Correspondence: J G Zaller, Institute of Ecology and Conservation Biology, University of Vienna, Althanstrabe 14, A-1090 Vienna, Austria. Tel: (+43)
1 4277 54221; Fax: (+43) 1 4277 9542; E-mail: jgzaller@yahoo.de
an infestation of grassland and arable land by R. crispus physiology (about 20%) and medical use of the two
and R. obtusifolius is also among the most important species (about 15%).
obstacles for farmers to switch from conventional to
organic farming in Central Europe (Dierauer & Stöp-
Biology and Ecology
pler-Zimmer, 1994).
Generally, both species are, to a certain extent, The majority of biological and ecological studies on the
indicators of mismanagement of agricultural land and of two Rumex species have focussed on topics in the field of
high soil nitrogen concentrations, and R. crispus vegetation ecology (including geobotany, growth
additionally indicates soil compaction (e.g. Ellenberg, responses, plant development, species distribution and
1986; Oberdorfer, 1990). Thus, one of the factors most interspecific competition between plants), followed by
closely associated with the occurrence of the two species studies investigating RumexÕ seed ecology (including
in agricultural land includes excessive application of seed production, soil seedbanks, germination, dor-
organic or synthetic nitrogen fertilizers (Haggar, 1980). mancy) and other topics like nutrient concentrations
The two Rumex species are particularly successful as and feeding value of Rumex plants, or Rumex as a host
weeds on agricultural land because of: (i) the ability to for diseases and parasites (Fig. 1). Remarkably few
flower several times a year, (ii) the production of large
number of seeds, which remain viable in the soil for
many years, (iii) their ability to quickly establish from
seeds and (iv) their germination whenever environmental
conditions provide a reasonable probability for the
seedling to survive (Cavers & Harper, 1964). Addition-
ally, these Rumex species show considerable ability to
re-grow from vegetative fragments left in the soil after
cultivation or cutting (Gwynne & Murray, 1985; Pino
et al., 1995).
The aim of this paper is to synthesize the available
literature on the ecology and non-chemical control of
R. crispus and R. obtusifolius based on about 700
references available through international scientific lit-
erature databases (e.g. Agris – http://www.fao.org/agris;
CAB International – http://www.cabi-publishing.org;
ISI Web of Science – http://www.isiknowledge.com)
and additional non-indexed sources (used especially for
publications before 1973). A large body of information
exists on these two species and a few reviews have been
published focussing mainly on their biology and non-
chemical control (Cavers & Harper, 1964; Foster, 1989)
or their seed ecology (Roberts & Totterdell, 1981).
Rather than just re-reviewing the vast literature, the
intention was to briefly summarize and critically evaluate
the published literature in order to pinpoint gaps of
knowledge in Rumex ecology and their non-chemical
control which could be filled by future research.
Of the reviewed publications, about 30% were of
descriptive nature (including reviews and model analy-
ses), about 20% of the papers report data from field
experiments, 20% of the studies stem from greenhouse
experiments and 30% from laboratory experiments. The
number of studies published on the two Rumex species
peaked during 1993–99 with more than 40 publications
appearing per year. Averaged over the past 30 years,
Fig. 1 Ecological topics covered by publications on Rumex crispus
most of the studies considered ecological aspects in a and R. obtusifolius available through scientific literature databases
broad sense (about 70%), followed by papers on (Agris, CAB International, ISI Web of Science; 1973–2003: total
chemical and non-chemical control (about 40%), 528 publications).
studies have addressed effects of soil properties on estimated that about 90% of Rumex seeds are lost
Rumex development and infestation (Fig. 1). because of dispersion, predation, death or decay (Wea-
Both species of Rumex are highly variable and show ver & Cavers, 1979a). Because of this high seed mortality
great phenotypic plasticity; therefore, the characteristics under field conditions, the contribution of the soil
given below can vary substantially depending on the seedbank to the infestion of fields with Rumex species is
environmental conditions. Both Rumex species are probably over-estimated even if some seeds remain
native to Europe and are up to 1.5 m tall perennial viable for about 80 years and accumulate in the soil
herbs, with up to 40 mm thick fleshy taproots which (Darlington & Steinbauer, 1961). Seed size of R. crispus
may extend up to 2.5 m in the soil (Kutschera et al., and R. obtusifolius is between 2.0–3.0 mm in length and
1992). At least for R. obtusifolius, it is known that it 1.0–2.0 mm in width; however, size varies as a conse-
hybridizes freely with many other members of the quence of environmental influences (e.g. predation,
subgenus Rumex, and R. crispus · R. obtusifolius is very position, defoliation) in which the mother plant was
common and widely distributed (Lousley, 1939). growing (Brouwer & Stählin, 1975). Whether seed size
Although no clear data are available on the longevity affects germination of both species is not clear. Small
of Rumex plants, it can be observed that some plants live seeds of R. crispus (Maun & Cavers, 1971) and
for decades. Readers more interested in the general R. obtusifolius (Cideciyan & Malloch, 1982) germinated
biology and ecology of the two species are referred to to higher percentages than large seeds. Lighter seeds of
Cavers and Harper (1964), who give a very thorough R. crispus and R. obtusifolius grew and competed less
description of these species. Neither of the two Rumex successfully than did seeds of normal mass (Cideciyan &
species do occur in higher abundance in native plant Malloch, 1982). Rumex seeds have also been shown to
communities (Ellenberg, 1986), but are definitely stimu- be eaten (Piearce et al., 1994) and vertically distributed
lated and distributed by human activities. in the soil by earthworms (J G Zaller, unpubl. obs.).
Seed viability of R. crispus was tested in a longevity
trial started by Beal in 1879, where R. crispus seeds
Flowering, seed production and soil seedbank
among seeds of other species were mixed with subsoil,
Flowering and seed production can occasionally happen placed in inverted bottles and buried to a depth of 50 cm.
in the seedling year but usually takes place in the second After 50 years, 50% of buried R. crispus seeds germina-
year starting in spring and may continue until a hard ted (Kivilaan & Bandurski, 1981), after 80 years, still 2%
frost in late autumn (Foster, 1989). It is reported that seeds germinated (Darlington & Steinbauer, 1961). In a
both species have a tendency to die after producing seeds similar experiment where seeds were buried at 1 m depth,
(Cavers & Harper, 1964); however, this may not be the germination of R. crispus was 80% after 1 year and 76%
case in all environments. Six days after the end of first after 10 years; germination for R. obtusifolius was 94%
flowering, 15% of the seeds are already viable, and after after 3 years and 83% after 21 years (Toole & Brown,
18 days, >90% of the seeds are able to germinate 1946). The seeds pass the digestive tracts of birds and
(Dierauer & Stöppler-Zimmer, 1994). The flowers have cattle without being harmed but are destroyed when fed
no nectar and are mainly wind pollinated. Fruits can be to chickens (Cooper et al., 1960).
blown by wind for a considerable distance, be carried
floating along with water or be distributed in dung after
Seed germination and dormancy
passing through the digestive system of grazing cattle.
Many fruits probably just fall to the ground and Particularly valuable contributions have been made
germinate in clusters near the parent plant. to the knowledge of the seeds of R. crispus and
The number of seeds set by Rumex is determined by R. obtusifolius by Harper (1965), Williams (1971) and
plant size at flowering and the duration of flowering; Baskin and Baskin (1985). The seeds have also been the
hence estimates vary greatly and range from 100 to choice of many workers for studying the phytochrome
>60 000 per year (Cavers & Harper, 1964). It is best to system (Vicente et al., 1962, 1968; Vergnano Gambi,
assume that even individual sparsely separated plants 1966; Le Deunff & Chaussat, 1968; Le Deunff, 1971).
can potentially add thousands of seeds to the soil There is a general agreement that seed germination of
seedbank. The continued infestation of pastures and both species is promoted by light and alternating
crops by Rumex is, to some extent, dependent on the temperatures. Temperature fluctuations can affect ger-
persistence of viable seeds on and below the soil surface. mination even in the absence of light (Isikawa & Fujii,
Pasture seedbank in the uppermost 15 cm of soil 1961; Kurimoto et al., 1974; Weaver & Cavers, 1979b;
contains about 5 · 106 Rumex seeds m)2 (Hunt & Takaki et al., 1981; Hand et al., 1982; van Assche &
Harkess, 1968); however, it might even be higher in Vanlerberghe, 1989). Under natural conditions, seeds of
pastures with a history of Rumex infestation. It is both Rumex species can germinate throughout the year;
however, there are clear flushes of germination in early available on whether a higher relative growth rate of
spring and early autumn when strong temperature Rumex compared with crop species is dependent on
fluctuations between day and night occur (Roberts & higher nutrient uptake kinetics of Rumex. There are also
Totterdell, 1981). The seedlings of R. obtusifolius are some doubts as to whether the two Rumex spp. are
somewhat slower in germination and early development nitrophilous in all developmental stages. For example,
than those of R. crispus (Cavers & Harper, 1964). soil nitrogen concentrations of microsites where
Generalizations about the germination behaviour of the closely related nitrophilous Rumex alpinus L.
both species have to be made carefully although, as light emerged were not higher than surrounding pasture soil
requirement for germination of buried seeds may (J G Zaller, unpubl. obs.). Generally, field studies on the
fluctuate dramatically with time (Taylorson & Hen- population dynamics of both Rumex species are seri-
dricks, 1972; Froud-Williams et al., 1984; Baskin & ously lacking. For example, it is not known to what
Baskin, 1985) and may be affected by the light extent climatic effects (e.g. drought or rain periods, cold
environment experienced by the seeds before burial winters, etc.) or previous field history contribute to the
(Fenner, 1985). Recently, Boyd and Van Acker (2003) variation in incidence of Rumex from one year to
found that seedling emergence of R. crispus was highest another.
when seeds were placed on the soil surface and soil
moisture was at field capacity. However, when the soil
Root ecology and regeneration
was drier, seedling emergence was similar for different
seeding depths ranging between 1 and 7 cm (Boyd & A thorough knowledge of the root system of Rumex is
Van Acker, 2003). imperative for the development of sustainable control
The time of seed production determines germination strategies against these species. Root growth of both
ability of R. obtusifolius; seeds developed in spring species is greatest in spring and plants which have
showed little germination ability, whereas germination overwintered often develop adventitious roots when
rate of seeds from summer was 95% (Adolf & Linke, highest growth recommences (Cavers & Harper, 1964).
