Review Consistent in Shape (Everist 1974) (Figure 2)

2 Plant Protection Quarterly Vol.24(1) 2009 2.1 m wide (Michael et al. 2006b). Leaves are alternate, dull dark green and generally variable in size (2–12 cm wide, 1–7 cm long) (Michael et al. 2006b), but fairly Review consistent in shape (Everist 1974) (Figure 2). Leaves have 5–7 deltate, crenate lobes (Dalby 1968) with leaf petioles (1.5–22 cm) longer than the blade (Michael 2006). Stip- ules are lanceolate to ovate, 2–5 mm long (Dalby 1968). The biology of Australian weeds Flowers emerge in axillary clusters (2–4 ﬂ owers per axil) on distinct peduncles 3–5 52. Malva parviﬂ ora L. mm long. Epicalyx segments are linear and 5–15 times as long as broad. The outer Pippa J. MichaelA, Kathryn J. SteadmanB and Julie A. PlummerC surface of the calyx has spreading stellate A School of Agriculture and Environment, Curtin University of Technology, hairs with 5–6 arms, each arm 0.05–0.3 mm Northam, Western Australia 6401, Australia. long (Barker 1977). Petals are white with B School of Pharmacy, University of Queensland, St Lucia, Queensland 4072, pinkish tips, oblong and slightly narrowed at the base, and have glabrous claws. They Australia. are 3–8 mm long, scarcely longer than the C School of Plant Biology, University of Western Australia, Crawley, Western Australia 6009, Australia.

(a) Name Flora, goddess of fl owers (Mitich, 1990). Malva (mallow) is derived from the Greek Common names in Australia for M. parvi- malache or malakos (soft), possibly referring fl ora are small-fl owered mallow, marsh- to either the downy leaves or its medicinal mallow, whorled mallow, whorlflower properties (Mitich 1990). The genus Malva mallow and ringleaf marshmallow, whilst is a member of the tribe Malveae of the overseas names include cheeseweed, little family Malvaceae (Corner 1976). In 1753, mallow and least mallow. Carolus Linnaeus was the fi rst to distinguish Malva species within the Malvaceae Description family based on their characteristic epica- Malva parvifl ora is a glabrous or pubescent lyx (Ray 1998). He originally identifi ed 15 annual (Dalby 1968) with a single long different Malva species. Currently, there taproot. Stems are either prostrate-ascend- are thought to be 25–40 species of Malva ing or erect (Figure 1) and can grow up to throughout the world. However, numer- 1.2 m high (Lamp and Collet 1984) and ous species initially placed in Malva have been transferred to other genera and there is much doubt about the correct nomen- clature of many Malva and other Malvace- ae species (Ray 1995). It was previously thought to be extremely unlikely that any species in the genus Malva naturally oc- (b) curred in Australia (Ray 1995); however three native Lavatera L. species have recently been changed to Malva (Ray 1998). Malva parvifl ora Linnaeus is one of several introduced Malva species that have naturalized in Australia. Linnaeus named parvifl ora from the Latin parvus (little), and

Figure 1. Malva parvifl ora showing Figure 2. Anatomy of Malva parvifl ora plant showing seedling (a) and its erect growth habit. fl owering stem (b) (Moerkerk and Barnett 1998). Plant Protection Quarterly Vol.24(1) 2009 3 epicalyx (Michael 2006). Flowers are bisex- Table 1. Distinguishing characteristics of the small-fl owered mallow group ual and unperfumed (Michael 2006). Pol- (information sourced from Hanf 1983). len sacs contain approximately 31 pollen grains, which are large (85 µm diameter) Character M. parvifl ora M. neglecta M. pusilla M. nicaeensis and sticky (Michael et al. 2006b). Corolla to calyx 1:1 2:1 1:1 2.5:1 The fruit (schizocarp) is a round cap- ratio sule approximately 1 cm in diameter containing between 8–12 mericarps. When Corolla colour pale pink pale pink/ pale pink/ pale violet ripe, mericarps change colour from green white white to dark brown (Michael et al. 2007). They Mericarp on dorsal distinctly faintly distinctly distinctly are narrowly and partially separated from face reticulate ribbed reticulate reticulate each other in the mature fruit, with angles ribbed ribbed ribbed on dorsal surface and toothed wing-like margins. Seeds are dark-brown when ripe Adjacent margins toothed smooth smooth smooth and variable in weight (0.008–0.17 g per 50 of the mericarp seeds). Seeds are reniform and non-hairy with an exotegmic seed coat comprised of six zones (Kumar and Singh 1991). Malva parviflora has a campylotropous ovule with integuments covering the embryo (a) (b) except for a small part of the chalazal end (c) (Corner 1976). The seed oil of M. parvifl ora is known to contain glycerides of cyclo- propene fatty acids (malvalic and sterculic), cyclopropane (dihydromalvalic and dihydrosterculic), epoxy and conjugated dienol acids (Ahmad et al. 1984). The dip- loid chromosome number for M. parvifl ora is 2n = 42 (Bidack and Brandham 1995).

