearby land. H. ranunculoides was also scattered throughout the
Two distinct communities occurred within and on floating mats of Cladium jamaicense at the swamp/
the edges of the lake: (i) a steep, exposed shoreline lake interface of Gweru Bay. The main inflows to
community and (ii) a swamp community in the the lake occurred at Gweru Bay. The rivers
shallow sheltered bays and inflows (Denny, 1973). provided a gentle flow through the swamp and at
Wind and wave action characterises the steep, the mouth oxygen readings indicated that the water
exposed shores. Only a narrow band of vegetation was well oxygenated except immediately above
occurred in this region, dominated by an outer belt the mud surface. H. ranunculoides, Polygonum
of the finnly rooted emergent species Phragmites salicifolium, Thelypteris squamigera and Cyperus
australis, usually with an inner belt of Cyperus nudicaulis were scattered amongst the Cladium
papyrus. Floating-leaved Potamogeton sp. and jamaicense and Cyperus papyrus rhizomes of the
Nymphaea caerulea sometimes occurred outside floating mats. Potamogeton x bunyonyiensis and
the emergent species, and submerged species, Nymphaea caerulea occurred around the outer
including Ceratophyl/um demersum, that grew fringes of the rafts, Utricularia vulgaris formed
down to 8 m depth. In the sheltered swamps, floating masses in the water around the rafts, and
floating mats of Cyperus papyrus and C ladium Hydrilla verticillata and Ceratophyllum demersum
jamaicense dominated. covered the bottom down to 5 m.

5
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In addition to these specific sites at Lake Bunyonyi,
= Laguna de La Herrera, Colombia
the C/adium-dominated floating mat communities Laguna de La Herrera is a 2.9 km2 lake that occurs
were a common feature of the lake as a whole. on the high plain of Bogota, Colombia, at an
The vegetation was examined on one large mat altitude of approximately 2550 m (Wijninga et al.,
floating in open water. The mat measured 75 m 1989). It experiences a bimodal distribution of rain
by 65 m and was composed of close mesh (occurring from April-June and from September-
interwoven roots and rhizomes of Cladium which November), with a mean annual rainfall of 635
formed a raft up to 0.64 m thick. The upper layers mm. Temperatures are cool, with a yearly average
of the mat were interspersed with dead and of 12.5°C and the occasional night frost. The lake
decaying organic detritus with little or no inorganic is polluted primarily by detergents draining from
sediment, and oxidising conditions prevailed. In nearby sloping pastures, and also by cattle dung
addition to Cladium jamaicense, Thelypteris and artificial fertilizers on nearby grasslands.
squamigera was abundant while other swamp Wijninga et al. (1989) conducted a detailed survey
plants including H. ranunculoides and Polygonim of the tropical mountain lake vegetation of Laguna
salicifolium were common. Floating-leaved and de La Herrera because it is the last big natural lake
submerged aquatic macrophytes occurred in pools on the high plain of Bogota, and therefore has
between the mats, and included Nymphaea conservation value.
caerulea, Potamogeton thunbergii, P. x
Wijninga et al. (1989) recognised ten community
bunyonyiensis, Lemna minor, Ceratophylla
types surrounding and within the lake (Table 1).
demersum and Utricularia vulgaris. Attached to
the outside of the raft were Hydrilla verticillata These communities were classified as either
terrestrial, helophytic or pleustophytic, a system
and Chara sp.
not found to be employed by any other authors.
The commonest swamp plant in tropical East Helophytes are defined by Cook (1990) as
Africa is Cyperus papyrus (Carter, 1955), and "essentially terrestrial plants whose
Denny (1973) attributes its reduced abundance at photosynthetically active parts tolerate long periods
Lake Bunyonyi to strong competition from other of either floating or being submerged", as distinct
emergents, particularly Cladium jamaicense and from hydrophytes, which are defined as "plants
Phragmites australis, at high altitude. P. australis, physiologically bound to water, with at least part
a pioneer species, appears to outcompete both C. of the generative cycle occurring in or on the
jamaicense and C. papyrus in exposed sites. surface of the water". However, these definitions
Presumably, P. australis is less successful in less do not appear to be adopted by Wijninga et al.
exposed areas because it is in direct competition (1989), since they used the terms helophytes and
with C. jamaicense and C. papyrus. hydrophytes interchangeably. Rather, it appears
The growth habits of C.jamaicense and C. papyrus that in this study, helophytes are regarded as
are similar, both forming rhizomatous mats which emergent or submerged plants. Furthermore, no
grow out over the surface of the water, and definitions of pleustophytes were located in the
fragment at their outermost fringes to form floating literature, although these species appear to correlate
islands. Denny (1973) suggests that these two to the free-floating aquatic plants of other
species could be direct competitors for swamp classifications (e.g. Gaudet, 1977).
sites, with the greater susceptibility of C.
jamaicense to fragmentation making it more likely
to float off in the rafts which accumulate in
windward bays. Interestingly, H. ranunculoides
also occupies a similar niche to these two species,
and it is therefore possible that its growth in the
Lake Bunyonyi system was kept in check by the
presence of competitors such as C.jamaicense and
C. papyrus, both of which are absent from the
Canning River environment.

6
------1------
Table 1.
=Comparison of presence/absence of Hydrocotyle ranunculoides, mean ECC and mean pH at
each of the community types recognised at the Laguna de la Herrera (Sabana de Bogota, Colombia) by
Wijninga et al. (1989).

COMMUNITY COMMUNITY PRESENCE/ MEAN MEAN

TYPE ABSENCE OF ECC pH
H. ranunculoides (us)
Hydrocotyle helophytic present (common, 106 6.0
ranunculoides dominant)
Polygonum punctatum helophytic present 187 6.2
(common)
Rumex obtusifolius I helophytic present 108 6.0
Polinonum punctatum (common)
Ridens laevis helophytic present 208 6.0
(moderate)
Phytolacca terrestrial absent - -
bogotensis

Scirpus californicus/ helophytic absent 130 6.0

Typha anius tifolia
Scirpus californicus helophytic absent 95 5.9

Eichhornia crassipes pleustophytic present (rare) 147 5.9

Limnobium pleustophytic absent 104 5.9

/aevigatum

Azolla filliculoides I pleustophytic absent 249 6.1

Lemna cf. miniscula

In one of the community types, H. ranunculoides filiculoides was dominant with a moderate to high
was the dominant species with a cover of 75% to cover, while other species (including Lemna cf.
100% (Fig. 1). Associated hydrophytes were gibba, Limnobium laevigatum and Ricciocarpus
Pennisetum clandestinum andLudwigia peploides, natans) were relatively rare. This community
although both were rare with a low percentage occupied shallow water of a depth ranging between
cover. Of the pleustophytic representatives, Azolla -40 cm to +22 cm.

Figure 1. Quadrat (2 m2) of the Hydrocotyle ranunculoides-dominated community of the Laguna de la

Herrera (Sabana de Bogota, Colombia). From Wijninga et al. (1989).

7
------•------
H. ranunculoides also occurred as an associate
species in five of the other community types, that
=
===
H. ranunculoides was present or absent (Table 1).
This species occupied both shallow swampy
is the shallow communities of Rumex obtusifolius/ regions and deeper regions of greater flow velocity.
Polygonumpictatum andPolygonumpictatum, and Mean pH was relatively consistent between the
the deeper communities of Bidens laevis, communities and no difference in ECC was
Eichhornia crassipes andLimnobium laevigatum. apparent. The communities in which H.
The Rumex obtusifolius (dominant) and ranunculoides occurred occupied a total of 64%
Polygonum pictatum (subdominant) community, of the lake, although its percentage cover was low
of which H. ranunculoides was a part, occupied in much of this area. The community in which it
shallow waters of a depth between -80 cm and 10 was dominant occupied only 1 %, and it was
cm. In the usually deeper conditions (5 cm to 60 common in communities occupying a total of 38%
cm) of the Polygonum punctatum community, H. of the lake.
ranunculoides was the associate species with other
helophytes being less frequent. This community
frequently occupied zones along marshy shores and
5.1.2. Temperate Zone Waterbodies
was not found in the centre of the lake. Mehuin Brook, Chile
The Bidens laevis community occurred in the
The brook of Mehuin supports permanently
course of the Bojaca River, and was persistently flowing fresh water in the region of Valdivia, south
the deepest part of the lake. H. ranunculoides was Chile (Steubing et al., 1980). It has a maximum
the associate species of this community, and depth of 40 cm, a mean pH of 6.2 (similar to the
displayed moderate cover. Of the other helophytic Laguna de La Herrera), gently sloping banks (5°C
species, only Ludwigia peploides and Rumex mean inclination), and a sand substrate. The area
obtusifolius were present, and had low cover. In in which it occurs receives high precipitation (mean
obvious contrast to the other communities, the annual rainfall of 2372 mm) and low temperatures
floating Azolla filiculoides layer was absent, (yearly mean of 12.1 °C). At the time of the study,
perhaps due to the stronger water currents. "
aquatic ecosystems of the Valdivia region were
The pleustophytic communities of Eichhornia oligotrophic, and without observable
crassipes and Limnobium laevigatum supported contamination. Mehuin Brook was totally covered
only a very low cover of H. ranunculoides. Other with floating-benthic hydrophytes of which the
helophytic species of very low cover that were dominant species were H. ranunculoides (70%
present in the Eichhornia crassipes community cover) Callitriche stagnalis (20% cover) Rorippa
included Polygonum punctatum and Ludwigia naturium-aquaticum (5% cover) and Mimulus
peploides, and associate pleustophytic species luteus (2% cover) (Steubing et al., 1980).
included Azolla filiculoides, Lemna cf. gibba and Marga-Marga Estero, Chile
Spriodela spp. Other helophytic species were
virtually absent from the Limnobium laevigatum The distribution patterns of the aquatic and marsh
community, while Azolla filiculoides and Lemna flora along the Marga-Marga stream in central
cf. gibba were common. Chile were studied during the 1982 summer-
-autumn season (Palma et al., 1987). H.
Communities in which H. ranunculoides was not
ranunculoides together with Elodea potamegeton
recorded included the terrestrial Phytoclacca
were the most abundant species. Distribution
bogotensis community (situated above the
patterns of the species were interpreted in terms of
waterline), the helophytic communities of Scirpus
substrate and physiochemical variables,
californicus!Typha angustifolius (average depth of particularly salinity and eutrophication of the water.
15 cm) and Scirpus californicus (depth of 40 to 50
Unfortunately, only the abstract is available in
cm), and the pleustophytic Azolla filiculoidesl
English and extraction of further information
Lemna cf. gibba community (depth of 15 to 40 requires translation.
cm).
Coastal United States of America
No marked differences in the measured
environmental parameters were apparent between H. ranunculoides occurs mostly on sandy
communities of the Laguna de La Herrera in which substrates in ponds, lakes and stream margins of

8
-----------•
= -----------
coastal USA (Fig. 2; Muenscher, 1944; Mason,
= r--r-,.·1:·.------ . 0- I\
r, ;--L·--,·___--·--:\t~~'---Y-ij;\V
\\, (~ ~·-•-
)•., .: • •-r••••-•/J.
1957; Aulbach-Smith et al., 1990). Unlike the high • ;\ --: ·. ·k~---- . · ·\I
altitude lakes of the tropical zones, it is restricted
to lower altitudes in temperate California (Mason, er:. f_·_· --
1957). In South Carolina, H. ranunculoides forms ~ \J::. /)/.:~<-~----- ~-----/···/t
large floating mats that can support other plants
such as Bidens spp., Polygonum densiflorum,
\ .·. __;._..•~---:
.
: .:·: ··---,,, : ..'
~ -,--,--···

·'.•.:_•):,.;.:._~;'(i
Alternanthera aquaticum and Myriophyllum 317 1'./
,....__ ~-- \
J
aquaticum (Aulbach-Smith et al., 1990), the latter
two species being noxious weeds in Australia Figure 2. Geographic distribution of Hydrocotyle
(Parsons and Cuthbertson, 1992). Extensive ranunculoides in the USA. From Muenscher
populations of H. ranunculoides have been (1944).
reported in South Carolina, occurring from autumn
to spring, but dying back during the summer after outflows, and in sheltered swamps. It is able to
flowering in April and May (Aulbach-Smith et al., establish itself off both steep (e.g. Lake Bunyonyi)
1990). This is in constrast to its growth pattern in and shallow banks (e.g. Lake Naivasha and Mehuin
the Canning River, where its maximal growth Brook). Gaudet ( 1977) claims that a gentle sloping
period is thought to occur during summer (Klemm shoreline is ideal for the development of a wet mud
et al., 1993). flora, although Klemm et al. (1993) observed that
H. ranunculoides was reported as a potential H. ranunculoides in the Canning River appears to
problem weed species in Ebenezer Creek, Georgia establish itself less successfully where the bank
(Newman and Thomaston, 1979). Ebenezer Creek has a gentler slope.
is a tributary of the Savannah River, and is a long, Similar pH occurred at the Laguna de La Herrera
narrow stream that drains large areas of low-lying and Mehuin Brook, both of South America, while
river swamp land. At the time of the report, the both pH and ECC was lower at the Laguna de La
creek was infested primarily with Alternanthera Herrera than the East African Lake Naivasha. In
philoxeroides, until control measures (such as coastal USA and Mehuin Brook, Chile, a sandy
introduction of the alligator flee beetle, Agasicles substrate is regarded as important for growth and
hygrophila, and the application ofKuron herbicide) establishment.
reduced its population to a few floating patches of
In all the studies, H. ranunculoides was a common
leafless shoots and stems. However, the decrease
component of the aquatic flora, but it was not
in Alternanthera philoxeroides resulted in a
recorded as proliferating to weed status as it does
considerable increase in the weeds H.
in the Canning River. Invading populations of H.
ranunculoides and Myriophyllum brasiliense.
ranunculoides may proliferate in the Canning River
Unfortunately, more recent reports on the status of
due to (i) high nutrient availability, (ii) the absence
H. ranunculoides in Ebenezer Creek could not be
of competitors, and (iii) the absence of herbivores.
found.
H. ranunculoides occupied both nutrient-enriched
(e.g. Laguna de La Herrera, Lake Naivasha) and
5.1.3. Conclusion: Ecology of H. oligotrophic systems (e.g. Lake Bunyonyi, Mehuin
ranunculoides Brook). Although it is difficult to compare the
productivity of H. ranunculoides between systems
Two distinct habitats can be recognised in which from the information provided in the literature, it
H. ranunculoides occurs: high altitude tropical was perhaps the most prolific in the polluted
lakes (of East Africa and South America) and low Laguna de La Herrera, where it was dominant or
altitude coastal regions of the temperate zone (of codominant over much of the lake, and in the
USA and South America). This species has been Marga-Marga Estero, where it was the most
observed attached to the banks of shallow pools, abundant species. This latter reference (Palma et
particularly in swampy regions, as well as in deeper al., 1987) is likely to contain information on the
water, often as art of a floating island community. effect of nutrients on the abundance of H.
It is present in both flowing water, such as river ranunculoides.