1992). Moreover, seeds from the second regeneration R. crispus could produce new plants from rootstocks
after cutting showed higher germination rates than those after 40 days, and R. obtusifolius were incapable of
from the third regeneration harvested in autumn (Adolf regrowth from rootstocks until 51 days (Monaco &
& Linke, 1992). Cumbo, 1972). Recent results of Pino and colleagues
Seed dormancy of Rumex spp. was reviewed by (1995) show, despite the reservations of other authors
Roberts and Totterdell (1981). The role played by (Cavers & Harper, 1964; Hongo, 1989), that R. obtusi-
dormancy in the germination ecology of R. crispus is not folius does have a true clonal growth system. Therefore,
clear (Baskin & Baskin, 1985). In general, Rumex clonal growth may be the usual regenerative system in
appears to have only short-term innate dormancy dense swards, where germination and seedling establish-
(several weeks to months), dormancy is more commonly ment are prevented and would maintain and increase the
induced for example by soil burial. occupancy of available space. Seed dispersal and seed-
ling establishment would remain the strategy for colon-
izing gaps in the sward and for maintaining genetic
Plant establishment
variability of the population. The main structure
Both species have low competitive abilities as seedlings, involved in clonal growth is the subterranean stem
and cannot become established in closed communities component (i.e. the rhizome) with its ability to produce
(Cavers & Harper, 1964; Weaver & Cavers, 1979a), adventitious roots and new taproots. Almost all 3-year-
additionally there is high seedling mortality over the first old plants develop secondary taproots, and 5-year-old or
winter (Cavers, 1967; Hongo, 1986, 1989). Surprisingly, older plants are usually heavily divided as secondary
no data seem to be available on the rate of germination taproots become the main root system (Pino et al.,
and seedling establishment in dense grassland commu- 1995). The result is a dense population of clones that
nities versus grassland gaps. Establishment of seedlings occupy an increasing area (i.e. phalanx strategy in clonal
and transplants of R. crispus and R. obtusifolius are also growth), a survival strategy common in plants from
thought to be dependent on historical abundance of closed communities (Harper, 1977). Root longevity of
Rumex on specific sites: new seeds or transplants had R. obtusifolius was estimated to be <2 years in fertile
been unable to survive where relics of Rumex popula- soil and >4 years in infertile soil (Cavers & Harper,
tions occurred (Cavers, 1967). 1964), but is most likely much longer.
Especially on soil with high nitrogen content, Rumex Regeneration of Rumex spp. from root fragments has
species can be observed to grow faster than most of been studied and it is now confirmed that regeneration
the other co-occurring species. However, no data are of shoots can take place only from the uppermost part of
the taproot (i.e. botanically the rhizome; Healy, 1953; future growth (Chapin et al., 1990). Mono- and disac-
Hudson, 1955; Pino et al., 1995). Regrowth ability seems charides can make up >50% of the concentration of
related to the previous presence of buds, as fragments total sugars in the roots of R. crispus and R. obtusi-
lacking buds (e.g. parts below the root-collar) have been folius (Kühbauch et al., 1980; Hatcher, 1996) and much
shown to be unable to regenerate (Hughes, 1938; cited in of the increase in root carbohydrate concentrations
Pino et al., 1995). However, there are also studies seems to occur during autumn (Voigtländer et al.,
showing that regeneration is possible from all root 1976). The average nitrate concentration in roots was
parts, ranging from below 10% regeneration from pieces more than six times higher than in above-ground parts
of root end and top parts, to a 90% regeneration of the (Gebauer et al., 1984). Generally, R. obtusifolius nitrate
middle parts of the roots (Adolf & Linke, 1992). It is content fluctuates rapidly as a consequence of nitrate
very likely that these contradicting results are due to a fertilization (Gebauer et al., 1984; Melzer et al., 1984).
confusion between roots and rhizomes which are As the accumulation of nitrate is a short-term process
difficult to distinguish. Root fragment size (Pino et al., only, R. obtusifolius cannot be considered as a nitrate-
1995) and mass (J G Zaller, unpubl. obs.) have been storage plant (Rehder, 1982). The ability of R. obtusi-
shown to be unrelated to regrowth, at least under folius to take up nitrate more efficiently than other
greenhouse conditions. Regenerative ability of Rumex plants is suggested to be responsible for the temporary
roots can additionally be explained by high contents of increase of endogenous nitrate (Melzer et al., 1984). No
certain chemicals (e.g. phenolics) in the roots which mycorrhizal fungi have been shown to be associated
actively suppress the growth of some fungi and bacteria with roots of these Rumex species (Cavers & Harper,
(Kasai et al., 1982) and thus resist decay. 1964). Clearly, if one seeks to find sustainable control
Morphological and anatomical responses of rhizomes strategies against these Rumex species, the enormous
of R. obtusifolius to different depths of burial in the field ability of vegetative regeneration needs to be taken into
showed that the rhizomes regenerated from 5, 10 and account.
20 cm depths (Roberts & Hughes, 1939), but failed to re-
grow when buried 30 cm below surface (Klimes, 1993).
Ecological interactions and global climate change
Burial depth had no effect on the number of internodes
produced during a growing season. It however led to a The two Rumex spp. appear to be vulnerable to
30-fold increased length of internodes with increasing ecological interactions especially in their early stages of
depths and in the case of deeply buried rhizomes, the their development. Once Rumex has produced the
carbohydrate reserves of the last-year segment were taproot it has a great advantage over the shallower-
nearly completely depleted (Klimes, 1993). Root frag- rooted grasses and herbs and is very difficult to eradicate
ment regeneration at different depths were also influ- (Jeangros & Nösberger, 1990; Niggli et al., 1993).
enced by soil moisture levels. Especially under Comparing both Rumex species based on their ecolog-
waterlogged conditions, root fragment regeneration ical indicator values (Ellenberg et al., 1991), R. crispus
was possible from the soil surface, but it was prevented shows less requirement for soil nutrients but a higher
when fragments remained buried (Pino et al., 1995). requirement for soil moisture than R. obtusifolius. Yet
Fragmentation of Rumex roots and stems may easily very few studies have addressed the effects of varying
occur as the result of ploughing or other soil cultivation soil fertility or moisture levels on the development of
methods thus giving rise to clusters of plants where both species. In one study, doubling of nitrogen fertil-
previously there was only one. The root system of ization increased Rumex growth especially when root
hybrids between R. crispus and R. obtusifolius seem to competition from grass species was present (Jeangros &
have slightly greater regenerative ability than the parents Nösberger, 1990).
(Cavers & Harper, 1964). Root regeneration has been Interactions between Rumex spp. and insect herbiv-
found to be dependent not only on abiotic conditions ory have been studied frequently (Bentley & Whittaker,
but also on seasonal control (Hudson, 1955). It has been 1979; Whittaker et al., 1979; Bentley et al., 1980; Smith
assumed that regeneration is possible only during the & Whittaker, 1980a,b; Whittaker, 1982) and will be
early spring (Hudson, 1955; Cavers & Harper, 1964), considered in more detail in a later section as it is more
although regeneration has been observed in greenhouse relevant for the development of biological control
conditions into late autumn (J G Zaller, unpubl. obs.). methods. Herbivory by Gastrophysa viridula Deg.
R. obtusifolius regrowth from rhizome fragments have (Coleoptera, Chrysomelidae) altered the outcome of
also been shown to be altered by competition of the competition between R. crispus and R. obtusifolius in
neighbouring grassland species (Zaller, 2004). grassland (Bentley & Whittaker, 1979) and also inter-
The pool size of reserves stored in roots is probably acted with competition by grasses resulting in a reduced
the best measure of the potential to contribute to R. obtusifolius mass (Cottam et al., 1986). The larvae of
certain weevils (Apion spp.) which live within the stalks was cultivated together with the closely related R. alpinus
of Rumex (Hopkins & Whittaker, 1980) and in turn play in special gardens in order to produce food for pigs or
host to hymenopterous parasites was also studied spinach-like food for humans (Stebler & Schröter, 1892).