Taxonomy Figure 3. Distinguishing characteristics of Malva parvifl ora (a) toothed Malva is traditionally distinguished from margins of the mericarp, (b) ribbed mericarp, (c) pale pink corolla with ratio other genera within the Malveae tribe of of 1:1 with calyx. Malvaceae by its characteristic triphyllum or three non-fused epicalyx bracts (Ray 1998). In particular, this character has been used to distinguish Malva from Lavatera, no longer exists and has been replaced. species have become widespread weeds a closely resembling genus with connate Bentham (1863) noted that four Europe- (Ray 1995). The centres of diversity are epicalyx bracts. However, it has been sug- an Malva species, M. rotundifolia L. (also most likely to be in the Mediterranean re- gested that the use of epicalyx characters known as M. pusilla Sm.), M. parvifl ora, M. gion and south-western Asia, extending to separate Malva and Lavatera is unten- verticillata L. and M. sylvestris L., were natu- as far as Turkmenistan and Afghanistan, able based on morphological and molecu- ralized as weeds in some parts of Australia (Hanf 1983, Jessop and Toelken 1986, Ma- lar analysis of the nuclear ribosomal-DNA and he described their diagnostic charac- kowski and Morrison 1989, Mitich 1990). Internal Transcribed Spacer region (Ray teristics (Barker 1977). In 1889, Schom- Malva has naturalized in many countries 1998). Recently, as a result of this genetic burgk observed that the four species had including Australia, Finland, Denmark, analysis, several native Australian Lavatera escaped from gardens in South Australia Korea, Japan, Argentina, Chile, India, Le- species have been reassigned to Malva. and ‘established themselves as in the old sotho, Namibia, Zimbabwe, South Africa Within the Malva genus, species which country, about hedges, roadsides, and in and the United States (Naqshi et al. 1988, have petals less than 12 mm long are regu- cultivated, as well as in waste grounds and Randall 2002, Hinsley 2007, USDA 2008). larly misidentifi ed and have proven diffi - pastures lands’ (Schomburgk 1889). It is It also occurs abundantly in New Zealand cult to distinguish. These include M. parvi- highly likely that M. parvifl ora was intro- (Low 1991). fl ora, M. pusilla Sm., M. nicaeensis All. and duced intentionally as a garden plant as The number of Malva species currently M. neglecta Wallr. Taxonomists frequently it was cultivated in Europe as a salad veg- present in Australia varies between differ- mistake M. parvifl ora for M. neglecta or M. etable and a ‘green manure’ (Dalby 1968, ent sources, with estimates of nine (Coun- pusilla in Canada (Makowski and Morri- Barker 1977). In 1922, it was recorded as cil of Heads of Australian Herbaria 2008), son 1989) and all M. parvifl ora samples in being present at high ‘luxurious’ densities six (Walsh and Entwisle 1996) and fi ve the Herbarium at the University of Kash- in agricultural areas of New South Wales, (Hnatiuk, 1990). Malva parvifl ora is thought mir, India have turned out to be M. neglecta where it was thought to have caused the to be the most common and widespread (Naqshi et al. 1988). However, several char- potentially fatal ‘staggers’ syndrome naturalized Malva species in Australia acteristics distinguish the small-fl owered in livestock (Dodd and Henry 1922). M. (Council of Heads of Australian Herbaria mallows (Table 1, Figure 3). parvifl ora is now naturalized in all states 2008) (Figure 4). It is more prevalent in the of Australia (Barker 1977, Hnatiuk 1990, southern half of Australia where climatic History Low 1991, Auld and Medd 1992, Lazarides conditions are temperate, which is con- The fi rst known reference of Malva in Aus- et al. 1997, Ray 1998, Council of Heads of sistent with its supposed Mediterranean tralia was in 1845 (Miquel 1845) with a de- Australian Herbaria 2008). origin. Its presence in humid tropical areas scription of a species called M. preissiana of Australia as well as its world wide dis- Miq. However, there is no further mention Distribution tribution indicates M. parvifl ora is highly of M. preissiana in any literary source ex- The natural origin of species within the adaptable to an extensive range of climatic amined, suggesting that the species name genus Malva is uncertain because many conditions. 4 Plant Protection Quarterly Vol.24(1) 2009 depths is 60%, with buried seeds produc- ing at least 13% more emergence than seeds on the surface. Seedling emergence decreases progressively as seeds are buried deeper than 2 cm, with no emergence occurring at 8 cm or deeper (Chauhan et al. 2006a). More seedlings emerge under no- tillage, when soil disturbance is limited to the sowing operation only, than minimum tillage systems, where there were two pre- sowing cultivations in addition to sowing (Chauhan et al. 2006b).

