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Ecology and non-chemical control of Rumex crispus

and R. obtusifolius (Polygonaceae): a review


J G ZALLER
Institute of Organic Agriculture, University of Bonn, Bonn, Germany

Received 26 February 2004


Revised version accepted 3 August 2004

are considered to be indicators of agricultural misman-


Summary
agement, there is a definite need for well-replicated, full-
Rumex crispus and R. obtusifolius are among the most factorial, long-term field experiments to assess the role
important perennial weeds in agriculture throughout the of management factors most often stated to be respon-
world. This literature survey revealed that the majority sible for Rumex infestation and distribution (e.g. high
of studies on Rumex ecology, which were conducted soil N and K levels, slurry and farmyard manure
under field conditions are of a descriptive nature, while application, sward disturbance, cutting frequency, gra-
experimental studies mainly report data derived under zing management, ploughing, soil compaction). It is
laboratory or greenhouse conditions. Areas on Rumex recommended that future research on non-chemical
ecology which deserve more research include: historical Rumex control should focus more on the efficacy of
development of Rumex infestations, field studies on seed combined applications of biological, mechanical and
and root ecology, population ecology, Rumex longevity, cultural control methods.
and impacts of global climate change on the perform-
Keywords: agricultural management strategies, arable
ance of these species. Although biological, mechanical
grassland, biological, cultural, mechanical, weed
and cultural control methods were often effective in
control, Rumex spp., research needs.
controlling Rumex populations, they were rarely suc-
cessful enough to eradicate the weeds. As both species

R. obtusifolius has a more distinct distribution and


Introduction
ranges from the sea level up to c. 1500 m in altitude.
Rumex crispus L. (curled dock) and Rumex obtusifolius These Rumex species are of agricultural significance
L. (broad-leaved dock) are among the most often because they compete with sown or native pasture
studied weed species worldwide, the latter is also species or arable crops and occupy area which could be
considered as one of the five most widely distributed utilized by more palatable crop species. Estimates of the
non-cultivated plant species in the world (Allard, 1965; area infested by these Rumex species are scarce: for the
see Hultén, 1950 and Meusel et al., 1965 for maps of UK it is estimated that c. 0.3–0.7 · 106 ha of grassland
their distribution). Both species are troublesome weeds are seriously infested with R. obtusifolius (Haggar, 1980;
in both grasslands (mainly pastures) and arable lands, Peel & Hopkins, 1980); for Germany, about seven of 10
but are also early colonizers of many disturbed areas in grassland farms are considered to have serious problems
lowland and upland. The species are present on almost with these species (Bach, 1992; Hofmann, 1992); in
all soil types but less often on peat and rarely on acid Japan (East-Hokkaido), 60% of all permanent grass-
soils. The range of altitude to which these species have lands are estimated to be infested with these species
become adapted is very great; a maritime ecotype of (Hongo, 1986). All those sources predict that Rumex
R. crispus grows on beaches another ecotype can also infestation will most likely increase in the future. For
be found at 2500 m above sea level in the Middle East Central Europe, it is estimated that more than 80% of
and south-western USA or at 3000 m in Iran all herbicides used in conventional grassland farming are
and 3500 m in Argentina (Hultén, 1950). Generally, used to control these species (Galler, 1989). The fear of

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

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Rumex ecology and non-chemical control 415

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).

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416 J G Zaller

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;

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Rumex ecology and non-chemical control 417

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

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418 J G Zaller

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

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Rumex ecology and non-chemical control 419

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

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420 J G Zaller

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.

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Table 1 Selected studies investigating biological control measures against Rumex crispus (R.c.) and R. obtusifolius (R.o.)

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)