(Hopkins et al., 1984). The indigenous people of northern California are said to
Allelopathic effects of R. obtusifolius aqueous shoot have used the seeds of Rumex species to make a cooked
extracts have been shown to inhibit the germination of mush (Holm et al., 1977). R. obtusifolius leaves served as
sunflower (Helianthus annuus L.), soyabean [Glycine max a popular antidote for nettle stings and a tea made
(L.) Merr.; Beres & Kazinczi, 2000] as well as native from the root was formerly used for curing boils
grassland species like Trifolium repens L. (J G Zaller, (Machatschek, 1999).
unpubl. obs.). Residues of R. obtusifolius decreased the In modern agriculture, both Rumex species are
germination of wheat (Triticum aestivum L.), barley undesirable in grasslands and in all types of crops mainly
(Hordeum vulgare L.), sunflower and soyabean by >80% because they decrease yields and reduce forage feeding
(Beres & Kazinczi, 2000), and decreased seedling growth value (Savory & Soper, 1970; Oswald & Haggar, 1976;
of maize (Zea mays L.) and soyabean (Beres & Kazinczi, Marten et al., 1987). Densities of 10 R. obtusifolius plants
2000). However, all these studies have been conducted m)2 or a ground cover of 30% can reduce total yield in a
under controlled conditions and it remains to be shown if perennial ryegrass sward by 30% (Oswald & Haggar,
findings are consistent under field conditions. 1983; Courtney, 1985). Forage nutritive value and
The two Rumex spp. have also been at the centre of palatability of Rumex crispus is, as with most cultivated
investigations on effects of elevated atmospheric CO2 or forages, negatively influenced by maturation and
ozone concentrations; however, there do not seem to be crude protein and in vitro digestibility concentration of
studies involving other factors of global climate change R. crispus declined 13 and 22%, respectively, from the
(e.g. global warming, ultraviolet-B radiation). An early vegetative to seed-set stages of development (Bosworth
study investigated the effects of ozone and smog on et al., 1986). Leaves of both Rumex species have only
Rumex plants and showed that leaves exposed to these 20% less palatability and digestibility than those of
pollutants partially turned red because they had been perennial ryegrass (Lolium perenne L.) (Courtney &
stimulated to form anthocyanin, while untreated plants Johnston, 1978). Compared with lucerne (Medicago
did not form the pigment (Koukal & Dagger, 1967). A sativa L.), R. crispus had similar crude protein concen-
later study found that R. obtusifolius grows less when trations (Marten et al., 1987) and R. obtusifolius showed
fumigated with increased ozone concentrations com- similar nitrogen and crude fibre concentration but higher
pared with plants fumigated with less ozone (Reiling & lignin concentrations (Waghorn & Jones, 1989). Despite
Davison, 1992). Rumex spp. grown at elevated the average nutritive value of the Rumex spp., rumicin
atmospheric CO2 concentrations showed greater root from the above-ground parts and chrysarobin from the
proliferation and increased water-use efficiency roots have been identified as biologically active sub-
(R. obtusifolius; Pearson & Brooks, 1995), reduced stances in the plant which may cause gastric disturbances
nitrogen and oxalate concentrations, increased C:N and dermatitis if large amounts are eaten by cattle (Holm
ratio in leaves (R. obtusifolius; Brooks & Whittaker, et al., 1977). There are also concerns over Rumex species
1998) and increased leaf areas (both Rumex spp.; Salt containing very high levels of nitrate and oxalate in their
et al., 1995) than those grown at ambient CO2. These foliage (Hatcher et al., 1995a; Bohner, 2001), and they
CO2-induced alterations in food quality of Rumex leaves are suspected of being fatal to livestock in some
and stimulated leaf-mining insects (Salt et al., 1995) circumstances (Kingsbury, 1964). However, research in
however, affected the herbivory of insect larvae only New Zealand showed that 10% of R. obtusifolius
little (Brooks & Whittaker, 1998). Because of its material in a portion of grass and lucerne inhibited bloat
reactions to elevated atmospheric CO2, it has also been in cattle, because of a decreased concentration on soluble
suggested that R. obtusifolius expands more than other crude protein in the rumen because of R. obtusifolius
species in an environment with higher CO2 and could (Waghorn & Jones, 1989). Composition of other
thus occupy habitats where nutrient availability is chemicals of both species varies considerably and is not
usually not limiting plant growth (e.g. agricultural soils; very different fom that of the cultivated species (Vengris
Grime, 1996). Clearly, this issue deserves more detailed et al., 1953; Fairbairn & Thomas, 1959).
research.
Non-chemical control in grassland and
Rumex for food and forage arable land
The importance of both species changed dramatically The long-term goal of control measures against the two
during the past 100 years. R. obtusifolius, in particular, Rumex species is to hinder the build-up of seeds and to
weaken their regrowth capacity by removing or destroy- biocontrol agents (Greaves, 1986). The most thoroughly
ing their above- and below-ground biomass. Of the studied organisms for RumexÕ biocontrol are the Co-
c. 250 reviewed studies published over the last 30 years leoptera Gastrophysa viridula Degeer and the rust fungus
which exclusively focus on non-chemical control meth- Uromyces rumicis (Schumach.) G. Winter. Of about 110
ods, contributions on biological control dominated studies published to date on biological control of Rumex
(range between 25 and 67% per year) followed by species, >50% used insects (mainly from the order of
cultural methods (between 22 and 100% per year) and Coleoptera), about a quarter used fungi and only a few
mechanical and direct control methods (between 7 and examined the effects of plant extracts or specific grazing
36% per year; Fig. 2). In particular, publications focus- by goats or sheep to control Rumex species (Table 1).
sing on mechanical measures increased in the last few Only a few researchers combined biological control
years (Fig. 2), which could be explained by the expan- methods in their work (see below).
sion of organic farming and the increased demand for While earlier studies explored mainly the potential of
non-chemical control measures. However, the very few control with Coleoptera in the laboratory, greenhouse or
contributions on mechanical control measures in the field (Bentley & Whittaker, 1979; Naito et al., 1979a,b;
1970s and 1980s, perhaps reflect the expectations in new Hopkins & Whittaker, 1980; Julien et al., 1982), later
chemical herbicides developed against Rumex species research additionally investigated effects on specific
during this period (Fig. 2). plant traits. Coleoptera have been shown to reduce
Rumex seed production (Bentley et al., 1980; DeGrego-
rio et al., 1991), regeneration (DeGregorio et al., 1991),
Biological control
leaf and shoot growth (Cottam et al., 1986) and to alter
Numerous predators and parasites associated with both Rumex cover (Kohout & Kohoutova, 1994; Kohout,
Rumex species have been found; however, only few have 1994). However, in summary, Rumex plants were rarely
a significant damaging effect on their hosts and can be so badly attacked by these beetles that they died out as a
considered for biological control. Cavers and Harper consequence, unless some other agent further weakened
(1964) mention 34 herbivorous insect species which can the plants. Very promising results have recently been
affect both Rumex spp., from collections in Italy; 198 reported from experiments under controlled conditions
insect species have been identified affecting mainly where Coleoptera beetles in clip cages were able to
R. crispus (Spencer et al., 1981). Additionally, around systemically reduce R. obtusifolius leaf area even after
20 fungal species have been identified as potential cages had been removed (Moore et al., 2003). To the
Fig 2. Relative contribution of publications covering non-chemical control methods against Rumex crispus and R. obtusifolius available
through scientific literature databases (Agris, CAB International, ISI Web of Science; 1973–2003: total 253 publications). No studies
on Rumex control were available in 1990.
Control Experimental
organism Control spp. Rumex spp. conditions Main results Reference
Coleoptera Apion miniatum Geom. R.o. Laboratory, field Herbivore regulates R.o. populations Kohout (1994); Kohout and Kohoutova (1994)
by leaf and root consumption
Apion violaceum Kirby R.o. Field Herbivore affects stem height and numbers Hopkins and Whittaker (1980)
Gastrophysa atrocyanea Mots. R.o. Field Herbivore shows potential as biocontrol agent Naito et al. (1979a,b)
Gastrophysa viridula R.c., R.o. Greenhouse R.o. preferred by herbivore over R.c. Bentley and Whittaker (1979)
G. viridula R.c, R.o. Field Herbivory reduced seed number (R.c., R.o.) Bentley et al. (1980)
and seed mass (R.o.)
G. viridula R.c. Field Herbivory reduced growth and survival Whittaker (1982)
and interacted with flooding
G. viridula R.o. Field Herbivory reduced growth only when Cottam et al. (1986)
grown in competition
Hypera rumicis L. R.c. Field Herbivory reduced growth, seed production DeGregorio et al. (1991)
and regeneration
Lixus cribricollis Boh. R.c. Laboratory R.c. food plant for herbivore Julien et al. (1982)
Rust fungus Ramularia rubella (Bonord.) Nannf. R.o. Greenhouse Rust infection decreased leaf area and Huber-Meinicke et al. (1989)
root dry matter content
Uromyces rumicis R.c. Field Rust infection reduced regrowth Inman (1970)
Cutting R.o. Shoots, roots Field, greenhouse Cutting in 3-weekly intervals reduced carbohydrate content Hidaka (1973)
in shoots and roots
R.o. Shoots Field, orchard Cutting two times/month reduced R.o. cover by 50% Guncan (1979)
R.o. Shoots Field, grassland 5–7 cuts/year reduced R.o. abundance by 60%, after Courtney (1985)
3–4 cuts/year by 40%
R.c., R.o. Shoots Field, grassland The higher cutting frequency the lower biomass of R.c. Hongo (1987)
and R.o.; R.o. showed more regrowth after cutting than R.c.