Reproduction Floral biology Malva parviflora plants can reproduce within two months of germinating. The fi rst fl owers open 51 days after germination (Michael et al. 2007), and reproduction occurs throughout the life of the plant (Cunningham and Mulham 1992). In Aus- tralia fl owering occurs from March (early autumn) through to December (early summer) (Western Australian Herbarium 2008). Flowers are bisexual and contain 10 anther sacs, 10 ovules and a central feath- Figure 4. Distribution of Malva parvifl ora in Australia. Data sourced from ery stigma. M. parvifl ora is predominately the Australian Virtual Herbarium (Council of Heads of Australian Herbaria inbreeding. Each pollen sac contains 31 2008). ±1.3 pollen grains, giving a pollen-ovule ratio of 31 and hence a very limited opportunity for outcrossing. Upon fl ower opening, anthers have already dehisced and Habitat either prostrate or erect, with prostrate pollen grains are attached to the stigma, Malva parvifl ora is widespread through- plants being generally associated with indicating prior self-pollination. In addi- out Australia, particularly the southern disturbed areas of frequent trampling or tion fl owers are small, unperfumed and half, and occurs over an extensive range mowing (Barker 1977). Erect plants can open for only a few days, and thus un- of climatic regions (Figure 4). Plants can grow up to 1.2 m high (Lamp and Col- likely to attract insect pollinators (Michael grow in areas with average rainfall as low let 1984) and 2.1 m wide (Michael 2006) et al. 2006b). as 315 mm (Merredin, Western Australia) and predominantly occur when resources (Michael et al. 2006b) and as high as 950 such as light and space are limited through Seed production and dispersal mm (Rockhampton, Queensland) (Coun- plant competition (i.e. livestock yards or Malva parvifl ora produces approximately cil of Heads of Australian Herbaria 2008). cropping situations) (P. Michael personal 10 seeds per fl ower (Michael et al. 2007). Plants can also grow in a broad range of observation). In cropping situations, M. Although there have been no studies of soil conditions including rocky or stony pusilla, a biologically similar species with total seed production per plant, closely soil, sand, loam and clay (Michael et al. the same chromosome number (2n = 42), related species from the Malveae tribe of 2006b, Western Australian Herbarium exhibits a more erect growth habit and Malvaceae produce many seeds. Produc- 2008). Although not usually considered grows to the height of the crop, but when tion of seeds from Abutilon theophrasti a halophyte, M. parvifl ora inhabits saline alone it spreads over the ground with Medik. ranges from 7000 to 17 000 seeds soils in Bahrain (Kivilaan and Bandurski branches over 1 m in length (Makowski (Winter 1960, Warwick and Black 1985). 1981) and parts of the Western Australian and Morrison 1989). The presence of wild Malva pusilla propagates exclusively by (WA) coastline (Western Australian Her- mustard, Brassica kaber (DC.) Wheeler, seeds (Makowski 1987, Makowski and barium 2008). It is most commonly found appears to inhibit the growth of M. parvi- Morrison 1989) and can produce between in disturbed ground and pastures of farm- fl ora under natural conditions, most likely 1000 to 5000 seeds per plant in a pure ing land, although it can occur in other ar- caused by allelopathy (Vicol and Dobrota stand and an average of around 300 seeds eas such as gardens, crops, roadsides and 1995). per plant when in competition with wheat wastelands (Barker 1977, Lamp and Collet Several sources state that seedlings (Pyasyatskene 1978, Carlson and Eberlein 1984, Auld and Medd 1992, Lazarides et emerge in autumn, indicating that the weed 1983). Given the large number of fl owers al. 1997, Western Australian Herbarium is a winter growing annual (Lazarides et on plants at any given time, the long pe- 2008). al. 1997, Michael et al. 2006a); however, riod of fl owering and high seed number M. parvifl ora can emerge throughout the per fl ower, seed production per plant of Growth and development year in some climates (Chorbadjian and M. parvifl ora is also likely to be in the thou- There has been limited study of the growth, Kogan 2002). In cropping systems, most sands. development or phenology of M. parvifl ora