9
------•------
-=
In the East African lakes, the growth of H.
ranunculoides may be limited by competition with
genera of Lake Naivasha, Lake Bunyonyi and the
Laguna de La Herrera include Polygonum, Bidens,
other species, particularly Cladiumjamaisence and Rumex and Lemna.
Cyperus papyrus, both of which occupy a similar It is possible that the absence of H. ranunculoides-
niche and have a similar growth form to H. s pec ifi c herbivores in the Canning River
ranunculoides. Other species typically co- contributes to its proliferation in this environment.
occurring with H. ranunculoides in these lakes Prevalent fauna! species in Lake Naivasha, Lake
include Typha latifolia, Ceratophyllum demersum, Bunyonyi and the Laguna de La Herrera are
Nymphaea caerulea, Hydrilla verticillata and the summarised in Table 2, although no mention was
introduced Salvinia molesta. Of these species, both made in any of the studies of these species
Hydrilla verticillata and Salvinia molesta have specifically consuming H. ranunculoides .
both occurred in the Canning River. Common

LOCATION SPECTES COMMONNAME COMMENTS

Lake Naivasha Myoco.stor coypus aquatic rodent population explosion,
(Gaudet, 1977) known to eat
Nvmphaea
Lake Naivasha Proco.mbus clarkii crayfish decline of sumberged
(Harper, 1992) .. macrophytes
attributed to
introduction of
crayfish
-
Lake Naivasha Hippopotamus major grazers at night
(Gaudet, 1977) on drawdown flora,
excepting Polygonum
plants
Lake Naivasha Cattle major grazers at day
(Gaudet, 1977)
Lake Bunyonyi Biomphallaria sp. Bilharzia snail prevalent on
CDenny, I973) Ceratophyllum
demersum
Lake Bunyonyi Haplochromis sp. and fish large numbers
(Denny, 1973) Clarias co.rsonii
Laguna de La Herrera cattle observed feeding on
(Wijininga et al., water plants, esp.
1989) Limnobium
laevi~atum

Table 2. Summary of fauna! species in freshwater habitats of Hydrocotyle ranuncu/oides.

5.2. Ecology of Other Hydrocotyle and shallow ponds along streams, particularly
Species along the Atlantic and Gulf Coasts (Muenscher,
1944; Fig. 3). Rejmankova (1992) examined the
Ecological studies of other Hydrocotyle species
growth strategies of this species, as well as four
include those of the aquatic H. verticillata, the
other emergent macrophytes, Ludwigia peploides,
semi-aquatic H. vulgaris, and the terrrestrial sand
Myriophyllum aquaticum, Nasturium officinale
dune species, H. novae-zeelandiae and H.
and Oeanthe sarmentosa, all of which commonly
bonariensis.
occur in nutrient rich pools, sloughs and irrigation
and drainage canals throughout central and
5.2.1. Hydrocotyle verticillata southern California. The author suggests that these
species be classified as 'creeping emergents', as
H. verticillata is native to USA, occupying marshes opposed to erect emergents (which include species

10
------•------
such as Typha latifolia, Scirpus acutus and
Phragmites australis). Much study has been
=
= Rejmankova (1992) claims that creeping
macrophytes, at least in the case of the study
devoted to erect emergents, while information on species, benefit from human impact on
creeping emergents is lacking. watercourses and lakes. It is suggested that the
creeping macrophytes behave as typical stress
r,.,1 - , .- -- -- - - tolerators, according to Grime's life history
,=. -·,····:-~ .:)
·... / \.,...... f··· .. ; ~ ·.rri;r.<~ strategies (Grime, 1979). Most are able to survive
\ (~.··r~r··\~~···'.·::· -\·1 · ·/~~ periods of low groundwater levels as well as water-
logged conditions. Furthermore, in undisturbed
\ '-f~·:'-t/?~\. ;, .... , -·-)~.~-.--ii:, nonpolluted habitats, these species are usually
!1.. • ! ( : <..~. T\~"'·· ....
318 '
e:~.: •
/--..... .
·--•.· ~ - · - 1 • ':• -_."i..;-...···. inconspicuous, but their behaviour changes
substantially once they get established in fertile,
nutrient-rich environments. Here they grow
Figure 3. Geographic distribution of Hydrocotyle vigorously, developing extensive dense mats and
verticillata in the USA. From Muenscher (1944). showing typical traits characteristic of plants from
fertile habitats: high relative growth rates and a
rapid response to nutrient supply. If left
Members of the creeping emergents group belong undisturbed, the creeping emergents are usually
to unrelated taxa but they develop from very similar slowly outcompeted by erect emergents. In
growth forms, probably as a response to a common contrast, creeping emergents are likely to
environment (Rejmankova, 1992). The most outcompete erect emergents in disturbed habitats,
characteristic feature of creeping emergents is their typically from periodic dredging and water level
creeping mode of growth. They root in the fluctuations.
substrate and send out long prostrate or ascending
stems freely rooting and branching at nodes and
often creating dense tangled mats. Adventitious 5.2.2 Hydrocotyle vulgaris
roots, which develop on these long stems, are often
without any direct contact with the substrate. From H. vulgaris is a wetland species with creeping
this description, it is apparent thatH. ranunculoides rooted stems bearing leaves held aloft on long
could also be classified as a creeping emergent. petioles (Grime et al., 1988). It is largely restricted
Rejmankova's (1992) study of five creeping to unshaded mire, although plants rooted on ditch
emergents revealed that rapid growth, high nitrogen banks may extend into still aquatic habitats where
accumulation, allocation of most biomass and the leaves become submerged. However, it persists
nitrogen into above-ground plant parts, and rapid only at positions along reservior margins at the
decomposition were characteristic of each species winter water level, suggesting an inability to exploit
in nutrient rich sites. Erect emergents, in contrast, sites which are waterlogged throughout the year.
have much higher standing crops, with a larger As with most wetland species, its distribution is
proportion of the biomass allocated to below largely restricted to gentle slopes. It is mainly
ground rhizomes. Ludwigia peploides was able to restricted to soils of pH 4.5 to 6.0, and is absent
grow for a prolonged period of time at high density from calcareous soils. The latitudes at which it
without showing any signs of senescence, being occurs are wide-ranging, but it is more abundant
attributed primarily to its high leaf area, which was within the upland regions (up to 530 m) due to
suggested to be a characteristic of the creeping habitat destruction in lowland districts. The
emergents in general. H. verticillata was geographical distribution of this species is
particularly noted for its rapid rate of summarised in Appendix 2.
decomposition, with the half time of decomposition H. vulgaris tends to occur in local monocultures
being 7 days. Rapid decomposition of creeping or in a turf with other species, particularly
emergents was attributed to a low C:N ratio, Sphagnum (Grime et al., 1988). It has only been
generally around 10:1, compared with 20-30:1 in recorded from vegetation providing an almost
erect emergents (Rejmankova, 1991). continuous vegetation cover, although its

11
-----------•-----------
=
stoloniferous growth can facilitate its spread onto shoot:root ratio and live leaf:live stem ratio at final
bare disturbed ground adjacent to established harvest were used to analyse growth response.
colonies. H. vulgaris only rarely occurs in shaded Creeping herbs were generally tolerant of burial,
situations, where it may form an understorey to growing to the surface from 1.33 H burial. The
species such as Juncus effusus. Canopy height is original plant parts mostly died, but survival was
determined by the length of the petiole, with petiole facilitated by the regrowth of small pieces of stolon.
length and leaf size decreasing in dry or grazed H. novae-zelandiae, a small creeping herb endemic
habitats. Marked changes in morphology are also to New Zealand, was the only species whose
associated with plants growing beside water; shoot:root ratio increased up to a burial depth of
individuals rooted under water often have a 1.33 H. For this species, there was a substantial
disproportionately long petiole and lack the hairs decrease in total dry weight, with much of the
present on the leaves of terrestrial forms original plant dying (Table 3). However, the plant
(Sculthorpe, 1967). was able to reach the surface via fine petiole
H. vulgaris overwinters with minute leaves, extensions of one or two new leaves as well as
resuming growth in late spring. This is followed slender adventitious rooting from the remaining
by self-pollinated flowering in June to August, with stolon. This was reflected in the increased
seed being set from August to October (Grime et shoot:root ratio at the lowest depth (Table 3) and
al., 1988). Like H. ranunculoides, H. vulgaris the higher leaf:stem ratio with partial burial. It
regenerates vigorously by the creeping shoots was suggested that this regrowth from a 'bank' of
which often form extensive patches. For both dormant buds is important in situations of complete
species, detached fragments root readily, and are burial in the sand dune environment.
probably important in the colonization of stream- A second experiment examined the length of time
and ditch-side sites following disturbance (Grime the sand dune species survived complete darkness
et al., 1988; Klemm et al., 1993). The fruits of H. in a lightproof box (Sykes and Wilson, 1990b).
vulgaris, which float and possibly adhere to the Survival length ranged from 19 to 141 days, with
feet of birds (Ridley, 1930), may also play a role H. novae-zelandiae being the longest survivor. The
in colonizing more distant sites. However, Grime authors suggested that for many of the species, and
et al. (1988) claim that regeneration by seed is H. novae-zelandiae in particular, etiolation assists
probably of infrequent occurrence since: (i) it in the progression of the buried plant to the sand
appears to produce few flowers when growing surface. In the case of H. novae-zelandiae, the
amongst tall vegetation, (ii) no persistent seed bank original aerial parts died in the dark, but etiolated
has been detected, and (iii) seed germination in leaves grew up from the stolons, some collapsing
the laboratory has been unsuccessful,. with seed and others replacing them. In conclusion, the
dormancy breaking mechanisms requiring ability of H. novae-zelandiae to tolerate both burial
investigation. and darkness appears to assist survival in a mobile
sand dune environment.