R.c., R.o. Shoots, roots Field, grassland Lower cutting height led to less seeds in R.c; R.o. seed Hongo (1988a)
production unresponsive to cutting height
R.c., R.o. Shoots, roots Field, grassland Viable seed production after fourth cutting higher in R.o. Hongo (1988b)
than R.c.; root mass higher in R.o. than R.c.
R.o. Leaves, shoots Greenhouse Regrowth not affected by cutting Nilsson and Hallgren (1991)
R.o. Shoots, roots Field, grassland Frequent cutting stimulated seedling growth and regeneration Hughes et al. (1993)
of tap roots and prevented seed production in adult plants
Novak (1995a,b)
Courtney (1985)
Nashiki (1995)
mainly aim at reducing or eliminating existing Rumex
infestations, cultural methods generally focus on the
Reference
prevention of infestations. Cultural control comprises
grazing, undersowing, mulching, crop rotation and the
choice of cultivars (Table 3). There is a general
R.c. emergence
cattle or goats
on bare ground and that their growth is negatively
Main result
arable land
arable land
long term Niggli and colleagues (1993) doubt that
Field, grassland
Field, grassland
grassland
grassland
grassland
grassland
Field, grassland
Field, grassland
Experimental
Shoots
Shoots
Shoots
Shoots
Shoots
Shoots
Shoots
Shoots
Shoots
Shoots
Shoots
grazing regime (five to seven cuts per year) Rumex
suppressed grass yields by no more than 16% whereas
under simulated silage cutting (three to four cuts per
Rumex spp.
R.o.
R.o.
R.o.
R.o.
R.o.
R.o.
R.o.
R.o.
R.c.
R.c.
research.
Grazing (simulated by 5–7 cuts/year)
Tillage
Crop cultivar and seeding density
Seeding mixture
Crop rotation
Seeding rate
of carefully managed rotovations timed to exhaust the Alopecurus pratensis L., increasing nitrogen fertilization
root of Rumex spp. (Cavers & Harper, 1964; Hunt & stimulated R. obtusifolius more than the competing
Harkess, 1968). However, tillage has also been reported grasses (Niggli, 1985), and an additional reduction of
to stimulate Rumex distribution in arable land (Dierauer cutting frequency from six to four per season stimulated
& Stöppler-Zimmer, 1994). Manipulative experiments R. obtusifolius especially at high nitrogen levels (Niggli,
testing the effects of different tillage methods on Rumex 1985). Fertilizer applications of 200 kg ha)1 year)1
populations and distribution are seriously lacking. The nitrogen increased leaf area of Rumex species in autumn
effect of tillage in darkness on seed germination of by 70% and resulted in a 19% higher leaf mass and a
Rumex species in arable land (Hartmann & Nezadal, 43% higher root mass compared with unfertilized plants
1990) is also unclear. Because Rumex is a light-sensitive (Hatcher et al., 1997a). The finding that an increase in
weed species, ploughing and tillage on bright days might added nitrogen from 200 to 400 kg ha)1 year)1 did not
provoke a flush of emerging seedlings in the subsequent affect Rumex biomass production (Hatcher et al.,
weeks. Therefore, soil tillage in darkness may reduce 1997a) suggests that even nitrophilous Rumex may not
germination and emergence of Rumex species by necessarily profit from excessive fertilizer applications.
excluding light during soil disturbance. Long-term Increasing nitrogen fertilization did also not allow
cultivation and different tillage systems produce im- R. obtusifolius to escape the effects of the insect and
portant changes in the composition and density of the fungus and did not alter the additive nature of the
soil seedbanks (e.g. Cardina et al., 1991) and the effects insect–fungus interaction (Hatcher et al., 1997a,c).
of different tillage systems take a very long time to
become evident (Feldman et al., 1997). By comparing
Grazing management
the soil seedbank of different tillage systems it could be
shown that the tillage system which caused more Effects of different grazing systems (e.g. rotational
disturbance (e.g. mouldboard ploughing) was more grazing, set stocking, continuous grazing) or stocking
advantageous to germination of R. crispus than the rate (i.e. grazing animals per hectare) on Rumex
systems of lesser disturbances (e.g. disc, chisel and infestation have rarely been studied (Table 3). Grazing
no-tillage; Feldman et al., 1997). The situation becomes management can influence the ability of Rumex to
even more complex as soil nitrate concentrations has invade pastures, primarily by altering the competitive-
been shown to influence the germination of Rumex; an ness of other pasture species as Rumex is usually
early application of nitrogen fertilizer just before or after avoided by grazers. It is generally assumed that both
soil disturbance is suggested to result in higher Rumex species are mainly refused or eaten only in very young
seedling emergence than a later application (Milberg, stages by cattle, sheep and horses, but is a favourite
1997). This is a very interesting area which should be food plant of deer (Cavers & Harper, 1964). However,
explored further and could result in recommendations there are also observations that even mature, up to
for how to best combine fertilization and tillage in order 1.5 m tall plants of R. crispus and R. obtusifolius are
to control Rumex species in arable land. Additionally, completely removed by certain sheep breeds (J G Zaller,
time of soil disturbance can make an enormous differ- unpubl. obs.). More studies need to be conducted on
ence to the dynamics of weed populations. For instance, how mixed grazing can be used to control Rumex
soil disturbance in late summer has been suggested to distribution in pastures, for instance simultaneous
actually promote the growth of Rumex populations grazing by cows and goats improved R. obtusifolius
rather than prevent it (Weaver & Cavers, 1979a,b). control better than single grazing by either cows or
However, soil disturbance during winter prevents goats (Sakanoue et al., 1995).
Rumex germination especially when daytime soil surface
temperature do not reach 15C (Roberts & Totterdell,
Other indirect control measures
1981).
The most important indirect control measures against
Rumex infestations according to Dierauer and Stöppler-
Fertilization
Zimmer (1994) are: (i) a balanced fertilizer use on arable
Although a high fertilization rate is commonly stated as land and grassland, (ii) avoiding poaching damage by
a key factor supporting Rumex infestations, it is grazing animals or using heavy machinery in wet soil
surprising that there have been very few studies on the conditions, (iii) removing single Rumex plants before
effects of different application rates of fertilizers (e.g. they reach the seeding stage, (iv) careful composting of
slurry, manure, compost, synthetic fertilizer) on the farmyard manure and (v) use of certified dock-free
occurrence of Rumex in grassland and arable land. In seeding material. However, no information is available
mixtures with Lolium perenne L., Poa pratensis L. and as to what extent these indirect measures actually
contribute to the control or prevention of Rumex in Rumex infestation on arable land during the last
infestations. decades should also be investigated. Although Rumex
When mainly organic fertilizer is used, composting of root ecology has been studied by several authors, there
farmyard manure is recommended. Temperatures within are still some general problems which need to be
the heap of 55C or higher have been shown to destroy addressed, regarding root dynamics and root ecology,
the seeds of Rumex spp. (Dierauer & Stöppler-Zimmer, for instance: (i) How long do Rumex roots and
1994), and aeration of slurry for a few days can populations live? (ii) How important is root herbivory?
completely inhibit the germination ability of Rumex (iii) How important is clonal growth in dense grassland
seeds (Vogtmann et al., 1979). Additionally, there are swards as opposed to arable land? (iv) What is the
indications that not only higher temperatures during spatial extension of Rumex clones in grassland and
composting are responsible for decreased Rumex germi- arable land? (v) How important is clonal integration in
nation but also antagonistic non-thermophilic micro- Rumex populations? (vi) How important are nutrient-
organisms during composting (J G Zaller, unpubl. obs.). rich microsites for Rumex growth and development in
Another important vector for the spread of Rumex comparison with crop plants? Surprisingly, only a few
on agricultural land is through seeds in herbage seed studies focussed on the effects of soil properties on the
material. Rumex seeds have been found in 21% of development of Rumex establishment and distribution.
commercial crop seed material with wide variation Those data would be especially interesting to estimate
between different crops (Evans & Yates, 1981, 1985). the long-term success of single plant control strategies.
According to the regulations of the European Union, Moreover, more research directed into allelopathic
a certified grass–clover seed mixture can still contain interactions of Rumex species and crop species would
up to 200 Rumex seeds kg)1 – in Switzerland the help in improving the efficacy of new non-chemical
threshold being 10 seeds kg)1 (Dierauer & Stöppler- control strategies.
Zimmer, 1994), and in Austria 0 seeds kg)1 (Krautzer, Responses of Rumex species to global environmental
2001). changes such as elevated CO2, global warming, sum-
mertime ozone concentration, nitrogen deposition, ele-
vated UV-B radiation, decreasing species diversity in
Research needs: Rumex ecology and non-
agro-ecosystems, have rarely been investigated and
chemical control
deserve more research effort. Because the two Rumex
Although the reviewed Rumex species are among the species are undesirable on agricultural land, the design
most frequently studied plant species worldwide, it is of ecological experiments should always include a link to
surprising how many aspects regarding their ecology interactions of particular ecological properties with
and non-chemical control have still not been addressed certain management practices.
in past research. Some specific areas in which further
research is needed are listed below.