seedlings of M. parvifl ora emerge within Malva parvifl ora seeds can be dispersed in Australia. However, under controlled two weeks of crop sowing (Chauhan et by biotic vectors, particularly agricultural fi eld conditions, dry weight of M. parvi- al. 2006a). Seeds buried at depths of 2 cm livestock. Up to 700 viable M. parvifl ora fl ora plants reaches a maximum of 340 g (Michael et al. 2006a) or 0.5–2 cm (Chauhan seeds can be passed daily through a horse (Michael et al. 2006b). Plants have a single, et al. 2006a) in the soil have higher emer- with recovery from manure peaking three long taproot that allows them to survive gence than seeds placed on the surface. to fi ve days after seed consumption and long periods of drought. They can grow Maximum seedling emergence from these gradually declining until 13 days, when Plant Protection Quarterly Vol.24(1) 2009 5 no further seeds are recovered (St John- after-ripening (Michael et al. 2007). Physi- tetrazolium staining. However, the seed Sweeting and Morris 1990). Hardseeded ological dormancy can also be partially coat was apparently not artifi cially scari- viable seeds (~20% of total consumed) can overcome using growth promoters, with fi ed so physical dormancy may still have be passed intact through sheep, with the 5 mM of KNO3 increasing germination by been present. Alternatively, as embryos majority being excreted within three days, 18% and 1 mM GA3 by 13% (Chauhan et become less vigorous with increasing age, although some seeds were recovered up al. 2006a), in seeds scarifi ed with 95% sul- there may have been insuffi cient time al- to seven days after initial seed digestion phuric acid for 60 minutes. Physiological lowed for germination or they may not (Michael et al. 2006c). As livestock are reg- dormancy occurs in other Malvaceae spe- have been able to exert suffi cient force to ularly moved within and between indi- cies (Egley and Chandler 1978, Warwick emerge from the seed coat. Further infor- vidual farms and agricultural regions, the and Black 1985, Baskin and Baskin 1998) mation on the longevity of M. parvifl ora is potential dispersal of seeds is vast. Seeds and in conjunction with physical dorman- lacking but viable seeds of M. rotundifolia can also be dispersed long distances by cy is known as combinational dormancy have been unearthed after 100 years (Kivi- birds (Proctor 1968), although no Austral- (Baskin and Baskin 2004). laan and Bandurski 1981) and 120 years ian studies have been conducted. Non-dormant seeds are able to germi- (Telewski and Zeevaart 2002) of burial nate over a wide range of temperatures in moist aerated soil, confi rming the ex- Physiology of seeds and germination (3.3–37°C) and a range of pH (pH 4–10) tensive persistence of species within the The main type of dormancy impeding ger- and have no light requirement (Chorbad- Malva genus. This suggests that M. parvi- mination in M. parvifl ora seeds is physical jian and Kogan 2004, Chauhan et al. 2006a, flora could potentially have seed bank dormancy, i.e. an impermeable seedcoat Michael et al. 2006a). Germination, howev- persistence of several decades in the event that prevents seeds from imbibing water er, is optimal at temperatures typically as- that fl uctuations in temperature are not (Michael et al. 2006a). The tissue respon- sociated with winter rainfall in Mediterra- enough to break physical dormancy. sible for physical dormancy (also known nean-type climates (15–20°C) (Muniz 2000, as hardseededness) in Malvaceae seeds Chorbadjian and Kogan 2004, Michael et Hybrids is a layer of palisade cells that develops al. 2006a). Germination declines as NaCl A hybrid between M. neglecta and M. parvi- from the exotegmen, the outer epidermis concentration increases; however, a small fl ora was recorded in 1932, but there is no of the inner integument (Corner 1976). In number of seeds can germinate at 160 mM other record of this crossing nor of any the fi eld, dormancy release occurs during NaCl (Chauhan et al. 2006a), indicating since (Dalby 1975). Dalby (1968) report- the summer months in response to natural the potential for salt tolerance in some ed the existence of other Malva hybrids fl uctuations in temperature. Replication populations. Germination is completely between M. pusilla × M. neglecta, M. alcea of summer temperatures