5.2.3. Hydrocotyle novae-zelandiae

Sykes and Wilson (1990 a,b) investigated the

ability of about thirty native and introduced New
Zealand sand dune species to sand burial.
Tolerance of sand deposition involves both the
plant's ability to grow up through the sand and to
survive darkness while it does so. In one
experiment, plants were grown for 15 weeks under
laboratory conditions in four different treatments:
surface (S), burial to two-thirds plant height (0.66
H), fully buried(l.0 H) and full burial plus a further
third of plant height (1.33 H) (Sykes and Wilson,
1990a). Measurements of total plant biomass,

12
-----------•
= ----------- = (temporal variation). Portions of a clone also
Table 3. Comparison of total plant dry weight and experience temporal variation in soil moisture
shoot:root ratio of Hydrocotyle novae-zelandiae availability as a result of small-sand erosional
plants not buried (surface) or buried to various patterns on a dune slope (Evans, 1991). Variation
proportions of their height (H). Depths with the in nitrogen availability is caused by a spatial array
same letter are not significantly different (p 0.05, of decaying pockets of organic matter in the rapidly
data log-transformed). From Sykes and Wilson leached, nitrogen poor, dune soil. Abrupt changes
(1990a). in resource availability can occur within a meter
in this environment, and low levels of each of these
resources have been shown to limit plant growth
BURIAL DEPTH TOTAL PLANT SHOOT ROOT
DRY WEIGHT (g) (DRY WEIGHT)
in dune systems (Kachi and Hirose, 1983; Barbour
et al., 1985; Gibson, 1988).
In this type of environment, the most abundant
Surface 6.90a 2.57 a plant species are usually clonal (Silander, 1985).
0.66H 3.43 a 5.14 ab Several studies (Evans, 1988; Evans, 1991; Evans,
1992) examine the possibility that the maintenance
1.33 H 0.08b 12.87 b of physiological connections between H.
bonariensis ramets allows a clone to integrate
water, light and nitrogen in environments where
these resources are patchy and that this integration
5.2.4. Hydrocotyle bonariensis may confer an adaptive advantage in the sand dune
environment. Experiments were conducted in
H. bonariensis is a rhizomatous herb native to, and
which H. bonariensis clones were grown across
common in, coastal dune communities of the
artificially created light, water and nitrogen
southeastern United States (Fig. 4; Evans, 1992).
gradients with and without ramet connections.
Its rhizomes grow linearly, with solitary leaves and
roots being produced at regularly spaced nodes. Evans (1988) demonstrated that acropetal
A branch meristem occurs in the axil of the leaf at translocation of nitrogen (old to young ramets)
each node, and an inflorescence bud occurs resulted in highly significant benefits to clones
opposite the leaf. Each node is regarded as a ramet growing from areas of high nitrogen to areas of
since they are the smallest potential physiologically low nitrogen. Ramets responded to increased
independent units that are capable of producing nitrogen availability, from either intraclonal
new clones when planted separately (Fig. 5). New translocation or immediate uptake from the soil,
ramets are generated vegetatively through the by producing primary, secondary and tertiary
forward extension and lateral branching of rhizome rhizome branches. In low nitrogen conditions,
meristems located in the leaf axils. In the field, clones allocated a greater percentage of biomass
rhizome connections between ramets can persist to the main rhizome and very little to branching.
for up to three years, although within this time, Evans (1991; 1992) demonstrated the basipetal
severing of connections may occur as a result of movement (young to old) of water, nitrogen and
trampling, grazing and exposure of rhizomes to photosynthates between two rhizome branch
extreme heat or cold (Evans, 1992). systems that interconnected hundreds of ramets
The branched rhizome systems of H. bonariensis within a clone. Ramet response to clonal
can be composed of over a thousand interconnected integration was shown to be a function of an
ramets covering over 100 square meters of the interaction of light, water and nitrogen conditions
coastal dune environment (Evans, 1988). This (Evans, 1991). In high light conditions, nitrogen
environment is characterised by highly patchy and transport was shown to be facilitated by the
unpredictable light, nitrogen and water resource existence of a water gradient. When both nitrogen
conditions. The patchiness of light in dune systems and water were limiting, the water potential
is generally associated with the presence of long- gradient drove the movement of water and nitrogen
lived evergreen shrubs (spatial variation) and the to the deficient portion of the clone, whereas when
accretion of sand over portions of the clone nitrogen was the only limiting factor, nitrogen

13
------------•
;;;;;;;;;;;;
------------
supply was limited to local uptake. However, this
interaction between nitrogen and water did not
occur in the presence of a light gradient. A
combination of both a water and light gradient
apparently resulted in less basipetal water
movement than in high light alone, since there was
probably less transpiration in the shade.
Translocation of water and nitrogen to portions of
a clone deficient in these resources resulted in a
significant net benefit to the clone in terms of
fitness-related traits such as total biomass, ramet
proliferation and seed production (Evans, 1991).
Clonal integration facilitated an increase in
reproduction (increase in inflorescence and seed
production) under low nitrogen conditions (Evans,
1992). Severed ramets (i.e. those in which
translocation from nonlimiting resources did not Figure 5. Four ramets are located on the main
occur) responded to low soil moisture by increasing rhizome segment of a Hydrocotyle bonariensis
root production and decreasing sexual clone. The oldest ramet (on left) has a secondary
reproduction. However, ramets in local low water branch with a newly established ramet. Ramets
conditions that were connected to ramets in high are centred around a node and consist of a leaf,
water conditions showed an increase in clonal roots, a branch meristem, rhizome and a potential
proliferation and sexual reproduction. inflorescence. From Evans (1992).

Figure 4. Geographic distribution of Hydrocotyle In addition, Evans and Whitney (1992) examined
bonariensis in the USA. From Muenscher (1944). the benefits associated with clonal integration
among interconnected ramets spanning a soil
salinity gradient. While H. bonariensis does not
typically grow in saline environments, it sometimes
expands into salt marshes from adjacent dunes. In
rhizome-severing experiments conducted under
both field and laboratory conditions, H. bonariensis
was shown to proliferate ramets under saline
conditions, provided that these ramets were
connected to other ramets growing in nonsaline
conditions. Supporting ramets incurred no net cost
The translocation of photosynthates across light
in terms of biomass or ramet production, but there
gradients also allowed for sexual reproduction and
was an increased percentage allocation to roots and
clonal expansion in the shade, although this
rhizomes. In contrast, ramets grown in saline
incurred a net cost to the clone in terms of ramet
conditions without the benefit of clonal integration
and seed production (Evans, 1991). Leaf
showed high mortality and produced little or no
characteristics such as intemode distance and
net clonal growth.
petiole height varied according to light intensity,
but were not affected by local water and nitrogen Thus, it is apparent from the results of these studies
conditions or resource integration (Evans, 1992). that H. bonariensis clones are highly integrated
It was suggested that increased petiole height under with respect to water, nitrogen, photosynthates and
low light conditions is critical for the placement salt. .Clonal integration results in increased ramet
of the blade above accreting sand or competing proliferation which in turn increases the production
vegetation, while short, rigid petioles produced in of resource acquisition units (leaves and roots),
the open dunes allow a leaf to endure persistent resource transport and storage units (rhizomes) and
wind exposure. sexual reproduction units (inflorescences), thereby

14
 ====
increasing total clonal fitness. This physiological 6. GENETICS, USES AND
integration of resources allows a clone to CHEMICAL PROPERTIES
effectively utilize the temporally and spatially
patchy resource conditions of its coastal dune Fedorov (1974) summarised the chromosome
environment. numbers of 20 Hydrocotyle species, including H.
ranunculoides which has a diploid chromosome
number of 24 (Table 4). Moore (1971) noted that
5.2.5. Conclusion: Ecology of other the genus Hydrocotyle displays a wide range of
Hydrocotyle Species polyploids, up to 15-ploid, together 'With
aneuploidy at all levels.
Ecological studies of Hydrocotyle species show the
French (1971) lists a number of Hydrocotyle
adaptation of particular species to their
species which are used as food and medicinal
environment. The creeping aquatic macrophyte,
resources, including H. alchemilloides (aromatic
H. verticillata, was suggested to display plastic
properties), H. bonariensis (used for skin diseases
behaviour according to the nutrient status of the
and toothache), H. bupleurifolia (astringent; used
water body. It is an inconspicuous component of
for diarrhoea), H. moschata (aromatic), H.
the vegetation in nonpolluted waters, but flourishes
sibthorpioides (leaves eaten, cough remedy), H.
with an increase in nutrient levels. This is also
umbellata (used for liver and kidney ailments) and
likely to be the case for H. ranunculoides, an
H. vulgaris (aperitive, detergent and a Danish home
aquatic species which also fits the description of a
remedy for whooping cough). H. asiatica (syn.
creeping emergent macrophyte. In a sand dune
for Centella asiatica) is eaten by humans in Asian
enviroment, survival of H. novae-zelandiae is
countries and has a number of medicinal uses in
assisted by its ability to tolerate darkness and burial
India, including use as a tonic, diuretic and skin
by sand, while H. bonariensis is known to use
disease treatment (French et al., 1971). Cho and
clonal integration to facilitate survival in a patchy,
Kim (1988) report that this species produces
unpredictable environment.
biologically active triterpenoids that have possible
In terms of moisture, the habitat of H. vulgaris is therapeutical uses in ulcerations, extensive wounds
intermediary, persisting in moist aquatic and eczemas. Other useful species include H.
environments that are not waterlogged throughout japonica, used as a folk medicine in Korea (Cho
the year. This is one of the few Hydrocotyle species and Kim, 1988); H. sibthorpioides and H.
in which germination studies appear to have been maritima, both having hemostatic and antitumor
attempted. Seed germination is thought to be of activities (Asakawa et al., 1982); H. tincture,
limited importance to the regeneration of H. known for its healing activity in the treatment of
vulgaris, with vigorous vegetative propogation wounds (Morisset et al., 1987); and H. podantha,
apparently being sufficient. Interestingly, Bell which has spasmolytic properties (Rao et al.,
(1971) notes that the few genera of the Apiaceae 1979).
with reduced umbels and inconspicuous flowers
Flavanols, rather than flavones, predominate in the
(including Hydrocotyle, Centella and Lilaeopsis)
less advanced genera of the Apiaceae, including
also have very effective methods of asexual
Hydrocotyle (Harbourne, 1971). The authors
reproduction, although no quantitative studies exist
suggest that the replacement of flavonol by flavone
to confirm this possible correlation. Other
appear to have evolutionary significance within the
somewhat scant information on seed germination
Apiaceae family, and probably among the
relates to H. ranunculoides, with Townsend (1989)
angiosperms generally. The flavinol glycoside
reporting that the fruit of H. ranunculoides become
Quertcetin (3-Glucoside) is common within the
buried in the mud at maturity. While this may
Apiaceae, including in the leaves of Hydrocotyle
sometimes be the case, seed collected from upright
species (Crowden et al., 1969). Other chemical
stalks from the Canning River were shown to
investigations have identified the flavonol
germinate at 25°C after 11 days (pers. observation).
glycoside,
hyperin, in H. maritima (Asakawa et al., 1982);
the flavonol glycoside, Quercetin 3-(6"-

15
------•------
caffeoylgalactoside) in H. sibthorpiodes
(Shigematsu et al., 1982); the flavanol
=
=
SPEOES DIPLOID
isorhamnetin-3-0-beta-D-galactoside in H. CHROMOSOME
japonica (Cho and Kim, 1988); trans-beta-
NO. (2n)
famesene as the major terpenoid consituent of H.
sibthorpiodes and H. maritima (Asakawa et al., H. acutifolia 48
1982); the lignan, L-sesamin, in H sibthorpioides H. asiatica 18, 22
(Ina et al., 1987); the lineoleic-oleic-palmitic type
H. benguetensis 96
of fatty acid composition in the seed oil of H.
asiatica (syn. forCentella asiatica) (Suhail et al., H. bonariensis 22
1987); and Madecassis acid in both H. asiatica H. chamaemorus 72
(syn. for Centella asiatica) and H. vulgaris, a
H. dichondroides 22
triperpernoid compound thought to assist in
defence against venous diseases, but absent in H. H. dissecta 44
ranunculoides (Tomei et al., 1989). H. javanica 72, 96
H. keeluniensis 24
7. HERBICIDE APPLICATIONS H. leucocephala 32, 48
H. mexicana 32
No information was found in the literature relating
to an effective herbicide for the management of H. moschata about36,48
Hydrocotyle ranunculoides. However, H. H. novae-zeeland iae 48
bonariensis, a dominant weed in the rubber
plantations of Bahia, was shown to be fairly H. poeppz":;(ii 48
effectively controlled by the application of H. ranunculoides 24
glyphosate (Lima and Pereira, 1985). The order H. rotundifolia 24
of herbicide efficiency based on the total weed dry
weight (composed of H. bonariensis as well as H. setulosa 48
other weed species) was (per hectare): 1.3 kg H. sibthorpioides 24,48,64
glyphosate> 1.5 kg oxyfluorfen + 0.2 kg paraquat H. umbellata 48?
> 2 kg diuron + hexazinone > 3 kg diuron +
ametryn + 0.2 kg paraquat. H. vul~aris 96

Trials on the North and South Islands of New Hydrocotyle sp. about 160
Zealand indicated that the herbicide, clopyralid (unidentified)
(Versatill™), was ineffective on the Hydrocoty/e
species that were present (Saville, 1987). Table 4. Diploid chromosome numbers of a
Furthermore, the aquatic herbicide, Sonar™ number of Hydrocotyle species. From Federov
(fluridone) was reported to kill many vascular (1974).
submersed weeds (includingHydril/a, Equisetum,
Potamogeton, Cabomba, Najas, Egeria and long-term environmental impact on the total
Utricularia spp.), but Hydrocotyle spp. were ecosystem. H. ranunculoides has been included
tolerant to the herbicide (Tarver, 1985). Other in at least two studies to determine its palatability
tolerant species included Chara, Nitella, to potential predators.
Pontederia cordata, Juncus effusus and Scirpus Firstly, H. ranunculoides was included in a study
spp. designed to test the host specificity of a small water
weevil, Neohydronomus pulchellus (DeLoach et
al., 1976). The weevil was shown to be host
8. HERBIVORY specific, feeding almost entirely on water lettuce
(Pistia statiotes, a serious aquatic pest in India,
Biological control is a favourable method of weed
Ghana, Loas and Thailand), and not feeding on H.
management in that it is likely to have the lowest
ranunculoides at all. Due to its demonstrated host