Non-chemical control
systems, investigations on the influence on Rumex of (Gastrophysa viridula) on seed production and quality in
crop rotation design and/or different methods of soil Rumex obtusifolius and Rumex crispus. Journal of Ecology
cultivation (e.g. ploughing, direct drilling, harrowing, 68, 671–674.
Beres I & Kazinczi G (2000) Allelopathic effects of shoot
tillage at darkness) are needed.
extracts and residues of weeds on field crops. Allelopathy
As the combination of different biocontrol agents (e.g. Journal 7, 93–98.
Coleoptera insects or rust fungi) has been demonstrated Bingaman BR & Christians NE (1995) Greenhouse screening of
to be effective to decrease Rumex biomass production corn gluten meal as a natural control product for broadleaf
under controlled conditions, more research on the and grass weeds. HortScience 30, 1256–1259.
combination of biological methods, also with mechanical Bohner A (2001) Physiologie und futterbaulicher Wert des
and cultural methods at varying environmental condi- Ampfers. In: 7. Alpenländisches Expertenforum zum Thema
ÔÔBestandesführung und Unkrautregulierung im Grünland –
tions, should be conducted (Hatcher & Melander, 2003).
Schwerpunkt Ampfer (ed. Bundesanstalt für alpenländische
Research on indirect control measures should also
Landwirtschaft, Gumpenstein), 39–44. Bundesanstalt für
include the influence of different composting methods alpenländische Landwirtschaft, Irdning, Austria.
on Rumex residue and seed viability and the effect of Bosworth SC, Hoveland CS & Buchanan GA (1986) Forage
certified Rumex-free seed material versus non-certified quality of selected cool-season weed species. Weed Science
seed material on the expansion of these Rumex species. 34, 150–154.
Boyd NS & Van Acker RC (2003) The effects of depth and
fluctuating soil moisture on the emergence of eight
Acknowledgements annual and six perennial plant species. Weed Science 51,
725–730.
I am very grateful to Peter Juroszek for critical Brooks GL & Whittaker JB (1998) Responses of multiple
comments on the manuscript. generations of Gastrophysa viridula feeding on Rumex
obtusifolius, to elevated CO2. Global Change Biology 4,
63–75.
References Brouwer W & Stählin A (1975) Handbuch der Samenkunde.
Adolf G & Linke G (1992) Zur Biologie des Stumpfblättrigen DLG-Verlag, Frankfurt/Main, Germany.
Ampfers (Rumex obtusifolius L.). In: Erfahrungen und Campiglia E, Temperini O, Mancinelli R, Marucci A & Saccardo
Ergebnisse bei der Bekämpfung von Ampfer (Rumex F (1998) Soil solarization in the Mediterranean environment:
obtusifolius L.) auf mehreren Grünlandstandorten unter effect on weed control and yield of cos lettuce (Lactuca sativa
Praxisbedingungen (ed. Martin-Luther-Universität L. var. longifolia Lam.). Italus Hortus 5, 36–42.
Halle-Wittenberg), 1–19. Martin-Luther-Universität, Cardina J, Regnier E & Harrison K (1991) Long-term tillage
Halle-Wittenberg, Germany. effects on seed banks in three Ohio soils. Weed Science 39,
Allard RW (1965). Genetic systems associated with colonizing 186–194.
ability in predominantly self-pollinating species. In: The Cavers PB (1967) Studies in the dynamics of plant populations.
Genetics of Colonizing Species (eds HG Baker & GL 1. The fate of seed and transplants introduced into various
Stebbins), 49–75.Academic Press, New York, USA. habitats. Journal of Ecology 55, 59–71.
Aquilina M & Clarke JH (1994) Effect of cutting date and Cavers PB & Harper JL (1964) Biological flora of the British
frequency on perennial broad-leaved weeds on set-aside. Isles. Rumex obtusifolius L. and R. crispus L. Journal of
Aspects of Applied Biology 40, Arable Farming Under CAP Ecology 52, 737–766.
Reform, Part 2, 541–546. Chapin FS, Schulze E-D & Mooney HA (1990) The ecology and
van Assche JA & Vanlerberghe KA (1989) The role of economics of storage in plants. Annual Review of Ecology
temperature on the dormancy cycle of seeds of Rumex and Systematics 21, 423–447.
obtusifolius L. Functional Ecology 3, 107–115. Cideciyan MA & Malloch AJC (1982) Effects of seed size on the
Bach A (1992) Ergebnisse der flächendeckenden Amp- germination growth and competitive ability of Rumex
ferbekämpfung in etablierten Beständen. In: Erfahrungen crispus and Rumex obtusifolius. Journal of Ecology 70,
und Ergebnisse bei der Bekämpfung von Ampfer (Rumex 227–232.
obtusifolius L.) auf mehreren Grünlandstandorten unter Cooper J, Maxwell T & Owens A (1960) A study of the passage
Praxisbedingungen (ed. Martin-Luther-Universität of weed seeds through the digestive tract of the chicken.
Halle-Wittenberg), 1–15. Martin-Luther-Universität, Poultry Science 39, 161–163.
Halle-Wittenberg, Germany. Cottam DA, Whittaker JB & Malloch AJC (1986) The
Baskin JM & Baskin CC (1985) Does seed dormancy play a role effects of chrysomelid beetle grazing and plant competition
in the germination ecology of Rumex crispus? Weed Science on the growth of Rumex obtusifolius. Oecologia 70,
33, 340–343. 452–456.
Bentley S & Whittaker JB (1979) Effects of grazing by a Courtney AD (1985). Impact and control of docks in grassland.
chrysomelid beetle, Gastrophysa viridula, on competition In: Weeds, Pests and Diseases of Grassland and Herbage
between Rumex obtusifolius and Rumex crispus. Journal of Legumes (ed. JS Brockman), 120–127. British Crop Protec-
Ecology 67, 79–90. tion Council, Croydon, UK.
Bentley S, Whittaker JB & Malloch AJC (1980) Field Courtney AD & Johnston R (1978) A consideration of the
experiments on the effects of grazing by a chrysomelid beetle contribution to production of Rumex obtusifolius in a
grazing regime. In: Proceedings 1978 of the British Differences in organs and diurnal changes. Oecologia 63,
Crop Protection Conference – Weeds, Brighton, UK, 136–142.
325–331. Gleason FK & Case DE (1986) Activity of the natural algicide,
Darlington H & Steinbauer GP (1961) The 80 year period of cyanobacterin, on angiosperms. Plant Physiology 80, 834–837.
Dr. Beal’s seed viability experiment. American Journal of Gonzalez L, Souto XC, Sanchez AM & Reigosa MJ (1995)
Botany 48, 321–325. Response of the weeds Rumex crispus, Plantago lanceolata,
DeGregorio RE, Ashley RA, Adams RG, Streams FA & Chenopodium album, Amaranthus retroflexus, Cirsium sp. and
Schaefer CW (1991) Biocontrol potential of Hypera rumicis Solanum nigrum to the addition of allelopathic compounds.
(L.) (Coleoptera: Curculionidae) on curly dock (Rumex In: Proceedings of the 1995 Congress of the Spanish Weed
crispus L.). Journal of Sustainable Agriculture 2, 7–24. Science Society (ed. Spanish Weed Science Society), 345–348.
Dierauer H-U & Stöppler-Zimmer H (1994) Unkrautregulierung Spanish Weed Science Society, Huesca, Spain.
ohne Chemie. Verlag Eugen Ulmer, Stuttgart, Germany. Gonzalez L, Souto XC & Reigosa MJ (1997) Weed control by
Dierauer HU & Thomas JM (1994) Efficiency of different Capsicum annuum. Allelopathy Journal 4, 101–110.
non-chemical methods of controlling broadleaf dock (Rumex Gough RE & Carlstrom R (1999) Wheat gluten meal inhibits
obtusifolius). In: Maitrise des adventices par voie non germination and growth of broadleaf and grassy weeds.
chimique. Communications de la Quatrieme Conference HortScience 34, 269–270.
Internationale I.F.O.A.M., (ed. IFOAM), 311–314. IFOAM, Greaves MP (1986) Developments in biologial control. British
Dijon, France. Crop Protection Society Monograph 29, 155–160.
Ellenberg H (1986) Vegetation Mitteleuropas mit den Alpen, 4th Grime JP (1996) The changing vegetation of Europe: What is
edn. Verlag Eugen Ulmer, Stuttgart, Germany. the role of elevated carbon dioxide? In: Carbon dioxide,
Ellenberg H, Weber HE, Düll R, Wirth V, Werner W & Populations, and Communities (eds C Körner & FA Bazzaz),
Paulissen D (1991) Zeigerwerte von Pflanzen in Mitteleuropa. 85–92. Academic Press, San Diego, CA, USA.
Verlag Erich Goltze, Göttingen, Germany. Guncan A (1979) The biology of Rumex obtusifolius L. and
Evans AW & Yates CW (1981) Survey of the distribution of its control in hazelnut plantations. Ataturk Universitesi
important weeds in herbage seed crops in England and Yayinlari 558, 28.