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U. rumicis R.c., R.o. Greenhouse, Rust infection reduced leaf number, root Schubiger et al. (1986)
laboratory and leaf biomass
Fungus Armellaria mellea Sum. R.o. Laboratory Fungus increased root roening Hughes et al. (1996)
Coleoptera and G. viridula and U. rumicis R.c., R.o. Field Reduced leaf area and biomass caused by Hatcher et al. (1994a,b, 1994c)
rust fungus rust-herbivore interactions
G. viridula and U. rumicis R.c., R.o. Field Reduced leaf area and regrowth additive Hatcher (1996)
effects of fungi and beetles
G. viridula and U. rumicis R.o. Greenhouse Herbivory alone caused greater growth Hatcher et al. (1997b)
reduction than rust alone; additive effects
of fertilization, herbivory and rust
G. viridula and U. rumicis R.o. Field Beetle herbivory reduced fungal infection; Hatcher and Paul (2000)
and R. rubella and systematic induced resistance by R.o.
Venturia rumicis (Desm.) G. Winter independent of nutrient fertilization
Plant extracts Capsicum annuum L. R.c. Laboratory Inhibition of germination and growth Gonzalez et al. (1997)
Maize gluten meal R.c. Laboratory Reduced survival, shoot length and Bingaman and Christians (1995)
root development
Maize pollen R.c. Laboratory Reduction of radicle growth Jimenez et al. (1983)
Cyanobacterin R.c. Laboratory Inhibition of germination Gleason and Case (1986)
Wheat gluten R.c. Laboratory Reduced germination and growth Gough and Carlstrom (1999)
Natural Benzonitrile, benzyl cyanide, R.c. Laboratory Stimulation of germination and French et al. (1986)
chemicals methyl salicylate growth of Rumex and rust
Phenolics R.c. Laboratory Inhibition of germination Gonzalez et al. (1995)
Rumex ecology and non-chemical control 421
422 J G Zaller

best of my knowledge, no study investigated the


Mechanical and other direct control measures
possibilities of using cages containing G. viridula to
control Rumex species in the field. Moreover, no Mechanical methods for Rumex control have changed
information is available on naturally occurring biocon- little over the years and are dominated by field research
trol agents like seed-feeding insects. on effects of cutting and pulling by hand (Table 2). A
Fungi (especially the rust fungus U. rumicis) were threshold, when single plant control in grassland
tested very early for efficacy to control Rumex species in becomes uneconomical, is considered to be around
field experiments and were shown to reduce regrowth 2000 Rumex plants per hectare (Pötsch, 2003).
(Inman, 1970), leaf and root growth (Schubiger et al., Cutting has been shown to deplete carbohydrates
1986; Huber-Meinicke et al., 1989) and to increase root stored in Rumex shoots and roots (Hidaka, 1973; Lang
rotting (Hughes et al., 1996), however, again never et al., 1974; Voigtländer et al., 1976; Nashiki et al.,
sufficiently effective to eradicate Rumex. 1998) and to decrease seed input into the soil seedbank
Several studies have examined the interactive effects (Pino et al., 1994). Not very surprisingly, the more
between foliar fungal pathogens and herbivorous frequently the cutting occurs, the more the Rumex
insects (Hatcher et al., 1994a,b, 1997a; Hatcher, 1996; species are affected (Table 2). Cutting frequency has
Hatcher & Paul, 2000). Results from those experiments also been shown to affect the two species differently.
show that U. rumicis is poor at infecting young Rumex R. obtusifolius showed more regrowth capacity (Hongo,
leaves (Hatcher et al., 1994b, 1995a,b) while these 1987) and seed production (Hongo, 1988a,b) after
leaves are preferred by G. viridula for oviposition cutting than R. crispus. Cutting height also influenced
(Hatcher et al., 1994c). Beetles alone removed up to morphology (Hopkins & Johnson, 2002) and seed
79% of leaf area of both Rumex spp. and rust infection production (Hongo, 1987) of regrowing Rumex but
caused a decrease of about 50% in leaf area (Hatcher, not necessarily its biomass production (Hopkins &
1996). The combination of herbivory and infection had Johnson, 2002). As a result of the very rapid replenish-
an additive effect, causing a reduction in leaf area of ment of root carbohydrates by R. obtusifolius within 2
92% of R. crispus and 88% of R. obtusifolius (Hatcher, (Hidaka, 1973) to 3 weeks (Roberts & Hughes, 1939)
1996). Previous herbivory reduced the area and mass of even five to seven cuts per season after 6 years was able
regrowth in the following spring by 78% in both to reduce Rumex abundance only by 60% (Courtney,
species, and here too the combined effects of herbivory 1985). Cutting every 2 weeks will therefore be needed to
and infection were additive, leading to a 80% reduction control this species.
in shoot dry mass and an average 77% reduction in Effects of hand and/or machine pulling has rarely
root dry mass and a reduced amount of total non- been the subject of more recent studies (Niggli & Irla,
structural carbohydrates in the roots (Hatcher, 1996). 1984; Pekrun et al., 2002). New developments include a
These studies clearly demonstrate the possibility to motor-driven dock pulling machine which can pull up
control Rumex spp. with a combined application of about 600 Rumex plants per hour (Pötsch, 2003).
herbivores and fungi. Further research on this issue Comparisons of the efficiency of Rumex removal showed
should explore how the two agents can be applied that machine pulling is fastest (6 s/plant), followed by
under field conditions by farmers and should also hand-pulling (23 s/plant) and hand infrared flaming
monitor the long-term effects on Rumex populations of (50 s/plant) (Pötsch, 2003). No data are available on the
this control method. long-term effects of pulling or flaming on Rumex
Extracts or natural chemicals derived from various populations. Methods to regulate heavier Rumex infes-
plant species have been shown to reduce germination tations in grassland also include ploughing and roto-
and shoot and root development of Rumex seedlings vating. Shallow ploughing or rotovating is thought to
(Table 1). However, as all experiments were conducted sever the plants below the shoot buds and thus prevent
under laboratory conditions, it remains to be investi- the taproots from regenerating. When grassland is
gated whether those effects also occur under field ploughed, recruitment from the seedbank and re-estab-
conditions (Table 1). Several other agents (e.g. vinegar, lishment of fragments produced during agronomic
K–Mg salt, NaCl) have been suggested as potential practices would be the most important way for regen-
control methods against Rumex spp.; however, results eration of Rumex populations (Pino et al., 1995).
have not been published. Regeneration of Rumex species has been observed from
Previous research also suggests that efficacy of root fragments from different soil depths and root parts
biological control might be greater when Rumex plants as small as 0.5 cm (Dierauer & Stöppler-Zimmer, 1994);
are already stressed by ecological or environmental therefore, long-term effects of soil cultivation in grass-
conditions (e.g. competition, flooding) (Whittaker, land seem only to be effective if root fragments are
1982). collected after cultivation.