16
specificity, the water weevil was suggested as a (Maxwell's Paddock) in 1982-1983, but was
possible biological control agent for water lettuce. present in a further sampling in 1986. A number
Secondly, Santos et al. (1988) examined the of other new species were recorded, and this was
attractiveness of extracts of macrophyte leaves to thought to be related to the natural succession of
the snailBiomphalaria tenagophila. Amongst the plants after clearing. Interestingly, the
macrophyte species examined, preference was in establishment of H. laxiflora only occurred in
the order of: Nasturium pumilum, Polygonum acre, treatments where the kanagaroo was excluded,
Commelina sp. = Echinochloa crusgalli. However, although the presence of the rabbit did not appear
extracts from Hydrocotyle ranunculoides and to affect establishment. Similarly, at another study
Heteranthera reniformis were unattractive to the site (Tidbinbilla River), the height of H. laxiflora
snail. It is therefore apparent that no suitable had dramatically increased in plots where the
biological control agents for H. ranunculoides in kangaroos had been excluded, although no physical
the Canning River are evident from the available signs of grazing were evident. It therefore appears
literature. In any case, a potential biological control that the grey kangaroo feed on H. laxiflora,
agent must be very host specific to H. although direct evidence is required to confirm this
ranunculoides, with particular care taken to select suggestion.
an agent that does not eat native species and In central Florida, the dipteran Coquillettidia
commercially important members of the Apiaceae, perturbans infested a solid vegetation mat
such as carrots. composed of Hydrocotyle umbellata, Ludwigia
Other Hydrocotyle species are known to serve as octovalvis and Panicum spp.(Slaff and Haefner,
food sources. For instance, H. asiatica (syn. for 1985). Unfortunately, only the abstract of this
Centella asiatica) is among the most popular paper was available, and this did not indicate
traditional Malaysian vegetables, and Mansor whether or not H. umbellata serves as a foodplant.
(1988) outlines the nutritional contents of this
species. Secondly, Rowell (1985) determined the
diet of eleven Costa Rican species of the 9. ECOPHYSIOLOGY
grasshopper genus Rhachicreagra by direct
H. ranunculoides, among other typical southern
observation and by faecal analysis. Plant species
USA wetland species, was analysed for its
of the Compositae were the principal component
contribution of methane emissions to the
foodplant of many of the grasshopper species,
atmosphere (Sebacher et al., 1985). Methane is
although in montane habitats, Hydrocotyle species
produced by anaerobic bacterial decomposition in
became important foodplants. Hydrocotyle was
the organic-rich sediments that support the
included in the natural diet of six grasshopper
hydrophyte root system. In this study, the rooted
species. In summary, Hydrocotyle spp. were the
aquatic plants were shown to play an important
principal foodplant of R. dryomocnemensis; both
gas exchange role in the carbon cycle between
Hydrocotyle spp. and Compositae were the
wetlands and the atmosphere. However, detectable
principal foodplants of R. gracilis; andHydrocotyle
levels of methane were not emitted by H.
spp., H. mexicana and· Iresine diffusa
ranunculoides, Alternanthera philoseroides and
(Amaranthaceae) were the principal foodplants of
Lemna sp., most likely because these species, being
R. obsidian. Hydrocotyle comprised a minor
classified by the authors as free-floating, were not
component of the diet of R. anchidiphalara, R.
rooted in the anaerobic sediments during the
brachysphagicerca and R. melanota.
experiment. However, other observations have
In the Tidbinbilla Nature Reserve, Australian shown that H. ranunculoides can be rooted to the
Capital Territory, the native H. laxiflora appeared sediment (Harper, 1992; Klemm et al., 1993), and
to be eaten by the grey kanagaroo, Macropus it is possible that under these conditions this species
giganteus (Neave and Tanton, 1989). A large emits higher levels of methane.
proportion of the Reserve was previously cleared
H. ranunculoides was also included in a study
to facilitate the grazing of sheep and cattle, and in
comparing carbonic anhydrase (CA) levels and
1971 the Reserve was declared a Public Park. H.
lacunar CO2 concentration in a range of submerged,
la:xiflora was absent from one of the study sites

17
------•------
floating and emergent aquatic macrophytes of the these plants may also contribute to low
United States (Weaver and Wetzel, 1980). In this productivity. The results also indicated that the
study, H. ranunculoides was classified as an internal lacunar concentrations of emergents were
emergent species. The results indicated an increase generally lower than those of submerged plants.
in the CA levels along the habitat gradient of Submerged macrophytes may accumulate CO2 due
submerged to floating to emergent growth forms, to the slow diffusion of gas across the leaf surface
with H. ranunculoides displaying the highest CA in water, particularly in unstirred water where a
activity of all the species (Fig. 6). boundary layer of gas lies immediately adjacent
A direct relationship was found between CA to the leaf surface. In contrast, gas diffusion into
activities and primary productivities of these plants and out of'the lacunar system of emergent plants
over the habitat gradient: both productivity and CA is much more rapid because of their aerial growth
activity was low in submerged and high in form. Efficient gas diffusion is also likely to be
emergent plants. While a number of factors may faciliated by wide open stomata, since the soil in
be responsible for the low productivity of which emergents are rooted is usually saturated
submerged plants (e.g. low light, excretion of (Weaver and Wetzel, 1980).
dissolved organic compounds), low CA levels in It has been suggested that a possible function of
-----.----,--r--.----.-.---.-,-ny"-- CA is in the refixation of respired and
3- c - • - photorespired CO2 in C3 plants in the same manner
c-- -
IJ (_,__. ..... that phosphoenolpyruvate carboxylase scavenges
D , .... .,••
e -_..........- and facilitates the refixation of CO2 in C4 plants
e ~ ·- ...... . (Everson and Slack, 1968). If so, CA is likely to
'i' be important in emergent macrophytes such as H.
~ ~ . . -..~
0 ·- ....._....
,
ranunculoides, but less important for submerged
G-- u .... .,i.,,. ,o
plants since their lacunar system may serve the
v. --··••"•

D •- K • . .~ "
same function of refixing respired and
~ "'- • ., ... qo•- --+- photorespired CO2 (Weaver and Wetzel, 1980).
~[ ci':•::·.:. In another ecophysiological study, both H.
bonariensis and H. umbellata were shown to have
adaptive physiological responses to their specific
environment (Overdieck and Strain,
- ~ .......... ,., __ .... )=3,a.i
1981). Ramets were collected from three
z
~ .... .,_.
..... tlo<_ ..
'i:320.1
contrasting habitats on Shackleford Banks, a small
i f'.to•clolfl - barrier island off the central North Carolina coast
,_
of USA. One ecotype of H. bonariensis was
~[
,.,
,_ ·-···--·
S.e< ... ,.,,
collected from coastal sand dunes, and another
ecotype of this species was collected from the deep
.DI N
...~-·-·-M·- shade of an evergreen maritime forest. The third
NI M.. - - ~ · · , - , " . , . , _ , ___..
type, H. umbellata, is found only in fresh or
L---2L--'-4--'6-...L8--',,10--,lcl=-2--:-l.L.-4--:'16':---7-18~/'--
brackish water ponds. Like H. ranunculoides, this
E. U. (10.({,~/4. -9/mg PROTEIN] species is rooted to the bottom of the pond with its
leaves extend 1-2 cm above the water surface.
Figure 6. Carbonic anhydrase (CA) activities (EU Ramets were grown under controlled
± SE) of aquatic macrophytes grouped along environmental conditions and measurements of net
habitat gradients. The emergent Hydrocotyle photosynthesis, transpiration and total leaf
ranunculoides has the highest CA activity. 2a= diffusion resistance were used to examine possible
CA comparison between Scirpus subtenninalis, effects of atmospheric humidity on the three
one of the few submerged species of this genus, Hydrocotyle types. The results of the study
and the emergent S. acutus. 2b= CA comparison indicated that increasing habitat dryness had large
between submerged and floating leaves of Nuphar effects on gas exchange (CO 2 and water vapour)
variegatum. From Weaver and Wetzel (1980).

18
------•------
and on total diffusion resistance of the pond type,
moderate effects on the forest type, and no
=
conducted in Florida, although these systems are
beginning to gain widespread use in other tropical
significant effects on the dune type. It therefore or subtropical regions of the world. Aquatic
appeared that the habitat gradient from wet to dry macrophytes have been cultured in nutrient-rich
corresponded to a decrease in sensitivity to dry waters of sewerage effluents, agricultural runoff
conditions. and drainage effluents, runoff from cattle feedlots,
methane digestor effluents, and eutrophic lakes and
streams (Reddy, 1984). It is anticipated that the
10. WASTEWATER TREATMENT
importance of aquatic macrophyte systems in
AND RESOURCE RECOVERY wastewater treatment and subsequent fuel
production will continue to increase (Wolverton,
The most intensive research into Hydrocotyle
1987).
appears to be in relation to its use in aquatic systems
for wastewater treatment and biomass harvesting Most of the aquatic wastewater treatment and
for fuel production. Both H. umbel/ata and H. biomass energy systems have focused on
ranunculoides have been evaluated as potential Eichhornia crassipes, an obnoxious weed of many
beneficial species in such systems, although more regions, including the United States. Its suitability
emphasis has been given to H. umbellata. The in these systems relate to its high productivity and
benefits of these aquatic plant systems are two- the ease with which it can be stocked and harvested
fold: (i) wastewater treatment by aquatic plants due to its floating habit (DeBusk and Reddy, 1987).
offer low-cost, low-energy solutions to pollution In order to optimize the efficiency of wastewater
control (Crites, 1979; Tchobanoglous and Culp, treatment and energy production systems, a
1980; Whingham, 1982), and (ii) the conversion continuous supply of biomass should occur. One
of plant material into methane through aerobic of the shortcomings of the E. crassipes system in
digestion is an economical use for the biomass by- Florida is its lack of tolerance to low winter
product of the wastewater treatment systems temperatures, which can retard growth and cause
(Wolverton and McDonald, 1981; Reddy and death (Reddy, 1984). This has prompted research
DeBusk, 1984). into other potential species to use in waste
treatment and resource recovery.
Much of the research on this subject has been

Table 5. Nitrogen and phosphorus uptake (g m-2 d· 1) by the floating aquatic macrophytes, Eichhornia
crassipes and Hydrocotyle umbellata, during summer and winter in central Florida. From DeBusk and
Reddy (1987).

SPECIES NITROGEN PHOSPHORUS

Summer Winter Summer Winter

Eichhornia 1.30 0.25 0.24 0.05
crassipes
Hydrocotyle 0.37 0.37 0.09 0.08
umbellata

H. umbellata has been widely accepted as a useful potential nitrogen and phosphorus removal rates
substitute for E. crassipes during cold winter of a H. umbellata system during winter were found
months (Reddy, 1984; Reddy and DeBusk, 1984; to be 321-645 mg N m-2 day 1 and 103-106 mg P
Reddy and Tucker, 1985; Clough et al., 1987; m-2 day· 1·respectively (Reddy and Tucker, 1985).
Reddy, 1988; Agami and Reddy, 1991). This Reddy (1983) demonstrated that both E. crassipes
species has been shown to play a significant role and H. umbellata removed about 50% of the
in removal of nutrients from polluted waters inorganic nitrogen in a eutrophic system over 4
(Reddy, 1983; Reddy and Tucker, 1985). The days.