Wales in 1978 season. Journal of the National Institute of Gwynne DC & Murray CB (1985) Weed Biology and Control
Agricultural Botany 16, Cambridge, UK, 289–309. in Agriculture and Horticulture. Batsford Academic and
Evans AW & Yates CW (1985) Survey of the distribution and Educational, London, UK.
occurrence of weeds in herbage seed crops in England and Haggar RJ (1980) Survey on the incidence of docks (Rumex
Wales in 1983 and a comparison with 1973 and 1978 seasons spp.) in grassland in 10 districts in U.K. in 1972. A.D.A.S.
already reported. Journal of the National Institute of Quarterly Review 39, 256–270.
Agricultural Botany 17, Cambridge, UK, 75–93. Hand DJ, Craig G, Takaki M & Kendrick RE (1982) Interaction
Fairbairn CG & Thomas B (1959) The potential nutritive value of light and temperature of Rumex obtusifolius. Planta 156,
of some weeds common to northeastern England. Journal 457–460.
of the British Grassland Society 14, 36–46. Hann P & Kromp B (2003) Der Ampferblattkäfer (Gastrophysa
Feldman SR, Alzugaray C, Torres PS & Lewis P (1997) The viridula Deg.) – ein natürlicher Gegenspieler des Stumpf-
effect of different tillage systems on the composition of the blättrigen Wiesenampfers (Rumex obtusifolius). Landbau-
seedbank. Weed Research 37, 71–76. forschung Völkenrode 255, 73–78.
Fenner M (1985) Seed Ecology. Chapman and Hall, London, Harper JL (1965) Establishment, aggression, and cohabitation
UK. in weedy species. In: The Genetics of Colonizing Species (eds
Fisher NM, Davies DHK & Williams GH (1993) Vegetation HG Baker & GL Stebbins), 243–266. Academic Press, New
succession and the effect of management on weedy species in York, USA.
an old arable field under set-aside. In: Proceedings 1993 of a Harper JL (1977) Population Biology of Plants. Academic Press,
Conference on Crop Protection in Northern Britain, Dundee, London, UK.
UK, 229–234. Hartmann KM & Nezadal W (1990) Photocontrol of weeds
Foster L (1989) The biology and non-chemical control of dock without herbicides. Naturwissenschaften 77, 158–163.
species Rumex obtusifolius and R. crispus. Biological Hatcher PE (1996) The effect of insect–fungus interactions on
Agriculture and Horticulture 6, 11–25. the autumn growth and over-wintering of Rumex crispus and
French RC, Kujawski PT & Leather GR (1986) Effect of various R. obtusifolius seedlings. Journal of Ecology 84, 101–109.
flavor-related compounds on germination of curly dock seed Hatcher PE & Melander B (2003) Combining physical, cultural
(Rumex crispus) and curly dock rust (Uromyces rumicis). and biological methods: prospects for integrated non-
Weed Science 34, 398–402. chemical weed management strategies. Weed Research 43,
Froud-Williams RJ, Drennan DSH & Chancellor RJ (1984) The 303–322.
influence of burial and dry-storage upon cyclic changes in Hatcher PE & Paul ND (2000) Beetle grazing reduces natural
dormancy, germination and response to light in seeds of infection of Rumex obtusifolius by fungal pathogens. New
various arable weeds. New Phytologist 96, 473–481. Phytologist 146, 325–333.
Galler J (1989) Grünlandverunkrautung. Ursachen, Vorbeu- Hatcher PE, Paul ND, Ayres PG & Whittaker JB (1994a) The
gung, Bekämpfung. Leopold Stocker Verlag, Graz, Austria. effect of an insect herbivore and a rust fungus individually,
Gebauer G, Melzer A & Rehder H (1984) Nitrate content and and combined in sequence, on the growth of two Rumex
nitrate reductase activity in Rumex obtusifolius L. I. species. New Phytologist 128, 71–78.
Hatcher PE, Paul ND, Ayres PG & Whittaker JB (1994b) Hongo A (1988b) Effect of cutting on growth and seed
Interactions between Rumex spp., herbivores and a rust production of Rumex obtusifolius L. and Rumex crispus L. in
fungus: Gastrophysa viridula grazing reduces subsequent eastern Hokkaido. II. Cutting at different maturing stages
infection by Uromyces rumicis. Functional Ecology 8, with respect to plants established from seeds. Weed
265–272. Research, Japan 33, 8–13.
Hatcher PE, Paul ND, Ayres PG & Whittaker JB (1994c) The Hongo A (1989) Survival and growth of seedlings of Rumex
effect of foliar disease (rust) on the development of obtusifolius L. and Rumex crispus L. in newly sown grass-
Gastrophysa viridula (Coleoptera: Chrysomelidae). Ecologi- land. Weed Research, Japan 29, 7–12.
cal Entomology 19, 349–360. Hopkins A & Johnson RH (2002) Effect of different manuring
Hatcher PE, Paul ND, Ayres PG & Whittaker JB (1995a) and defoliation patterns on broad- leaved dock (Rumex
Interactions between Rumex spp, herbivores and a rust obtusifolius) in grassland. Annals of Applied Biology 140,
fungus – the effect of Uromyces rumicis infection on leaf 255–262.
nutritional quality. Functional Ecology 9, 97–105. Hopkins MJG & Whittaker JB (1980) Interactions between
Hatcher PE, Ayres PG & Paul ND (1995b) The effect of Apion species (Coleoptera: Curculionidae) and Polygona-
natural and simulated insect herbivory, and leaf age, on the ceae. II. Apion violaceum Kirby and Rumex obtusifolius
process of infection of Rumex crispus L. and R. obtusifolius L. Ecological Entomology 5, 241–247.
L. by Uromyces rumicis (Schum.) Wint. New Phytologist 130, Hopkins MJG, Williams P & Fisher JP (1984) The parasite
239–249. complex associated with stem-boring Apion (Col., Curcu-
Hatcher PE, Paul ND, Ayres PG & Whittaker JB (1997a) lionidae) feeding on Rumex species (Polygonaceae).
Added soil nitrogen does not allow Rumex obtusifolius to Entomologist’s Monthly Magazine 120, 187–192.
escape the effects of insect–fungus interactions. Journal of Hopkins A, Jones EL, Bowling PJ & Johnson RH (1997) Cultural
Applied Ecology 34, 88–100. methods of dock control in permanent pasture. In: British
Hatcher PE, Paul ND, Ayres PG & Whittaker JB (1997b) Grassland Society Fifth Research Conference (ed. British
Nitrogen fertilization affects interactions between the com- Grassland Society), 39–40. University of Plymouth, Newton
ponents of an insect–fungus–plant tripartite system. Func- Abbot, Devon, UK.
tional Ecology 11, 537–544. Huarte HR & Arnold RLB (2003) Understanding mechanisms
Hatcher PE, Paul ND, Ayres PG & Whittaker JB (1997c) of reduced annual weed emergence in alfalfa. Weed Science
The effect of nitrogen fertilization and rust fungus 51, 876–885.
infection, singly and combined, on the leaf chemical Huber-Meinicke G, Defago G & Sedlar L (1989) Ramularia
composition of Rumex obtusifolius. Functional Ecology 11, rubella (Bon.) Nannf. as a potential mycoherbicide against
545–553. Rumex weeds. Botanica Helvetica 99, 81–89.
Healy A (1953) Control of docks. New Zealand Journal of Hudson J (1955) Propagation of plants by root cuttings. 2.
Science and Technology, Section A 34, 473–475. Seasonal fluctuation of capacity to regenerate from roots.
Hidaka M (1973) Effect of cutting on the total nonstructural Journal of Horticultural Science 30, 242–251.
carbohydrates (TNC) contents in the roots and crowns of Hughes EW (1938) Studies into the biology of some Rumex
Rumex obtusifolius L. Journal of the Japanese Grassland species. MSc thesis, University of Wales, Cardiff, Wales,
Society 19, 313–317. UK.
Hofmann F (1992) Der Stumpfblättrige Ampfer und Hughes CNG, Froud Williams RJ & Fox RTV (1993) The effects
Möglichkeiten seiner Bekämpfung. In: Erfahrungen und of fragmentation and defoliation on Rumex obtusifolius and
Ergebnisse bei der Bekämpfung von Ampfer (Rumex obtusi- its implication for grassland management. In: Proceedings
folius L.) auf mehreren Grünlandstandorten unter Praxisbed- 1993 of the Brighton Crop Protection Conference – Weeds,
ingungen (ed. Martin-Luther-Universität Halle-Wittenberg), Brighton, UK, 767–772.
1–13. Martin-Luther-Universität, Halle-Wittenberg, Hughes CNG, West JS, Fox RTV, Moran VC & Hoffmann JH
Germany. (1996) Control of broad-leaved docks by Armillaria mellea.
Holm L, Plucknett DL, Pancho JV & Herberger JP (1977) The In: Proceedings 1996 of the 9th International Symposium on
World’s Worst Weeds: Distribution and Biology. University Biological Control of Weeds, Stellenbosch, South Africa,
Press of Hawaii, Honolulu, HI, USA. 531–534.
Hongo A (1986) Infestation of Rumex obtusifolius L. and Hultén E (1950) Atlas över växternas utbredning i norden.
distribution pattern of its individual plants in sown grass- Generalstabens Litografiska Anstalts Förlag, Stockholm,
lands in Eastern Hokkaido. Weed Research, Japan 31, Sweden.