in the laboratory inhibited at osmotic potentials of −0.6 to L. × M. moschate L. and M. sylvestris × M. confi rms that these fl uctuations are impor- −1 MPa, suggesting moderate sensitivity neglecta in Europe. Yet again, there are no tant, as dry seeds lose dormancy under al- to osmotic stress (Chauhan et al. 2006a). other records of these crossings and it is ternating 50/20°C but not under constant Changes in seed germinability and dor- not stated whether these were natural or temperatures (Michael et al. 2006a). Scan- mancy during development on the parent artifi cial. Dalby (1968) suggested that the ning electron microscopy of the seed coat plant have been comprehensively studied apparent variability of some Malva species shows structural differences in the chala- in M. parvifl ora (Michael et al. 2007). Seed is due, at least in part, to hybridization. zal region of permeable and impermeable moisture content decreases as seeds de- However, due to the lack of other records seeds, indicating the importance of this re- velop, whereas fresh (maximum 296 mg) of natural Malva hybrids, it is unclear as gion in the physical dormancy breakdown and dry weight (maximum 212 mg) in- to whether these ‘hybrids’ were real or a of M. parvifl ora seeds (Michael et al. 2006a). crease to a peak at 12–15 and 21 days after case of misidentifi cation. In experiments The structure of this chalazal region, a slit- fl owering (DAF) respectively. Thus, physi- with artifi cial Malva crosses, no spontane- like discontinuity, is consistent with that ological maturity occurs at 21 DAF, when ous hybrids occur, though some artifi cial observed in other species belonging to the seed moisture content is 16–21%. Seeds are crosses produce slightly fertile offspring Malveae group of Malvaceae, confi rming capable of germinating early in develop- (i.e. M. pusilla × M. neglecta, M. parvifl ora × the signifi cance of the chalazal region in ment, reaching 63% at 9 DAF, but germi- M. neglecta) (Kristofferson 1926). There is regulating physical dormancy of seeds nation declines as development continues, no evidence for hybridization in Canadian within the Malva genus. Bypassing natural presumably due to imposition of physi- collections of Malva species (Makowski dormancy release is possible by physical ological dormancy. Physical dormancy de- and Morrison 1989). Malva sylvestris pollen scarifi cation of the seedcoat using tweez- velops at or after physiological maturity, placed on the stigma of Gossypium L. (cot- ers (Sumner and Cobb 1967, Chauhan et once seed moisture content declines below ton) does not germinate at all (Kamalova al. 2006a, Michael et al. 2006a) or a scalpel 20%. Seeds are able to tolerate desiccation 1985) indicating that intergeneric hybrids (Sumner and Cobb 1967, Chauhan et al. from 18 DAF; desiccation hastens develop- are also very unlikely. Furthermore, as M. 2006a) to allow imbibition. Chemical scari- ment of physical dormancy and improves parviflora is predominantly inbreeding, fi cation has variable results on germina- germination, indicating that weed control with self-pollination occurring prior to tion of hardseeded M. parvifl ora. Placing measures imposed after fl owering are un- fl ower opening, hybrids are not expected seeds in 71% sulphuric acid or 5% sodium likely to prevent viable seeds entering the to form easily (Michael et al. 2006b). hypochlorite for up to 18 hours does not soil seed bank. enable imbibition, but 95% sulphuric acid Malva parvifl ora seeds can remain viable Population dynamics for 60 minutes or boiling seeds in water for many years in soil without germina- Malva parvifl ora is morphologically vari- for 5–10 seconds followed by immersion tion if their physical dormancy is retained. able between geographical locations. in ice water is partially effective (Sumner Seeds of M. parvifl ora were discovered in Plants collected from northern popula- and Cobb 1967). bricks of adobe buildings built between tions within the wheatbelt of WA (north In addition to physical dormancy, mi- 1769 and 1837 in California and New of 29°22’S) fl ower earlier, have shorter nor physiological dormancy is observed Mexico, a dry environment with reduced calyxes, are narrower at fl owering and in freshly matured M. parvifl ora seeds, temperature fluctuation and