19
 -----------•
= ----------- = uptake, oxygen transport capacity is also an
~ 10001 important design criteria for aquatic plant
wastewater treatment and resource recovery
,:_ 800~
systems (Reddy et al., 1989). Aquatic plants
-6
~ 600!-
I Penny wort transport oxygen from foliage to roots, resulting
0l
-
I
I
in an oxidised environment in the rooting zone
en I (Armstrong, 1964). Rhizosphere oxidation has
~ 400L

! 2ool """iii' ' J[[[] Cl

Wa1erhyac1n1n
important implications in wastewater treatment
since it detoxifies hydrogen sulphide and the
reduced forms of iron and manganese, and activates
biogeochemical processes such as the oxidation of
0
o 12 24 36 48 60 72 days soluble organic compounds and nitrification of
January February I March NH4-nitrogen (Reddy et al., 1989).
Several studies have revealed that H. umbellata
has a high oxygen transport capacity. Moorhead
Figure 7. Biomass recovery rates of Eichhornia
and Reddy (1988) measured oxygen transport by
crassipes (waterhyacinth) and Hydrocotyle
plants as the net change in dissolved oxygen with
umbellata (pennywort) after a freeze in January
time, and demonstrated a greater transport rate by
1982 (Florida) for three consecutive nights (- l 7°C,
H. umbellata than E. crassipes (3.49 and 1.24 g
-3.3°C, and -2.8°C, respectively). From Reddy
0 2 kg· 1 dry root mass h- 1 , respectively) under
(1984).
laboratory conditions. Reddy et al. (1989)
measured oxygen transport rate as the oxygen
During a Florida summer, both productivity and consumed by the effluent (biological 0 2 demand,
nutrient uptake of E. crassipes exceeds H. or BOD5>and also found a greate: transport capacity
umbellata during the summer (Reddy and DeBusk, by H. umbellata than E. crassipes (10.land 2.75
1985). However, while nutrient uptake by H. mg 0 2 g- 1 plant dry wt, respectively) under
umbellata remains relatively constant throughout laboratory conditions. Clough et al. (1987)
the year, E. crassipes displays a sharp drop in measured percentage BOD 5 removal by both
nutrient assimilation during winter (Table 5). species in outdoor tanks in Florida. Typical BOD 5
Furthermore, Reddy (1984) demonstrated that H. reductions in the wastewater effluent varied from
umbellata recovered much faster from a hard freeze 70 to 90% during the study, with the highest BOD5
over three consecutive nights than E. crassipes, removal observed in tanks containing H. umbellata
resulting in significantly higher biomass yields during both warm and cool seasons.
(Fig. 7).
H. umbellata has also been successfully utilized
Interestingly, the standing crop of both species was in thin film systems for contaminant removal of
shown to be greater when grown together than polluted effluent (Dierberg et al., 1987a,b). A thin
when grown in monoculture (Agarni and Reddy, film system consists of vascular plants grown in
1991). H. umbellata was able to coexist with the shallow raceways through which a thin layer of
highly productive E. crassipes due to its elongated wastewater is passed. The plants develop a dense
petioles (avoiding shading by E. crassipes) and its mat of roots which, together with the colonizing
ability to outcompete E. crassipes during winter. bacteria, reduce suspended solids and BOD5 •
It was claimed that the improved growth of H. Dierberg et al. (1987a) showed that H. umbellata
umbellata in the presence of E. crassipes was due was able to substantially decrease the concentration
to the physical support provided by the E. crassipes of two common chlorinated compounds, 2,4-
plants. Thus, the results of this study suggest that dichlorophenol and monochloramine, from
the combined use of these two species in secondary domestic effluent under long-term,
wastewater treatment and biomass production continuous feeding conditions. The authors
would be successful in providing year-round claimed that this system is superior to the use of
optimum performance. chemical dechlorinating agents because it is
In addition to plant productivity and nutrient cheaper and circumvents the deoxygenation of

20
------------1
=
====
------------
water caused by dechlorinating agents. nitrogen and total suspended solids in the effluent
In a similar thin film system, H. umbellata was was about half of that in the influent. However,
shown to remove the heavy metals, copper and effluent water still had appreciable levels of all
lead, at levels of 69% and 85%, respectively measured variables except the amount of total
(Dierberg et al., 1987b). However, the high copper suspended solids, nitrate and nitrite after passing
loading proved to be ultimately toxic to the plants, through the aquatic plant treatment ponds. It was
and a subsequent decline in treatment efficiency suggested that the effectiveness of these species
occurred after one month's operation. The authors may be improved by aeration, since anaerobic
suggest that lower metal loadings to such thin-film conditions were created by the extensive cover of
systems are likely to enable greater heavy metal both species in the treatment ponds.
reductions by H. umbellata. H. ranunculoides was found to offer some
The results of such research into the use of aquatic advantages as a wastewater treatment species over
macrophytes in wastewater treatment and resource E. crassipes. Firstly, E. crassipes is a serious weed
recovery indicate that, while the productivity and species in the aquatic systems of Alabama, while
nutrient uptake of E. crassipes is the greatest of all H. ranunculoides is relatively uncommon in this
tested macrophytes under optimal growth region. Secondly, E. crassipes suffered winter kill
conditions, H. umbellata is a useful substitute while H. ranunculoides overwintered. In Alabama,
during the cooler months in E. crassipes-based perpetuation of natural stands of E. crassipes is
systems. Production of both species is higher when dependent upon seed germination or reintroduction
they are grown together than in monoculture, and the following growing season (Dinges, 1982). Like
the inclusion of both species facilitates efficiency H. umbellata, H. ranunculoides appears to be more
throughout the year. In addition to greater growth cold tolerant, successfully overwintering during a
and nutrient uptake than E. crassipes during winter, winter when E. crassipes was killed (Boyd and
H. umbellata was also shown to provide greater Bayne, 1988). The use of an aquatic plant that
BOD5 removal efficiency during both warm and does not require annual restocking, such as H.
cool seasons. ranunculoides, is economically desirable.
Less extensive work has been conducted on the Thirdly, H. ranunculoides showed less root
potential use of H. ranunculoides as an aquatic diminuation than E. crassipes. At the beginning
macrophyte for wastewater treatment and resource of the study, the root length of E. crassipes
recovery. Boyd and Bayne (1988) compared the exceeded that of H. ranunculoides, and by the end
efficacy of the use of H. ranunculoides and E. of the study, the reverse was true. Boyd and Bayne
crassipes in the treatment of wastewater discharged (1988) suggest that root mass maintenance by
from a livestock waste lagoon in Alabama. aquatic wastewater treatment plants is likely to be
Initially, the study examined E. crassipes only, but an important characteristic since the microscopic
after an incident of mass mortality due to an flora and fauna associated with the roots are
unknown cause, it was noticed that small quantities important in the improvement of water quality.
of H. ranunculoides that had invaded the tanks A further benefit of H. ranunculoides related to its
were not adversely affected. Consequently, this potential use as livestock food. While plant
species was also included in a subsequent trial. In nutritive content of both H. ranunculoides and E.
this second trial, both species were treated with crassipes compared well with other common
the insecticide, Sevin, and both remained healthy livestock forage plants, the former had a higher
throughout the growing season . crude protein and digestibility than E. crassipes
Maximum growth rate and standing crop was 0.009 (Table 6). Crude protein, fat and ash values of
lb ft· 2 day· 1 and 8.20 tonnes a· 1, respectively, for H. both species were similar to previous findings by
ranunculoides, and 0.0084 lb fr 2 day· 1 and 13.02 Wolverton and McDonald (1981).
tonnes a· 1, respectively, for E. crassipes. Both
plants were found to be effective in removing
pollutants from the water. Usually, concentrations
of total ammonia nitrogen, phosphate, biological
oxygen demand, chemical oxygen demand, organic

21
------•------ =
Table 6. Comparison of nutritive variables (dry weight basis) of Hydrocotyle ranunculoides and Eichhornia
crassipes grown in wastewater treatment ponds with common livestock forage plants. Unbracketed values
are from Boyd and Bayne (1988), bracketed values are from Wolverton and McDonald (1981).

SPECIES %CRUDE %FAT %ASH %ACTD % DIGEST-

PROTEIN DETERGENT IBILITY
;
FIBER
Hydrocotyle
rimunculoides 24.2 1.7 18.3 29.4 69.3
(23.4) (2.19) (17.4)
Eichhornia
crassipes 15.1 1.1 16.7 34.3 47.9
(14.7) (1.59) (11.1)
Alfalfa
(Medicago 18.9 3.0 10.6 35.0 55.4
sativa)
Bahia grass
(Paspalum 8.2 2.1 6.4 41.0 -
notatum)
Bermudagrass
(Crynodon 6.0 2.3 6.6 38.0 56.7
dact11lon)
Fescue
(Festuca · spp.) 9.5 2.0 10.0 39.0 51.4

While H. ranunculoides may be more suitable than it is particularly well represented in North and
E. crassipes for wastewater treatment (Boyd and South America. It is not recognised as a
Bayne, 1988), E. crassipes appears more suitable particularly weedy genus, but has been recorded
for subsequent fuel (methane) production as a weed in Chile (H. ranunculoides), Argentina
(Wolverton and McDonald, 1981). The efficient (H. bonariensis), Brazil (H. umbellata), and
anaerobic digestion process of both species is likely Australia (H. ranunculoides, H. bonariensis, H.
to be facilitated by their low lignin content. laxiflora and H. tripartita).
However, the content of hemicellulose, which is Weediness is determined by two major factors -
regarded as more amenable to bacterial degradation the environment and aspects of the plant's life
than cellulose, was greater in E. crassipes than in history (Mitchell and Bowmer, 1990). Important
H. ranunculoides. Furthermore, the maximum abiotic environmental factors include temperature,
volume of methane production was greater for E. salinity and nutrient availability, while major biotic
crassipes (0.198 m 3 kg 1 or 3.17 ft 3 lb· 1) than for H. environmental factors include the presence of
ranunculoides (0.146 m 3 kg· 1 or 2.34 ft 3 lb· 1). It is competitors, predators and diseases. In general,
therefore apparent that, under the prevailing life history characteristics such as high
experimental conditions, E. crassipes is easier to reproductive rate, rapid growth, short life-span,
digest anaerobically than H. ranunculoides, multiple reproductive strategies, and effective
suggesting its greater suitability as an energy dispersal mechanisms for propagules are associated
source. with weediness. Physiological adaptations which
may be of importance include an ability to
withstand water and salt stress, and a wide
11. CONCLUSION
tolerance to a range of temperatures and nutrient
Plants of the genus Hydrocotyle adopt a creeping levels.
habit, with many species being restricted to aquatic Relatively little information is available on the life
or moist terrestrial habitats. The geographic history characteristics, ecology and management
distribution of the genus is cosmopolitan, although of H. ranunculoides. It is not a widely known

22
- - - - - - - - - - - ===
•-----------
=
weed species, but does co-occur with other weed Few studies have been conducted which investigate
species, including Eichhornia crassipes, the effect of abiotic and biotic environmental
Alternanthera philoxeroides, Myriophyllum factors on the growth and survival of Hydrocotyle
brasiliense and Salvinia molesta. It appears to have species. Physical and chemical limits on the
a flexible growth response, for example, it survival of H. ranunculoides are difficult to
overwinters to avoid low temperature. It may also determine from the literature. It is consistently a
show morphological plasticity with respect to freshwater species, and is common in two distinct
nutrients, being known to undergo rapid growth in habitats, both of cool temperatures - high altitude
the high-nutrient conditions of wastewater tropical lakes and low altitude coastal regions of
treatment systems and the Canning River of the temperate zone. Within these habitats, it
Western Australia. The rapid growth rates of other attaches itself to a sandy or muddy substrate, or
aquatic creeping emergent macrophytes (e.g. H. forms part of a floating island community. No
verticillata) under nutrient non-limiting conditions preferences for water velocity, water depth, bank
are thought to facilitate vigorous invasion, slope, pH or nutrients were apparent from the
particularly in habitats disturbed by man. These literature. This species therefore appears to be a
creeping emergents are evidently inconspicuous in 'generalist' in its ecological responses within the
undisturbed habitats, but become weedy in limits of cool fresh water bodies.
eutrophic, disturbed habitats. Further research is
The absence of competitors and predators in the
required to determine whether the growth rate of
Canning River may also contribute to the success
H. ranunculoides also increases with an increase
of H. ranunculoides in this environment. It was
in nutrients as it is possible that the eutrophic
suggested that the growth of H. ranunculoides in
condition of the Canning River is at least partly
East African lakes was limited by competition from
responsible for this species' prolific growth.
other species of similar life form (e.g. Cladium
H. ranunculoides is capable of both sexual and jamaisence and Cyperus papyrus). Species of
asexual reproduction, although the contribution of similar life form in direct competition with H.
seedlings to population growth appears to be ranunculoides may be absent from the Canning
currently unknown. Little other information is River. Centella asiatica (syn. for H. asiatica) is
available on the seed biology of the genus except morpholgically similar to H. ranunculoides, being
forH. vulgaris which, however, is still thought to a rhizomatous plant which roots at the nodes.
depend primarily on vegetative propagation for However, this species tends to occupy more
regeneration. Indeed, the creeping rhizomatous terrestrial, rather than permanently aquatic, habitats
growth form of Hydrocotyle species is an extremely of the Canning River. Limited experimental work
effective method of propagation and may decrease on the palatability of H. ranunculoides suggests
the selection pressure for energy input into sexual that it species may be rather resistant to herbivory.
reproduction. In the aquatic environment, small Hydrocotyle species which have been shown to be
ramets detached from a Hydrocotyle mat are able palatable to herbivores include (i) H. asiatica (syn.
to be transported by water to colonize another site for Centella asiatica), which is a common salad
(e.g. H. ranunculoides, H. verticillata, H. vulgaris). vegetable in Asia, (ii) H. mexicana, which is a
In the terrestrial sand dune environment, H. common component of a number of grasshopper
bonariensis is able to dominate a patchy species of the genus Rhachicreagra and (iii) H.
environment by the transportation of resources laxiflora, which appears to be eaten by the grey
(such as nutrients, photosynthates and water) from kangaroo Macropus giganteus.
a part of the clone in nonlimiting conditions to a
Despite the detrimental effect the extensive mats
part of the clone in stressful conditions. It is
of H. ranunculoides are claimed to have on the
possible that clonal integration also occurs within
Canning River environment (such as reduced
the extensive rhizomatous mats of H.
oxygen levels, reduced recreational opportunities,
ranunculoides. From a management perspective,
increased erosion and exclusion of native aquatic
clonal integration may assist in the efficient
plant species), this species has also been recognised
translocation of herbicides such that lower rates
for its beneficial characteristics. Both H.
of application may be needed than supposed.
ranunculoides and H. umbellata have been shown

23
------•------
to effectively reduce the nutrient levels of polluted
water, being particularly useful during the winter
=
months when the otherwise highly productive
Eichhornia crassipes is affected by low
temperature. It is therefore noteworthy that H.
ranunculoides may actually be beneficial to the
Canning River environment with respect to its
ability to reduce the nutrient concentrations of this
eutrophic system, provided the accumulated
biomass is removed from the system to prevent
recycling of nutrients. The resulting biomass from
aquatic plant wastewater systems can then be used
as a cheap source of gaseous fuel (methane).
Furthermore, H. ranunculoides has been shown to
have high digestibility and greater nutritive content
than typical livestock forage plants. Many other
Hydrocotyle species are used by man as medicines
and food sources.
In conclusion, H. ranunculoides appears to possess
a number of typical weed characteristics, including
high growth rates (probably in high nutrient
conditions such as wastewater treatment systems,
the Canning River and certain polluted water
bodies such as the Laguna de la Herrera), effective
vegetative propagation (fragmentation and
possibly clonal integration), plasticity in growth
response (e.g. overwintering) and possibly high
resistance to herbivory. Further studies of the
biology and ecology of H. ranunculoides in the
Canning River are required to achieve successful
long-term control or eradication of this species. In
particular, information which is likely to assist in
devising an effective management strategy include
the determination of successional status, the
relative importance of asexual and sexual
reproduction on population growth, possible
ecotypic variation (as shown by H. bonariensis
ecotypes), and the effect of environmental
parameters such as salinity, temperature and
nutrients on growth, survival and reproduction of
H. ranunculoides.