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Table 2 Main results of selected studies investigating mechanical and other direct non-chemical control measures against Rumex crispus (R.c.) and R. obtusifolius (R.o.)

Rumex spp. Plant part Experimental


Control measure tested studied conditions Main result Reference

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

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R.o. Leaves, shoots Field, grassland 4-weekly cutting interval more effective than 6-weekly; Niggli et al. (1993)
interaction with fertilization
R.o. Shoots, roots Field, grassland Cutting only effective when 10 cm below ground Dierauer and Thomas (1994)
R.o. Shoots Field, grassland Six cuts per year led up to 300% increase R.o. cover Aquilina and Clarke (1994)
R.o. Shoots Field, arable land Cutting reduced seed input into soil seedbank Pino et al. (1994)
R.o. Shoots, roots Field, grassland Four cuts per year decreased R.o. cover and Novak (1995a,b)
increased feeding value of herbage
R.o. Roots Field, grassland Cutting reduced carbohydrate content Nashiki et al. (1998)
R.o. Shoots Field, grassland Cutting advanced sprouting in following year Martinkova and Honek (2001)
Cutting and soil aeration R.c., R.o. Shoots Field, grassland Soil aeration and frequent cutting reduced Hopkins et al. (1997)
Cutting and manuring R.o. Leaves, shoots Field, grassland Height of cutting influenced ramet and leaf density but not Hopkins and Johnson (2002)
biomass production; 4-weekly cutting reduced R.o. in herbage
Cutting and drilling R.o. Shoots, roots Field, grassland The more root and shoot mass removed Pekrun et al. (2002)
the higher success; hand driller successful
Pulling R.o. Shoots, roots Field, grassland Drilling reduced R.o. cover Niggli and Irla (1984)
R.o. Shoots Field, grassland Automatic pulling reduced R.o. cover Pötsch (2001, 2003)
Flaming R.o. Shoots Field, arable land Flaming had little effect on Rumex cover Dierauer and Thomas (1994)
Solarization R.c. Shoots Field, arable land Suppression of R.c. cover by solarization Campiglia et al. (1998)
Hot water application R.o. Shoots Field, orchard Frequent application reduced R.o. cover Kurfess and Kleisinger (2000)
Rumex ecology and non-chemical control 423
424 J G Zaller

Parr and Brockman (1985)


Control through management practices (cultural

Huarte and Arnold (2003)

Pino et al. (1996, 1999);


Hann and Kromp (2003)
control)

Sakanoue et al. (1995)


Hopkins et al. (1997)
Nashiki et al. (1993);

Mineta et al. (1997)


Fisher et al. (1993)
While both biological and mechanical control methods

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

High seeding rate of L. perenne suppressed number of R.o. seedlings


consensus that the essential principle of any cultural

Lucerne increased R.o. cover, cereal cropping decreased R.o. cover


Nondormant cultivar of lucerne and higher seeding density reduced
Rumex control programme for grasslands must be the
provision of a dense, vigorous and competitive sward,