microbial have heavier seeds at senescence than as scarifi ed seeds are slow to germinate activity (Spira and Wagner 1983). Seeds populations from more southern popula- (Sumner and Cobb 1967, Michael et al. extracted from these bricks and placed on tions (Michael et al. 2006b). Two popula- 2006a). Release of physiological dorman- agar for 19 days did not germinate, but a tions from the western side of the wheat- cy naturally occurs after a short period of third of the seeds were viable according to belt (Mingenew and Wagin), which has 6 Plant Protection Quarterly Vol.24(1) 2009 higher rainfall, lose physical dormancy So although potentially toxic, M. parvifl ora and the Mediterranean region (Abousabaa more slowly than populations from the poses only a small risk to livestock if the 2001). Analysis of M. parvifl ora leaves indi- eastern side (Morawa and Hyden), which ‘staggers’ condition is recognized early. cate that they are high in protein (36.2%), has lower rainfall (Michael et al. 2006a). However, if the animal is continuously carbohydrates (12.4%) and soluble fi bre In South Australia, pedicel length var- pushed it may die. It is not known what (4.2%) (Ranhotra et al. 1998) and seeds ies considerably between plants (Barker causes ‘staggers’ although researchers contain considerable amounts of anti- 1977). In Jordan, growth is dependent on have suggested that plant nitrate content oxidants (Abousabaa 2001). M. parvifl ora the climatic region in which plants grow may be a contributing factor (Webb 1948, has also been used in traditional medicine (Eideh and Elkarmi 2005). Plants growing Gardner and Bennetts 1956, Kingsbury for hundreds of years for its anti-bacterial in the cool region grew taller in preference 1964, Everist 1974). Consumption of M. properties (Grierson and Afolayan 1999). to increasing stem width, leaf length, leaf parvifl ora can also be detrimental to poul- Recently it has been discovered that M. width and petiole length, whilst plants try. Sterculic and malvalic acid, fatty acids parvifl ora contains important anti-bacteri- growing in the warm region exhibited the found in M. parvifl ora plants (Shenstone al and anti-fungal proteins and genes for opposite, with plant height increase being and Vickery 1959, Ahmad et al. 1984), pro- these could potentially be transferred to lower. duce the typical symptoms of ‘pink-white improve resistance of otherwise pathogen- condition’ in the whites of poultry eggs. sensitive plants (Wang and Bunkers 2000, Importance The disorder produces a mottled, discol- Wang et al. 2001, Shale et al. 2004, 2005). Detrimental oured yolk, a pasty condition when eggs Four potent anti-fungal proteins purifi ed There has been little research into the com- are kept at low temperatures, and higher from seeds were found to severely in- petitive effects of M. parvifl ora; however than normal yolk water content (Shenstone hibit the activities of Fusarium graminearu Malva species are very good competitors and Vickery 1959). This condition has se- Schwabe and Phytophthora infestans (Mont.) with both agricultural (Makowski and vere commercial consequences through a de Bary (Wang and Bunkers 2000, Wang et Morrison 1989, Makowski 1995) and hor- reduction in overall egg quality. al. 2001). Hexane, methanol and water ex- ticultural crops (Agamalian and Kurtz Malva parvifl ora acts as host for many tracts made from M. parvifl ora with a pros- 1989, Abusteit and Shehata 1993) as well as pests and diseases including the South Af- trate growth form inhibit the growth of other weed species such as wild oats (Ave- rican cassava mosaic virus (Berrie 2001), Gram-positive and Gram-negative bacte- na ludoviciana Durieu) (Bilalis et al. 2001). the faba bean necrotic yellow virus (Al- ria, while extracts made from plants with In particular Malva species are especially Nsour et al. 1998), cotton tipworm (Croci- an upright growth form inhibit the growth competitive in minimum tillage systems dosema plebejana Zell.) (Hamilton and Gage of Gram-positive bacteria only (Shale et al. where tap roots are allowed to grow freely. 