24
------1
= ------
=
12. REFERENCES

Agami, M., and K. R. Reddy. 1991. Interrelationships between Eichhornia crassipes and Hydrocotyle
umbellata. Aquatic Botany 39: 147-157.
Agnew, A. D. Q. 1974. Upland Kenya Wildflowers: a flora of the ferns and herbaceous flowering plands
of upland Kenya. London: Oxford Univ. Press.

Anderson, L. W. J. 1990. Aquatic weed problems and management in western United States and Canada.
Aquatic Weeds: the Ecology and Management of Nuisance Aquatic Vegetation,, A. H. Pieterse, and K. J.
Murphy (eds.), pp 371-391. Oxford: Oxford Univ. Press.

Anderson, L. C. 1983. Hydrocotyle bowlesioides in Georgia - new to United States. Castanea 48: 317.

Armstrong, W. 1964. Oxygen diffusion from the roots of some British bog plants. Nature 204: 801-802.

Asakawa, Y., R. Matsuda, and T. Takemoto. 1982. Mono- and sesquiterpernoids from Hydrocotyle and
Centella species. Phytochemistry 10: 2590-2592.
Aulbach-Smith, C. A., S. J. de Kozlowski, and L.A. Dyke. 1990. Aquatic and Wetland Plants of South
Carolina. South Carolina Aquatic Plant Management Council.

Auld, B. A., and R. W. Medd. 1987. Weeds: an Illustrated Botanical Guide to the Weeds of Australia.
Melbourne: Inkata Press.

Barbour, M. G., T. M. De Jong, and B. M. Pavlik. 1985. Marine beach and dune plant communities.
Physiological Ecology ofNorthAmerican Plant Communities,, B. F. Chabot, and H. A. Mooney (eds.), pp.
296-322. New York: Chapman and Hall.
Beadle, I. C. 1932. The waters of some East African Lakes in relation to their flora and fauna. J. Linn. Soc.
Lond. Zoo!. 38: 157-211. (Cited from Harper, 1992).

Beadle, N. C. W., 0. D. Evans, R. C. Carolin, and M. D. Tindale. 1982. Flora of the Sydney Region. NSW:
Reed.
Bell, C. R. 1971. Breeding systems and floral biology of the U mbelliferae or evidence for specialisation in
unspecialised flowers., V. H. Heywood (ed.), pp 93-108. London: Academic Press.

Black, J.M. 1957. Flora of South Australia. Adelaide: Govt. Printer.

Boyd, C. E., and D.R. Bayne. 1988. Use ofWater Hyacinths (Eichhornia crassipes) and Water Pennywort
(Hydrocotyle ranunculoides) in Treatment of Poultry Wastewater. Alabama: Dept. Fisheries and Allied
Aquacultures, Auburn Univ.

Burbidge, N. T., and M. Gray. 1970. Flora of the Australian Capital Territory. Canberra: Aust. Nat. Univ.
Press.

Carter, G. S. 1955. The Papyrus Swamps of Uganda. Cambridge: W. Heffer & Sons. (Cited from Denny,
1973).

Cho, E. H., and I. H. Kim. 1988. Studies on the pharmaco-constituents of Hydrocotyle japonica. Journal
of the Pharmaceutical Society of Korea 32: 281-286.

Clough, K. S., T.A. DeBusk, andK. R. Reddy. 1987. Model water hyacinth andpennywort systems for the
secondary treatment of domestic water. Aquatic P !ants for Water Treatment and Resource Recovery,, K. R.
Reddy, and W. H. Smith (eds.), 775-781. Florida: Magnolia Publ.
Constance, L., and M. 0. Dillon. 1990. A new peltate Hydrocotyle (Umbelliferae) from Northern Peru.
Brittonia 42: 257-259.

25
------•------
Cook, C. D. K. 1990. Aquatic Plant Book. The Hague, Netherlands: SPB Academic.

Cook, D. K., B. J. Gut, E. M. Rix, J. Schneller, and M. Seitz. 1974. Water Plants of the World: a manual for
the identification of the genera offreshwater macrophytes. The Hague: Dr. W. Junk Publ.
Costa, C. S. B., and U. Seeliger. 1988. Leaf demography of Hydrocotyle bonariensis, arhizomatous perennial
herb, in coastal dunes of Rio Grande do Sul, Brazil. Revista Brasileira de Biolog_ia 48: 443-452.

Crites, R. W. 1979. Economics of aquatic treatment systems. Aquaculture Systems for Wastewater Treatment:
Seminar Proceedings and Engineering Assessment,, R. Bastian, and S. Reed (eds.), pp 475-485. Washington,
DC: USA Environmental Protection Agency.
Crowden, R. K., J.B. Harbourne, and V. H. Heywood. 1969. Chemosystematics of the Umbelliferae: a
general survey. Phytochemistry 8: 1963-1984.

DeBusk, T. A., and K. R. Reddy. 1987. Wastewater treatment using floating aquatic macrophytes:
contaminant removal processes and management strategies. Aquatic Plants for Water Treatment and
Resource Recovery,, K. R. Reddy, and W. H. Smith (eds.), pp. 643-656. Florida: Magnolia Publ.

DeLoach, C. J., A. D. DeLoach, and H. A. Cordo. 1976. Neohydronomus pulchellus, a weevil attacking
Pistia stratiotes in South America: biology and host specificity. Annals of the Entomological Society of
America 69: 830-834.
Denny, P. 1985. Submerged and floating-leaved aquatic macrophyts (euhydrophytes). The Ecology and
Management of African Wetland Vegetation,, P. Denny (ed.). Geobotany 6, The Netherlands: Junk.

_ . 1973. Lakes of south-western Uganda. IL Vegetation studies on Lake Bunyonyi. Freshwater Biology
3: 123-135.
_ . 1972. Lakes of south-western Uganda. 1. Physical and chemical studies on Lake Bunyonyi. Freshwat.
Biol. 2: 143-158. (Cited from Denny, 1973).

Dierberg, F. E., N. A. Goulet, and T. A. DeBusk. 1987a. Removal of two chlorinated compounds from
secondary domestic effluent by a thin film technique. J. Environ. Qua/. 16: 321-324.
Dierberg, F. E., T. A. DeBusk, and N. A. Goulet.1987b. Removal of copper and lead using a thin film
technique. Aquatic Plants for Water Treatment and Resource Recovery,, K. R. Reddy, and W. R. Smith
(eds.). Florida: Magnolia Publ.
Dinges, R. 1982. Natural System for Water Pollution Control. New York: Reinhold.

Drude, 0. 1897. Umbelliferae. Die naturlichen Planzenfamilien 3: 63-150. (Cited from Tseng, 1967).

Eichler, H. 1987a. Nomenclatural and bibliographical survey of Hydrocotyle L. (Apiaceae). Part I. Feddes
Repertorium 98: 1-51.

_ . 1987b. Nomenclatural and bibliographical survey of Hydrocotyle L. (Apiaceae). Part II. Feddes
Repertorium 98: 145-196.

Evans, J. P. 1992. The effect of local resource availability and clonal integration on ramet functional
morphology in Hydrocotyle bonariensis. Oecologia 89: 265-276.

_ . 1991. The effect of resource integration on fitness related traits in a clonal dune perennial, Hydrocotyle
bonariensis. Oecologia 86: 268-275.

_ . 1988. Nitrogen translocation in a clonal dune perennial, Hydrocotyle bonariensis. Oecologia 77: 64-
68.

26
------•------ ==
=
Evans, J.P., and S. Whitney. 1992. Clonal integration across a salt gradient by a nonhalophyte, Hydrocotyle
bonariensis (Apiaceae). American Journal of Botany 79: 1344-1347.
Everson, R. G., and C.R. Slack. 1968. Distribution of carbonic anhydrase in relation to the C-4 pathway of
photosynthesis. Phytochemistry 7: 581-584.
Ewart, A. J. 1930. Flora of Victoria. Melbourne: Govt. Printer.

Ewart,A. J., and 0. B. Davies. 1917. The Flora of the Northern Territory. Darwin: Mccarron & Bird.
Fedorov, A. 1974. Chromosome Numbers of Flowering Plants. Koenigstein: Otto Koeltz Science.
Fernandez, 0. A., D. L. Sutton, V. H. Lallana, M. R. Sabbatini, and J. H. Iriogoyen. 1990. Aquatic weed
problems and management in South and Central America. Aquatic Weeds: the Ecology and Management
of Nuisance Aquatic Vegetation,, A.H. Pieterse, and K. J. Murphy (eds.), pp 406-425. Oxford: Oxford
Univ. Press.
Foreman, D. B., and N. G. Walsh. 1993. Flora of Victoria. Vol. I: Introduction. Melbourne: Inkata.
French, D. H. 1971. Ethnobotany of the Umbelliferae. The Biology and Chemistry of the Umbelliferae,, V.
H. Heywood (ed.), pp. 385-412. London: Academic Press.
Gaudet, J. J. 1977. Natural drawdown on Lake Naivasha, Kenya, and the formation of papyrus swamp.
Aquatic Botany 3: 1-47.
Gibson, D. J. 1988. The maintenance of plant and soil heterogeneity in dune grassland. J. Ecol. 76: 497-
508.
Godwin, H. 1984. History of the British Flora. Cambridge: Cambridge Univ. Press.
Grime, J.P. 1979. Plant Strategies and Vegetation Processes. Chichester: Wiley.
Grime, J.P., J. G. Hodgson, and R. Hunt. 1988. Comparative Plant Ecology: a Functional Approach to
Common British Species. London: Unwin Hyman.

Harbourne, J. B. 1971. Flavenoid and phenylpropanoid patterns in the Umbelliferae. The Biology and
Chemistry of the Umbelliferae,, V. H. Heywood (ed.), pp. 293-314. London: Academic Press.

Harper, D. M. 1992. The ecological relationships of aquatic plants at Lake Naivasha, Kenya. Hydrobiologia
232: 65-71.
_ . 1984. Recent changes in the ecology of Lake Naivasha, Kenya. Verh. int. Ver. Theoret. Angewandlte
Limnol. 22: 1192-1197. (Cited from Harper, 1992).
Hickey, M., and C. King. 1988. One Hundred Families ofFlowering Plants, 2nd ed. Cambridge: Cambridge
University Press.
Ina, H., A. Asai, H. Iida, and T. U shida. 1987. Chemical investigation of Hydrocotyle sibthorpioides.
Planta Medica 53: 228.
Jessop, J. 1981. Flora of Central Australia. Sydney: Reed.
Johnson, P., and P. Brooke. 1989. Wetland Plants in New Zealand. Wellington: D.S.I.R.

Kachi, N., and T. Hirose. 1983. Limiting nutrients for plant growth in coastal sand dune soils. J. Ecol. 71:
937-944.
Kaplan, K. 1990. The sundew and marsh pennywort community in artificial ponds of lower Saxony and
Westphalia (Germany). Drosera 90: 23-28.

27
------•------ =
=
Klemm, V. V., N. L. Siemon, and R. J. Ruiz-Avila. 1993. Hydrocotyle ranunculoides: a control strategy for
the Canning River Regional Park. Swan River Trust Report 6.

Lima, A., and R. C. Pereira. 1985. Evaluation of herbicide efficiency in a mature rubber plantation in the
south of Bahia. Communiado Tecnico No. 43, Cetro Nacional de Pesquisa de Seringueira e Dende.

Linblad, R., and B. Staul. 1989. Notes on Apium inudatum (Apiaceae) Baldellia ranunculoides
(Alismataceae) in Northern Halland, southwestern Sweden. Svensk Botanisk Tidskrift 83: 211-217.

Lind, E. M., and M. E. S. Morrison. 1974. East African Vegetation. London: Longman.

Lock, J.M. 1973. The aquatic vegetation of Lake George, Uganda. Phytocoenologia 1: 250-262.