Mixed grazing reduced R.o. cover more than grazing by either


R.o. mass in grassland herbage lower after simulated grazing
High seeding rate with Lolium perenne reduced R.o. cover,
achieved by continuous undersowing and/or high seed-

Cutting time in consideration of dock beetle life cycle can


ing rates, moderate fertilization and a site-specific
stocking system (Parr & Brockman, 1985; Fisher et al.,

Mulching and no-tillage increased R.c. populations


1993; Nashiki, 1995; Novak, 1995b; Hopkins et al.,
1997). For arable land, a diverse crop rotation including

Seeding mixture prevented R.o. germination


regular weed control measures like harrowing, a high

fertilization had no effect on R.o. yield


seeding/planting density and a continuous crop cover

High sward density suppressed R.o.


seem to be important to prevent Rumex infestations.

L. perenne suppressed R.o. cover

reduce R.o. cover and regrowth


Recently, it was also demonstrated that lucerne culti-
vars with low levels of winter dormancy and an
Table 3 Selected studies investigating cultural control measures against Rumex crispus (R.c.) and R. obtusifolius (R.o.)

than after silage cutting


increased planting density could significantly reduce
the emergence of R. crispus (Huarte & Arnold, 2003).
The findings that Rumex seedlings are often found only

R.c. emergence

cattle or goats
on bare ground and that their growth is negatively
Main result

affected by shading indicate that control through in autumn


competition for light should be successful. The competi-
tion of grasses and herbs may also restrict Rumex
regrowth in the short term (Zaller, 2004), but for the
Field, arable land

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

effective control by competition of grasses can be


conditions

achieved, because of the tremendous regrowth potential


of Rumex species.
Field,
Field,
Field,
Field,
Field,
Field,
Sward management has been shown to be able to
significantly affect suppression of grass production by
Plant part

Rumex species. In a 6-year trial, with a simulated


studied

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.

year) grass yield was reduced by up to 70% (Courtney,


tested

1985). This is an area which definitely requires more


R.o.

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)

Mixed grazing by cattle and goats

Tillage
Crop cultivar and seeding density

A tillage operation is often recommended as the ultimate


non-chemical control measure to respond to heavy
Rumex infestations on arable land and even on grass-
Cutting management
Mulching, no-tillage

land. If a tillage operation removes the upper 7 cm of


Control measure

Seeding mixture

the root and then destroys it by desiccation or freezing,


Sward density
Undersowing

Crop rotation
Seeding rate

it is suggested that it is effective because the remaining


root cannot regenerate (Holm et al., 1977). Another
suggestion to eliminate Rumex spp. is to use a succession

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Rumex ecology and non-chemical control 425

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

 European Weed Research Society Weed Research 2004 44, 414–432


426 J G Zaller

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

An impressive body of literature is already available


Rumex ecology
on non-chemical control methods. However, the
Research on Rumex ecology was mainly focused on majority of research focused mainly on short-term
seeds and vegetation ecology derived from laboratory effects of control measures against Rumex species.
and greenhouse studies. However, there is still a lack of Thus, there is a definite need for long-term manipu-
data on germination and dormancy of seeds under field lative field experiments to assess the combined effects
conditions. Once the seeds germinated, survival rates of of various non-chemical control measures on Rumex
Rumex seedlings on open ground as opposed to shaded in a variety of grassland and arable agro-ecosystems.
sward gaps or dense sward communities are unclear. Moreover, more long-term research should focus on
Interesting for the development of control strategies the factors most often stated to be responsible for the
would also be to know whether more diverse grassland development of Rumex infestations (e.g. high soil
communities are better able to prevent the development nitrogen and potassium levels, fertilization with slurry
of Rumex infestations than less diverse grasslands. and farmyard manure, sward disturbance, cutting
There is still a fundamental lack of knowledge on the frequency).
variation of Rumex populations across different geo- Surprisingly little is known about the effects of
graphical locations and their adaptations to climatic different grazing systems (e.g. rotational grazing, set
conditions (e.g. annual precipitation, mean air tempera- stocking, continuous grazing), and/or stocking rate, the
ture) and spatial and temporal patterns of nutrient impact of different species or varieties of grazers on
availability on agricultural land. Moreover, the increase Rumex infestations in pasture ecosystems. For arable

 European Weed Research Society Weed Research 2004 44, 414–432


Rumex ecology and non-chemical control 427

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.
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