300–305. Hunt I & Harkess R (1968) Docks in grassland. Scottish Journal
Hongo A (1987) Effects of treading and cutting on the growth of Agriculture 47, 160–162.
of broadleaved dock (Rumex obtusifolius L.) and curly dock Inman RE (1970) Observations on the biology of Rumex rust
(Rumex crispus L.). Research Bulletin of Obihiro University Uromyces rumicis (Schum.)Wint. Botanical Gazette 131,
15, 107–112. 234–241.
Hongo A (1988a) Effect of cutting on growth and seed Isikawa S & Fujii T (1961) Photocontrol and temperature
production of Rumex obtusifolius L. and Rumex crispus L. in dependence of germination of Rumex seeds. Plant and Cell
eastern Hokkaido. I. Cutting at different maturing stages Physiology 2, 51–62.
and different heights with respect to transplanted plants. Jeangros B & Nösberger J (1990) Effects of an established sward
Weed Research, Japan 33, 1–7. of Lolium perenne L. on the growth and development of
Rumex obtusifolius L. seedlings. Grass and Forage Science 45, Le Deunff Y (1971) Mise en évidence du phytochrome chez
1–7. les smences de Rumex crispus L. Annales de Physiologie
Jimenez JJ, Schultz K, Anaya AL, Hernandez J & Espejo O Ve´ge´tale 9, 201–208.
(1983) Allelopethic potential of corn pollen. Journal of Le Deunff Y & Chaussat R (1968) Etude de la dormance
Chemical Ecology 9, 1011–1025. secondaire des semences chez Rumex crispus L. Annales de
Julien MH, Kassulke RC & Harley KLS (1982) Lixus Physiologie Ve´ge´tale 10, 227–236.
cribricollis (Col.: Curculionidae) for biological control of the Lousley JE (1939) Notes on British Rumices. I. Botanical
weeds Emex spp. and Rumex crispus in Australia. Entomo- Society and Exchange Club of the British Isles Report 12,
phaga 27, 439–446. 118–157.
Kasai T, Okuda M, Sano H, Mochizuki H, Sato H & Sakamura S Machatschek M (1999) Nahrhafte Landschaft. Ampfer,
(1982) Biological activity of the constituents in roots of Kümmel, Wildspargel, Rapunzelgemüse, Speiselaub und
ezo-no-gishigishi (Rumex obtusifolius). Agricultural and andere wiederentdeckte Nutz- und Heilpflanzen. Böhlau
Biological Chemistry 46, 2809–2813. Verlag, Vienna, Austria.
Kingsbury, JM (1964) Poisonous Plants of the United States and Marten GC, Sheaffer CC & Wyse DL (1987) Forage nutritive
Canada. Prentice-Hall, New York, USA. value and palatability of perennial weeds. Agronomy Journal
Kivilaan A & Bandurski RS (1981) The one hundred year 79, 980–986.
period for Dr. W. J. Beal’s seed viability experiment. Martinkova Z & Honek A (2001) Regeneration of Rumex
American Journal of Botany 68, 1290–1292. obtusifolius L. after cutting. Rostlinna Vyroba 47, 228–232.
Klimes L (1993) Morphometry of leaves of Rumex crispus, R. Maun MA & Cavers PB (1971) Seed production and dormancy
obtusifolius and their hybrid (R. x pratensis). Preslia 65, of Rumex crispus. I. The effects of removal of cauline leaves
219–224. at anthesis. Canadian Journal of Botany 49, 1123–1130.
Kohout V (1994) The weevil (Apion miniatum Germar) – a Melzer A, Gebauer G & Rehder H (1984) Nitrate content and
biological regulator of distribution of broad-leaved docks. nitrate reductase activity in Rumex obtusifolius L. II.
Ochrana Rostlin 30, 79–81. Responses to nitrate starvation and nitrogen fertilization.
Kohout V & Kohoutova SN (1994) Possibilities of utilization of Oecologia 63, 380–385.
species Apion miniatum Germar in biological control of Meusel H, Jäger E & Weinert E (1965) Vergleichende
genus Rumex. Zeitschrift für Pflanzenkrankheiten und Chorologie der Zentraleuropäischen Flora. Karten Gustav
Pflanzenschutz 14, 217–220. Fischer Verlag, Jena, Germany.
Koukal J & Dagger W (1967) Anthocyanin formation as a Milberg P (1997) Weed seed germination after short-term light
response to ozone and smog treatment in Rumex crispus L. exposure: Germination rate, photon fluence response and
Plant Physiology 42, 1023–1024. interaction with nitrate. Weed Research 37, 157–164.
Krautzer B (2001) Saatgutqualität als Grundlage für ampfer- Mineta T, Hidaka K, Enomoto T & Oki Y (1997) Changes in
freie Nach- und Neuansaaten im Grünland. In: 7. Alpen- weed communities in direct-seeded paddy fields under
ländisches Expertenforum zum Thema ÔÔBestandesführung Astragalus sinicus L. living mulch and no-tillage cultivation
und Unkrautregulierung im Grünland – Schwerpunkt during three years. Journal of Weed Science and Technology
Ampfer (ed. Bundesanstalt für alpenländische Landwirt- 42, 88–96.
schaft Gumpenstein), 45–50. Bundesanstalt für alpenländi- Monaco T & Cumbo E (1972) Growth and development of curly
sche Landwirtschaft, Irdning, Austria. dock and broadleaf dock. Weed Science 20, 64–67.
Kühbauch W, Imhoff H & Voigtländer G (1980) Verteilung Moore JP, Taylor JE, Paul ND & Whittaker JB (2003) The use
der 14C-Aktivität nach 14CO2-Applikation auf die Nicht- of clip cages to restrain insects reduces leaf expansion
strukturkohlenhydrate im Stumpfblättrigen Ampfer, systemically in Rumex obtusifolius. Ecological Entomology
Wiesenkerbel und Bärenklau in verschiedenen Wachs- 28, 239–242.
tumsstadien. Zeitschrift für Acker- und Pflanzenbau 149, Naito A, Miyazaki M & Emura K (1979a) Studies on the
488–502. biological control of Rumex obtusifolius L. a grassland
Kurfess W & Kleisinger S (2000) Wirkung von Heißwasser auf weed by Gastrophysa atrocyanea Mots. (Chrysomelidae,
Unkrautpflanzen. Zeitschrift für Pflanzenkrankheiten und Coleoptera) 1. On the host specificity of the insect. Bulletin
Pflanzenschutz 17, 473–477. of the National Grassland Research Institute Japan 14,
Kurimoto S, Otake S, Takihiro T & Kimura H (1974) 117–123.
Ecological studies on the control of broad leaf dock, Naito A, Miyazaki M & Kanda K (1979b) Studies on the
Rumex obtusifolius L., in the grassland, 1: Some biological control of Rumex obtusifolius L., a grassland
properties of germination. Bulletin of the Hiroshima weed, by Gastrophysa atrocyanea Mots. (Coleoptera:
Prefectural Agricultural Experiment Station (Japan) 33, Chrysomelidae). II. Dispersal of overwintered adult insects.
57–61. Applied Entomology and Zoology 14, 51–55.
Kutschera L, Lichtenegger E & Sobotik M (1992) Wurzelatlas Nashiki M (1995) Competition of the weed, Rumex obtusifolius
mitteleuropäischer Grünlandpflanzen. Band 2. Pteridophyta L., with pasture plants for early control in new grassland.
und Dicotyledoneae (Magnoliopsida). Teil 1: Morphologie, Bulletin of the Tohoku National Agricultural Experiment
Anatomie, Ökologie, Verbreitung, Soziologie, Wirtschaft. Station 90, 93–153.
Gustav Fischer Verlag, Stuttgart, Germany. Nashiki M, Meguro R & Suyama T (1993) Control of growth of
Lang V, Voigtländer G & Kühbauch W (1974) Zum Reserve- Rumex obtusifolius L. during grass establishment by
stoffwechsel von Stumpfblättrigen Ampfer (Rumex obtusi- manipulation of seeding rate. Weed Research, Japan 38,
folius L.). Weed Research 15, 153–158. 205–213.
Nashiki M, Meguro R & Fukuda E (1998) Effect of cutting on regeneration of Rumex obtusifolius L. Weed Research 35,
carbohydrate content of Rumex obtusifolius L. Journal of 141–148.
Weed Science and Technology 43, 274–276. Pino J, Sans FX, Masalles RM et al. (1996) Population
Niggli U (1985) Bekämpfung von Rumex obtusifolius L. und dynamics of Rumex obtusifolius under contrasting alfalfa
Bestandeslenkung in intensiv bewirtschafteten Naturwiesen. cropping systems. In: Proceedings 1996 of the Second
PhD Thesis, Eidgenössische Technische Hochschule, International Weed Control Congress, Copenhagen,
Zürich, Switzerland. Denmark, 165–170.