Mansor, P. 1988. Traditional salad vegetables of Malaysia. Teknologi Sayur Sayuran 4: 1-5.

Marchant, N. G., J. R. Wheeler, B. L. Rye, E. M. Bennett, N. S. Lander, and T. D. Macfarlane. 1987. Flora
of the Perth Region. Part 1. Perth: WA Herbarium.
Mason, H. L. 1957. A Flora of the Marshes of California. Berkeley: University of California Press.

Mathius, M. E. 1971. Systematic survey of New World Umbelliferae. The Biology and Chemistry of the
Umbelliferae,, V. H. Heywood (ed.), pp 13-30. London: Academic Press.
_ . 1936. The genus Hydrocotyle in northern South America. Brittonia 2: 201-237.

Mitchell, D.S. 1980. Aquatic weeds. An Ecological Basis/or Water Resource Management,, W. D. Williams
(ed.), 81-89. Canberra: Australian National Univ. Press.

Mitchell, D. S., and K. H. Bowmer. 1990. Aquatic weed problems and management in Australasia. Aquatic
Weeds: the Ecology and Management of Nuisance Aquatic Vegetation,, A. H. Pieterse and K. J. Murphy
(eds.), pp 355-363. Oxford: Oxford Univ. Press.

Monteferrante, F. J. 1987. Plant water relations of four coastal Louisiana dune and swale species. Dissertation
Abstracts International, B. Science and Engineering 47: 9.

Moore, D. M. 1971. Chromosome studies in the Umbelliferae. The Biology and Chemistry of the
Umbelliferae,, V. H. Heywood (ed.), pp. 233-255. London: Academic Press.
Moorhead, K. K., and K. R. Reddy. 1988. Oxygen transport through selected aquatic macrophytes. J.
Environ. Qua!. 17: 138-172.

Morisset, R., N. G. Cote, J.C. Panisset, L. Jemni, P Camirand, and A. Brodeur. 1987. Evaluation of the
healing activity of Hydrocotyle tincture in the treatment of wounds. Phytother. Res. 1 (117-121):

Muenscher, W. C. 1944. Aquatic Plants of the United States. New York: Comstock Publ.

Murata, G. 1973. New or interesting plants from Southeast Asia. I. Hydrocotyle chiangdaoensis. Acta
Phytotaxonom. Geobot. 25: 97-106.

Neave, H. M., and M. T. Tanton. 1989. The effects of grazing by kangaroos and rabbits on the vegetation
and the habitat of other fauna in the Tidbinbilla Nature Reserve, Australian Capital Territory. Australian
Wildlife Research 16: 337-351.
Newman, M. J., and W.W. Thomaston. 1979. Aquatic weed control in Georgia. Proc. South. Weed Sci.
Soc. 32: 271-279.
Ohwi, J. 1965. Flora of Japan. Washington: Smithsonian Institute.

Olsen, G., and E. Skitmore. 1991. State of the Rivers ofthe South West Drainage Division. Western Australian

28
------•------
Water Resources Council.
=
=
Overdieck, D., and B. R. Strain. 1981. Effects of atmospheric humidity on net photosynthesis, transpiration,
and stomata! resistance: Hydrocotyle umbellata andHydrocotyle bonariensis. Int. J. Biometeor. 25: 29-38.

Palma, B., C. San-Martin, M. Rosales, L. Zuniga, and C. Ramirez. 1987. Spatial distribution of the aquatic
flora and marsh vegetation in the Marga-Marga Estero in central Chile. Ana/es def Instituto de Ciencias
de/ Mar Y Limnologia Universidad Nacional Autonoma.de Mexico 14: 125-132.

Parsons, W. T., and E. G. Cuthbertson. 1992. Noxious Weeds of Australia. Melbourne: Inkata Press.

Rao, M. N. A., G. K. Patnaik, and R. P. Rastogi. 1979. Spasmolytic principle of Hydrocotyle podantha.
Indian Drugs 16: 185-187.
Reddy, K. R. 1988. Water hyacinth (Eichhornia crassipes) biomass cropping system. I. Production. Methane
from Biomass: a Systems Approach,, W. H. Smith, and J. R. Frank (eds.). London: Elsevier.

_ . 1984. Water hyacinth (Eichornia crassipes) biomass production in Florida. Biomass 6: 167-181.
_ . 1983. Fate of nitrogen and phosphorus in a wastewater retention reservoir containing aquatic
macrophytes. J. Environ. Qua/. 12: 137-141.

Reddy, K. R., E. M. D 'Angelo, and T. A. DeBusk. 1989. Oxygen transport through aquatic macrophytes:
the role in wastewater treatment. J. Environ. Qua/. 19: 261-267.

Reddy, K. R., and W. H. Smith. 1987. Aquatic Plants for Water Treatment and Resource Recovery. Florida:
Magnolia Publ.

Reddy, K. R., and W. F. DeBusk. 1984. Growth characteristics of aquatic macrophytes cultured in nutrient-
enriched water: I. Water Hyacinth, Water Lettuce and Pennywort. Economic Botany 38: 229-239.

Rejmankova, E. 1992. Ecology of creeping rnacrophytes with special reference to Ludwigia peploides.
Aquatic Botany 43: 283-299.

_ . 1991. Vegetation and Water Chemistry of Oso Flaco Lakes. Report for State of California Dept. Parks
and Recreation, Sacramento. (Cited from Rejmankova, 1992).

Richard, A. 1820. Monographie du Genre Hydrocotyle. Ann. Sci. Phys. 4: 145-225. (Cited from Mathius,
1936).

Ridley, H. N. 1930. The Dispersal of Plants Throughout the World. Ashford: Reeve. (Cited from Grime et
al., 1988).

Rodriguez, R. L. 1971. The relationships of the Umbellales. The Biology and Chemistry of the Umbelliferae,,
V. H. Heywood (ed.), pp 63-91. London: Academic Press.

Rodway, L. 1903. The Tasmanian Flora. Hobart: Govt. Printer.

Rowell, C. H. F. 1985. The feeding biology of a species-rich genus of rainforest grasshoppers

(Rhachicreagra, Orthoptera, Acrididae). Oecologia 68: 87-98.

Santos, M. B. L., and J. R. de Freitas. 1988. Preference of Biomphalaria tenagophila among macrophytes
and their periphytons determined through the degree of attractiveness. Revista do Instituto de Medicina
Tropical de Sao Paulo 30: 264-269.
Saville, G. W. 1987. Turf weed control with clopyralid herbicide. Proceedings of the New Zealand Weed
and Pest Control Conference,

Sculthorpe, C. D. 1967. The Biology of Aquatic Vascular Plants. London: Edward Arnold.

29
------•------
= =
Sebacher, D. I., R. C. Harriss, and K. B. Bartlett. 1985. Methane emissions to the atmosphere through
aquatic plants. J. Environ. Qua!. 14: 40-46.

Shaffer, G. P., C. E. Sasser, J. G. Gosselink, and M. Rejmanek. 1992. Vegetation dynamics in the emerging
Atchafalaya Delta, Louisiana, USA. Journal of Ecology 80: 677-687.

Shevchic, V. L. 1986. New habitat of Hydrocotyle vulgaris (Apiaceae) in the European part of the USSR.
Bot. Zh. 71: 1271-1277.
Shigematsu, N., I. Kouno, and N. Kawano. 1982. Quercetin from Hydrocotyle sibthorpioides.
Phytochemistry 21: 2156-2158.
Silander, J. A. 1985. Microevolution in clonal plants. The Population Biology and Evolution of Clonal
Organisms,, J.B. G. Jackson, L. W. Buss, and R. E. Cook (eds.), pp. 107-152. Connecticut: Yale Univ.
Press.

Slaff, M., and J. D. Haefner. 1985. Seasonal and spatial distribution of Mansonia dyari, Mansonia titillans
and Coquillettidia perturbans (Diptera: Culicidae) in the central Florida, USA, phosphate region. Journal
of Medical Entomology 22: 624-629.
Steubing, L., C. Ramirez, and M. Alberdi. 1980. Energy content of water- and bog-plant associations in
the region of Valdivia (Chile). Vegetatio 43: 153-161.

Steward, K. K. 1990. Aquatic weed problems and management in the eastern United States. Aquatic Weeds:
the Ecology and Management ofNuisance Aquatic Vegetation,, A.H. Pieterse, and K. J. Murphy (eds.), pp
391-405. Oxford: Oxford Univ. Press.

Suhail, A., M. H. Ansari, M. Ahmad, and S. M. Osman. 1987. Studies on minor seed oils. XII. Fett.
Wissenschaft-Technologie 89: 154-156.
Swarbrick, J. T. 1973. A Working List ofAustralian Weeds. Lawes: QLD Ag. College. (Cited from Mitchell,
1978).

Sykes, M. T., and J.B. Wilson. 1990a. An experimental investigation into the response of New Zealand
sand dune species to different depths of burial by sand. ActaBotanicaNeerlandica 39: 171-181.

_ . 1990b. Dark tolerance in plants of dunes. Functional Ecology 4: 799-805.

Tarver, D. P. 1985. Status report on Sonar as an aquatic herbicide. Proceedings of the Southern Weed
Society,
Tchobanglous, G., and G. Culp. 1980. Wetland systems for wastewater treatment: an engineering assessment.
Aquaculture Systems for Wastewater Treatment: an Engineering Assessment,, S. Reed, and R. Bastian
(eds.), pp 13-42. Washington, DC: US Environmental Protection Agency.

Thompson, A., and R. Dodson. 1963. Geology of the Naivasha Area. Geological Survey of Kenya, Report
No. 55,. Nairobi: Govt. Printer. (Cited from Gaudet, 1977).
Thompson, K. 1985. Emergent plants of permanent and seasonally-flooded wetlands. The Ecology and
Management of African Wetland Vegetation,, P. Denny (ed.), pp. 43-107. The Netherlands: Junk.
Tomei, P. E., C. L. Cioni, and I. Morelli. 1989. Geobotanical and phytochemical comments on the subject
of Hydrocotyle vulgaris, Hydrocotyle ranunculoides and Centella asiatica. Plantes Medicinales et
Phytotherapie 23: 38-45.
Townsend, C. C. 1989. Flora of Tropical East Africa: Umbelliferae. Rotterdam: Balkema.

30
------•------
-=
Tseng, C. C. 1967. Anatomical studies of flower and fruit in the Hydrocotyloideae (Umbelliferae). Univ.
Calif. Pub!. Bot. 42: 1-58.
Wade, P. M. 1990. General biology and ecology of aquatic weeds. Aquatic Weeds: the ecology and
management of nuisance aquatic vegetation,, A.H. Pieterse, and K. J. Murphy (eds.). Oxford: Oxford
Univ. Press.

Wakefield, W. A. 1951. Notes on some Australian species of Hydrocotyle. Victorian Naturalist 68: 7-9.

Weaver, C. I., and R. G. Wetzel. 1980. Carbonic anhydrase levels and internal lacunar carbon dioxide
concentrations in aquatic macrophytes. Aquatic Botany 8: 173-186.

Webb, C. J. New Zealand species of Hydrocotyle (Apiaceae) naturalised in Britain and Ireland. 18 93-95.
Webb, C. J., and P. N. Johnson. 1982. Hydrocotyle (Umbelliferae) in New Zealand: a three-foliate species.
New Zealand Journal of Botany 20: 163-168.
Webb, C. J., W.R. Sykes, and P. J. Gamock-Jones. 1988. Flora of New Zealand. Vol. 14. Naturalised
Pteridophytes, Gymnosperms, Dicotyledons. Christchurch: D.S.I.R.
Webb, D. H., and W. M. Dennis. 1981. Additions to the flora of Tennessee: Najas gracillima, Cyperus
difformis, Eleocharis tortilis, Hyrdocotyle umbellata. Sida Contibutions to Botany 9: 184-185.
Welsh, S. L., N. D. Atwood, S. Goodrich, and L. C. Higgins. 1987. A Utah Flora. Utah: Birmingham
Young Univ.

31
Appendix 1. Occurence of Hydrocotyle species in Australasia. (ACT=Australian Capital Territory;
NSW=New South Wales; Vic=Victoria; SA=South Australia; QLD=Queensland; Tas=Tasmania;
NT=Northem Territory; WA=Westem Australia; NZ=New Zealand).

SPEOES LOCATION CO:M:MENTS REFERENCE

H. acutiloba southern NSW & Wakefield (1951)
eastern Vic
Sydney widespread Beadle et al. (1982)
H. ala ta WA winter-wet depressions Marchant et al. (1987)
H. asiatica Throughout Australia in or near water Black (1957)
(ind. SA) & NZ; Asia;
Africa; America
Throughout NZ, very common in swamps Rodway (1903)
Australia, & most warm
parts of world
Vic; Australasia medicinal, narcotic Ewart (1930)
H. blepharocarpa WA near-coastal sands & limestones Marchant et al. (1987)
H. boruiriensis NSW & SA maritime habitats; sandy soil Auld & Medd (1987)
lawns
Sydney sandy soils near beach; common; Beadle et al. (1982)
introd. from South America
WA naturalised in disturbed sandy soil Marchant et al. (1987)
at Bunbury; native to Nth & Sth
America
H. callicarpa Southern Australia Rodway (1903)
temperate Australia, Ewart (1930); Black
incl. SA (1957)
WA; Vic; Tas; NSW moist soil Marchant et al. (1987)
H. capillaris Tas; Vic; SA; WA Rodway (1903)
temperate Australia Ewart (1930); Black
I (1957)
H. comocarpa Kangaroo Is. Black (1957)
H. crassiuscula Kangaroo Is. Black (1957)
H. diantha SA; WA wet soil Black (1957)
WA sandy or granitic soils Marchant et al. (1987)
H. dissecta NZ endemic; forest margins and river Webb et al. (1988)
terraces
H. elongata NZ endemic; wet areas of forest Webb et al. (1988)
margins or clearings
H. geranifolia Sydney widespread; sheltered places Beadle et al. (1982)
Vic; NSW Ewart (1930)
H. vammatocarpa NT Ewart & Davies (1917)
H. heteromeria NZ endemic;damporshadedlawns Johnson & Bro~)k (1989)
NZ endemic; lawn weed; damp shady Webb et al. (1988)
places
H. hirta Tas, Vic, NSW, SA, WA Wakefield (1951)
Tas; extra-tropical very_ common Rodway (1903)
Australia
Vic; temperate moist places Ewart (1930); Black
Australia; QLD (1957)
H. hispidula WA sandy soils Marchant et al. (1987)
H. hydrophila NZ endemic; lake, streams, swamp Webb & Johnson (1982);
margins Johnson & Brooke (1989);
Webb et al. (1988)
H. intertexta WA endemic; King George's Sound & Wakefield (1951)
Pemberton
 H. /axiflora ACT; widespread in SE common in pastures, waste ground, Burbidge & Gray (1970);
Australia & Tas woodland & dry forest Auld & Mcdd (1987)
Vic dominant understorey species of dry Foreman & Walsh
sclerophyll forest (1993)
Vic; QLD; NSW; SA; weed of damp pastures; foul smell Ewart (1930); Black
(1957); Willis et al.
(1975); Beadle et al.
0982)
H. lcmnoides WA apparently endemic to Perth Marchant et al. (1987)
region; swamps; rarely collected
H. medicaginoides Vic; SA; WA Ewart (1930); Black
(1957)
H. microphylla NZ endemic; damp open or shady Johnson & Brook (1989);
places; lawn weed in wetter areas Webb et al. (1988)
H. moschata NZ endemic; forest margins & Webb et al. (1988)
clearings; lawn weed
H. mucosa Tas, southern Vic, SA Wakefield (1951)
1--·
H. novac-zelandiae NZ endemic; damp tussockland, Johnson & Brooke (1989);
grassland, sedge and swamp; Webb et al. (1988)
sometimes submerged
1--I. peduncularis ACT, QLD to Tas wet ground under shrubs Burbidge &Gray (1970)
Sydney widespread Beadle et al. (1982)
Tas, Vic, NSW, SA, Wakefield (1951)
QLD
H. pilifera SA; WA Black (1957)
WA moist soils Marchant et al. (1987)
H. plebeia SA;WA in water Black (1957)
H. pterocarpa NZ indigenous to NZ & Australia; Webb et al. (1988)
swamps, lake margins & stream
edges
Tas; Vic; SA Radway (1903)
I
Vic; SA; Tas; NZ in water Ewart (1930); Black
(1957)
H. ranunculoides WA naturalised in permanent fresh Marchant et al. (1987)
water streams; native to Europe
H. ru~losa SA; WA Black (1957)
H. sulcata NZ endemic; tussock grassland, Webb & Johnson (1982);
swamps, lakes and stream edges Johnson & Brooke (1989);
Webb et al. (1988)
H. tetraKonocarpa WA occurs on sands Marchant et al. (1987)
H. trach1/carpa Central Australia Jessop (1981)
H. tripartita Eastern Australia common in moist places Rodway (1903); Black
(1957)
NZ naturalised (Australian) Webb & Johnson (1982);
Johnson & Brooke (1989);
Webb et al. (1988)
QLD to Vic Wakefield (1951)
Sydney widespread; margins of streams; Beadle et al. (1982)
sheltered places; lawn weed
Vic; QLD; NSW; SA Ewart (1930)
H. vulgaris most of Australia, incl. near water Black (1957)
SA; Europe; Africa
Sydney un::orrun:n Beadle et al. (1982)
Vic; temperate poisonous Ewart (1930)
Australia, except
Tasmania
Appendix 2. The occurrence of Hydrocotyle species throughout the world.

SPECIES LOCATION COMJ\fENTS REFERENCE

H. aconitifolia Colombia; Venezuela Constance & Dillon
(1990)
South America Mathius (1936)
H. acuminata South America Mathius (1936)
H. acutifolia South America Mathius (1936)
H. alchemilloides South America Mathius (1936)
H. boliviana South America Mathius (1936)
H. bonariensis Louisiana coastal dunes Monteferrante (1987)
North Carolina common coastal Evans (1988)
(USA) species
Rio Grande do Sul, dominant species of Costa & Seeliger
Brazil coastal sand dunes (1988)
South America irrigation ditch Mathius (1936)
banks; along shaded
stream banks; moist
soil
H. bor.plandii South America Mathius (1936)
H. bonplandii var. glabra South America Mathius (1936)
H. bonplandii var. South America Mathius (1936)
hirtipes
H. bowlesioides Georgia, USA; Costa moist loamy soil Anderson (1983)
Rica; Paraguay; under partial shade;
Brazil similar to H.
sibthorpioides
H. chiangdaoensis ,South-East Asia Murata (1973)
H. dahlgreni South America moist sandy banks Mathius (1936)
H. dichondroides Japan Ohwi (1965)
H. fjlipes South America Mathius (1936)
H. geraniodes South America Mathius (1936)
H. globiflora Peru Constance & Dillon
(1990)
South America Mathius (1936)
H. vacilis South America Mathius (1936)
H. grossulariaefolia South America Mathius (1936)
H. gunnerifolia South America Mathius (1936)
H. hederacea South America Mathius (1936)
H. heucheraefolia South America Mathius (1936)
H. hexagona Ecuador Constance & Dillon
(1990)
H. hexagona South America Mathius (1936)
H. hirta Chile emergent life form; Steubing et al. (1980).
associated with
Juncetum procerii
H. hitchcockii South America moist ground Mathius (1936)
H. humboldtii Colombia; Ecuador; Constance & Dillon
Peru (1990)
South America ravines Mathius (1936)
H. incrassata South America Mathius (1936)
H. japonica Korea Cho & Kirn (1988)
H. javanica Japan; Tropics of woods in low Ohwi (1965)
Asia mountains
H. langsdorf Brazil Constance & Dillon
(1990)
H. Iehmannii Columbia Constance & Dillon
(1990)
South America Mathius (1936)
H. leucocephala South America Mathius (1936)
H. lon~ipes South America Mathius (1936)
H. mannii East Africa woodland, near Lind & Morrison
mountain lakes (1974)
Uganda, tropical occurs in swamp (1 % Denny (1973)
East Africa cover) with H.
ranunculoides
(insignificant %
cover)
upland Kenya quite common on the Agnew (1974)
forest floor in &
below the bamboo
zone
H. maritima Japan; Korea; China; lawns & roadsides in Ohwi (1965)
Bonins lowlands & hills;
cnrran:m
·H. mexicana South America Mathius (1936)
H. minutifolia South America Mathius (1936)
H. monticola upland Kenya muddy & peaty Agnew (1974)
stream banks in the
upper forest levels
into the alpine zone
H. multifida South America Mathius (1936)
H. palmata South America Mathius (1936)
H. 'T!ennellii South America along stream in Mathius (1936)
clearing
H. peruviana South America Mathius (1936)
H. pusilla South America Mathius (1936)
H. quinqueloba Brazil; Peru Constance & Dillon
(1990)
South America Mathius (1936)
H. ramiflora Japan; Korea lawns & sunny places Ohwi (1965)
in lowlands & low
mountains
H. rotundifolia South America damp ground Mathius (1936)
H. sagasteguii Peru Constance & Dillon
(1990)
H. sibthorpioides Japan; widely shaded places in the Ohwi (1965)
distributed in lowlands; common
tropical & warm-
temperate Asia
Japan rornmon weed Shigematsu et al.
(1982)
Japan medicinal Ina et al. (1987)
H. sphenoloba South America Mathius (1936)
H. tambalomaensis South America Mathius (1936)
H. tenerrima South America Mathius (1936)
H. umbel la ta Nova Scotia, Canada rare, coastal plain Roland (1991)
South America street gutters; Mathius (1936)
pastures; edge of
pools; meadows
Tennessee Webb & Dennis
(1981)
throughout edges of ponds, Mason (1957)
California, east to ditches, marshy
.Atlantic, south to grouoo
Mexico; southern
Africa
H. urbaniana South America Mathius (1936)
H. venesuelensis South America· Mathius (1936)
H. verticil lat a California aquatic; commonly Rejrnankova (1992)
occur in irrigation
and drainage canals
California; Atlantic Mason (1957)
states, Mexico, West
Indies
Utah moist ground or in Welsh et al. (1987)
water in Washington
Country; S. America
north to
Massachusetts &
California
H. verticillata var. South America Mathius (1936)
racemosa
H. vestita South America Mathius (1936)
H. vulgaris British Isles Godwin (1984)
! British Isles; W, C & Grime et al. (1988)
S Europe northwards
to Scandanavia &
southwards to
Portugal & Grecse; N
Africa; introduced to
NZ
Europe Tomei et al. (1989)
Germany codominant with Kaplan (1990)
Drosera intermedia
Sweden small pools near Lindblad & Stahl
coast; sandy substrate (1989)
Ukraine Zelerichuk (1987)
USSR endan2-ered species Shevchic (1986)
H. yabei Japan woods in hills & Ohwi (1965)
mountains
H. yanghuangensis Ecuador Constance & Dillon
(1990)
South America woods, at base of Mathius (1936)
rotten trunks ·
Hydrocotyle sp. Louisiana, USA coastal deltaic Shaffer et al. (1992)
wetlands
 DISTRIBUTION OF THE GENUS HYDROCOTYLE
THROUGHOUT THE WORLD

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~~---,._. -~ ~ - - ~

BRITISH
ISLE ..
VASCOTIA

"""
CALIFORNIA

0 ·1/ NORTH AFRICA ~

> Bonin Is.
\
..
'••

COLUMBIA-----J-
ECUADOR- ,._ J..
PERU-

,>
ii
\
,o
,,
CHILE---4,_\

SOUTHERN AFRICA

0
_SOUTH AMERICA <:)

......

~
- ---..J'---v--·
~
.,-.JV\. --· -~..----.A-._' - ~ /

..._. ~ 0 . /~ / ---- ~_,-1, 1rf_.-~ /_,--··· /

'',, .)'vv-" ···-·--- ··-

-...... ·-- .
(.(/
·-.__ -- _____ ::>-
Appendix 3. The occurrence of Hydrocotyle ranunculoides throughout the world.

LOCATION COMMENTS REFERENCE

Africa -- Lock (1973); Thompson (1985)
California shallow pools & muddy shores, lower altitudes Mason (1957)
Central America Cook et al. (1974)
Chile one of dominant species (70% cover) of fresh, clear water Steubing et al. (1980)
brooklets, brooks and small rivers in the Central valley &
pacific littoral region of south Chile
Chile (central) codominant with Elodea potamo~eton Palma et al. (1987)
Colombia a dominant species, upt to 75-100% cover Wijninga (1989)
Europe Tomei et al. (1989)
Florida Weaver & Wetzel (1980)
Georgia, USA potential weed Newman & Thomaston (1979)
Kenya (upland), East Africa locally common in ponds & marshes, growing in water Agnew (1974)
between the reeds
Kenya, East Africa a dominant water-edge species colonising the Salvinia Harper (1992)
moles/a (weed) mats; floating community moving in rafts or
islands due to wind action; covering about 25% of lake surface
Kenya, East Africa occurs in shallow water & recently exposed shore of Lake Gaudet (1977)
Naivasha (tropical freshwater lake in closed basin)
northern South America Pennsylvania & Washington south to southern South Mathius (1936)
America at coastal & low elevations; common name:
Somberito de· a~ua
South Carolina restricted to coastal plain Aulbach-Smith & de
Kozlowski (date??)
Tropical East Africa appears to be geocarpic, the peduncle bending over to push Townsend (1989)
the frttlts into the mud
Uganda Denny (1973)
USA in ponds, lakes & margins of streams; mostly on sandy Muenscher (1944)
bottom. Atlantic coast; local in the South & westward.
Distribution map provided
 DISTRIBUTION OF HYDROCOTYLE RANUNCULOIDES
THROUGHOUT THE WORLD

CALIFORNIA SOUTH CAROLINA

io
--E GEORGIA
FLORIDA o•'

.....

COLUMBIA ;c':
0... ....
\•
I

d
,... ' ,
~
.I

CH ILE---+\ PERTH

[7 ✓p
0
'::)