Niggli U & Irla E (1984) Neuere Geräte zur Einzelstock- Pino J, Sans FX & Masalles RM (1999) Effect of crop length on
Bekämpfung von Blacken (Rumex obtusifolius). Mitteilungen the population dynamics of Rumex obtusifolius in an alfalfa-
für die Schweizerische Landwirtschaft 1/2, 25–27. winter cereal crop rotation. In: SEMh Congreso 1999:
Niggli U, Nösberger J & Lehmann J (1993) Effects of nitrogen Sociedad Española de Malherbologia, Actas (ed. Weed
fertilization and cutting frequency on the competitive Control Spanish Society), 127–131. Weed Control Spanish
ability and the regrowth capacity of Rumex obtusifolius L. Society, Logroño, Spain.
in several grass swards. Weed Research 33, 131–137. Pötsch EM (2001) Wissenswertes zur mechanischen und
Nilsson H & Hallgren E (1991) Chemical and mechanical chemischen Ampferbekämpfung. In: 7. Alpenländisches
control of dock (Rumex obtusifolius). A greenhouse experi- Expertenforum zum Thema ÔÔBestandesführung und
ment. Swedish Crop Protection Conference Weeds and Weed Unkrautregulierung im Grünland – Schwerpunkt Ampfer
Control 32, 267–276. (ed. Bundesanstalt für alpenländische Landwirtschaft
Novak J (1995a) Changes in ruderal grassland after no-tillage Gumpenstein), 75–81, Bundesanstalt für alpenländische
resowing with cocksfoot. Acta Fytotechnica 51, 95–104. Landwirtschaft, Irdning, Austria.
Novak J (1995b) The influence of utilization and the changes of Pötsch EM (2003) Möglichkeiten der mechanisch/biologischen
nitrogen content in eutrophicated soil on the retreat of Ampferbekämpfung. Landbauforschung Völkenrode 255,
Rumex obtusifolius. Acta Fytotechnica 51, 73–80. 63–68.
Oberdorfer E (1990) Pflanzensoziologische Exkursionsflora, 6th Rehder H (1982) Nitrogen relations of ruderal communities
edn. Verlag Eugen Ulmer, Stuttgart, Germany. (Rumicion alpini) in the Northern Calcareous Alps.
Oswald AK & Haggar RJ (1976) The effect of asulam on two Oecologia 55, 120–129.
Lolium perenne swards containing Rumex obtusifolius. Weed Reiling K & Davison AW (1992) The response of native,
Research 16, 7–10. herbaceous species to ozone: growth and fluorescence
Oswald AK & Haggar RJ (1983) The effects of Rumex screening. New Phytologist 120, 29–37.
obtusifolius on the seasonal yield of two mainly Roberts RA & Hughes EW (1939) Biological studies in the
perennial ryegrass swards. Grass and Forage Science 38, control of docks (Rumex spp.). Welsh Journal of Agriculture
187–191. 15, 218–237.
Parr TW & Brockman JS (1985) The control of weed Roberts EH & Totterdell S (1981) Seed dormancy in Rumex
populations during grass establishment by the manipulation species in response to environmental factors: review article.
of seed rates. In: Weeds, Pests and Diseases of Grasslands and Plant, Cell and Environment 4, 97–106.
Herbage Legumes (ed. JS Brockman), 20–28. British Grass- Sakanoue S, Kitahara N & Hayashi H (1995) Biological control
land Society Occasional Symposium No. 18, British Crop of Rumex obtusifolius L. by goat grazing. Japan Agricultural
Protection Council, Croydon, UK. Research Quarterly 29, 39–42.
Pearson M & Brooks GL (1995) The influence of elevated CO2 Salt DT, Brooks GL & Whittaker JB (1995) Elevated
on growth and age-related-changes in leaf gas-exchange. carbon-dioxide affects leaf-miner performance and plant-
Journal of Experimental Botany 46, 1651–1659. growth in docks (Rumex spp). Global Change Biology 1,
Peel S & Hopkins A (1980) The incidence of weeds in grassland. 153–156.
In: Proceedings 1980 of the British Crop Protection Con- Savory BM & Soper D (1970) Factors affecting the control of
ference – Weeds, Brighton, UK, 877–890. docks (Rumex spp.) with Asulam. In: Proceedings 1970 of the
Pekrun C, Jund D, Hofrichter V, Wagner S, Thumm U & 10th British Weed Control Conference, Brighton, UK, 358–
Claupein W (2002) Pflanzen- und ackerbauliche Maßnahmen 365.
zur Ampferbekämpfung auf Acker- und Grünlandflächen Schubiger FX, Défago G, Kern H & Sedlar L (1986) Damage to
unter den Produktionsbedingungen des Ökologischen Rumex crispus L. and Rumex obtusifolius L. caused by the
Landbaus. Zeitschrift für Pflanzenkrankheiten und rust fungus Uromyces rumicis (Schum.) Wint. Weed Research
Pflanzenschutz 18, 533–540. 26, 347–350.
Piearce TG, Roggero N & Tipping N (1994) Earthworms and Smith RW & Whittaker JB (1980a) Factors affecting
seeds. Journal of Biological Education 28, 195–202. Gastrophysa viridula populations (Coleoptera: Chrysome-
Pino J, Sans FX, Masalles RM & Thomas JM (1994) The lidae) in different habitats. Journal of Animal Ecology 49,
effect of periodical shoot removal on Rumex obtusifolius L. 537–548.
establishment in irrigated alfalfa (Medicago sativa L.) Smith RW & Whittaker JB (1980b) The influence of habitat
crops. In: Maitrise des adventices par voie non chimique. type on the population dynamics of Gastrophysa viridula
Communications de la quatrieme conference internationale Degeer (Coleoptera: Chrysomelidae). Journal of Animal
I.F.O.A.M (ed. IFOAM), IFOAM, Dijon, France, 2, Ecology 49, 225–236.
319–324. Spencer NR, Rosenthal SG & Hostetter N (1981) Exploration
Pino J, Haggar RJ, Sans FX, Masalles RM, Hamilton RNS & for biotic agents for the control of Rumex crispus. In:
Sackville Hamilton RN (1995) Clonal growth and fragment Proceedings 1981 of the 5th International Symposium on
Biological Control of Weeds, Brisbane, Queensland, Hygienisierungseffekt durch die Belüftung von Gülle.
Australia, 125–151. Schweizerische landwirtschaftliche Monatshefte 57, 225–234.
Stebler FG & Schröter C (1892) Beiträge zur Kenntnis der Voigtländer G, Lang V and Kühbauch W (1976) Zum
Matten und Weiden der Schweiz. X. Versuch einer Übersicht Reservestoffwechsel des Stumpfblättrigen Ampfers (Rumex
über die Wiesentypen der Schweiz. Landwirtschaftliches obtusifolius L.) und des Wiesenknöterich (Polygonum
Jahrbuch der Schweiz 6, 95–212. bistorta L.). Landwirtschaftliche Forschung 29, 109–117.
Takaki M, Kendrick RE & Dietrich SMC (1981) Interactions of Waghorn GC & Jones WT (1989) Bloat in cattle. Potential of
light and temperature on the germination of Rumex dock (Rumex obtusifolius) as an antibloat agent for cattle.
obtusifolius L. Planta 152, 209–214. New Zealand Journal of Agricultural Research 32, 227–235.
Taylorson RB & Hendricks SB (1972) Phytochrome control of Weaver SE & Cavers PB (1979a) Dynamics of seed populations
germination of Rumex crispus L. seeds induced by of Rumex crispus and R. obtusifolius (Polygonaceae) in
temperature shifts. Plant Physiology 50, 645–648. disturbed and undisturbed soil. Journal of Applied Ecology
Toole E & Brown E (1946) Final results of the Duvel buried 16, 909–917.
seed experiment. Journal of Agricultural Research 72, Weaver SE & Cavers PB (1979b) The effects of date of
201–210. emergence and emergence order on seedling survival rates in
Vengris J, Drake M, Colby WG & Bart J (1953) Chemical Rumex crispus and R. obtusifolius. Canadian Journal of
composition of weeds and accompanying crop plants. Botany 57, 730–738.
Agronomy Journal 45, 213–218. Whittaker JB (1982) The effect of grazing by a chrysomelid
Vergnano Gambi, O (1966) Influence of light and temperature beetle, Gastrophysa viridula, on growth and survival of
on the germination of some Rumex seeds. Webbia 21, Rumex crispus on a shingle bank. Journal of Ecology 70, 291–
461–474. 296.
Vicente M, Engelhardt M & Silberschmidt K (1962) The Whittaker JB, Ellistone J & Patrick CK (1979) The dynamics
influence of temperature on the germination response to of a chrysomelid beetle, Gastrophysa viridula, in a hazardous
light of seeds of Rumex obtusifolius. Phyton, Buenos Aires 19, natural habitat. Journal of Animal Ecology 48, 973–986.
163–167. Williams J (1971) Seed polymorphism and germination. 2. The
Vicente M, Noronha AB, Silberschmidt K & Meneghini M (1968) role of hybridization in germination polymorphism of
Successive reversion of the effect of temperature on Rumex crispus and R. obtusifolius. Weed Research 11, 12–21.
germination of Rumex obtusifolius L. by far-red light. Zaller JG (2004) Competitive ability of Rumex obtusifolius
Phyton, Buenos Aires 25, 11–16. against native grassland species: above- and belowground
Vogtmann H, Quante G, Plakolm G & Hurni B (1979) Die allocation of biomass and nutrients. Journal of Plant
Reduktion der Keimfähigkeit von Unkrautsamen und Diseases and Protection 19, 345–351.