1986), rust fungus (Puccinia malvacearum 2005). Antibacterial activity of M. parvi- Malva pusilla has been studied extensively Bertero ex Mont.) (Elarosi et al. 1974), the fl ora (leaves and roots) initially increases in this regard. It is listed as a noxious weed thrip Frankliniella occidentalis Perg. (Bautis- with storage, with the highest inhibitory in Canada (Makowski and Morrison 1989), ta et al. 1995) and root lesion nematodes activity being measured after 3–6 months where it is a major weed of less competi- (Pratylenchus neglectus L. and P. thornei L.) storage, after which activity gradually de- tive fi eld crops such as lentil and fl ax (Ma- (Vanstone and Russ 2001). As the weed can creased (Shale et al. 2004). kowski and Mortensen 1992). It doubled emerge all year round (Chorbadjian and Whilst M. parvifl ora is host to many in abundance on cultivated land between Kogan 2002), it is likely to be able to host pathogens and diseases, it also provides 1980 and 1985 in Alberta, Canada (Ma- biotrophic fungi, such as powdery mildew shelter, food and refuge to benefi cial pred- kowski 1995). It is capable of substantially and rusts. An Egyptian study recorded ators such as Amblyseius victoriensis Wom., reducing crop yields and quality, particu- the presence of 39 different fungal species a predatory mite found in peach orchards larly if it emerges before the crop (Friesen from 20 genera on a single M. parvifl ora of NSW (James 1989, 1990) and Coccinel- et al. 1992). In Canada, wheat yields are re- plant (Abdel-Hafez and El Naggar 2001). la arcuata Fabr. (beetle), Nabis capsiformis duced by 30% at densities of 139 plants m−2 These pests compound the consequences Germ. (pale damsel bug) and Deraeocoris and densities of 39 plants m−2 reduce the of M. parvifl ora infestation. signatus Dist. (brown smudge bug) found yield of fl ax by more than 90% (Makowski in cotton fi elds of south-east Queensland and Morrison 1989). Malva pusilla remains Benefi cial (O’Brien 1978). So whilst caution should green in the autumn and causes problems In certain countries, M. parvifl ora is a use- be exercised in its use, M. parvifl ora has by interfering with both the threshing and ful and valuable plant species. It has been many benefi cial attributes. cleaning processes of the harvester (Ma- used as feed for poultry (Asar et al. 1972), kowski and Morrison 1989). It is also a carp fi sh (Labib et al. 1994) and grazing Legislation problem as a crop seed contaminant. The animals (Guerrero 1999). Malva parvifl ora Malva parvifl ora is not currently a declared highly competitive nature of Malva spe- has chemical feed quality attributes com- or noxious weed in any Australian state cies throughout the world suggests that parable to alfalfa, with weed-infested pad- or territory (Australian Weeds Committee M. parvifl ora also has the potential to cause docks in eastern California able to sustain 2008) and there is no legislation governing signifi cant detriment to agricultural sys- more lamb grazing days and produce more its movement or control. tems within Australia. kilograms of lamb gain per hectare of land Consumption of M. parvifl ora can be than weed-free paddocks (Guerrero 1999). Weed management harmful to livestock. In Australia there is There is the opportunity for M. parvifl ora Herbicides evidence that ingestion of very large quan- to be marketed as a livestock food source There are currently 192 herbicide products tities of M. parvifl ora (1.4–5.5 kg day−1) in as long as the potential for ‘staggers’ is containing 54 active ingredients registered addition to hard driving can cause ‘stag- taken into account by ensuring livestock for use on M. parvifl ora, including paraquat gers’ in sheep, horses and cattle (Dodd are not unduly exerted or pushed. (135 g L−1) and diquat (115 g L−1), trifl uralin and Henry 1922, Hurst 1942, Everist 1974, Malva parvifl ora is an important part (400 g L−1), simazine (500 g L−1), diuron (500 Main and Butler 2006). Affected animals of the human diet and provides essential g L−1) and glyphosate (450 g L−1) (APVMA cannot walk and display trembling and nutrients for many underprivileged peo- 2008). In order to be registered, herbicides quivering of the limbs, shallow breathing ple in countries such as Morocco, where must provide evidence of reasonable weed and rapid pulse (Everist 1974). If the ani- only the young shoots are eaten, (Tanji control. However, control of M. parvifl ora mal is allowed to rest, the condition will and Nassif 1995), Egypt (Badawi and El- with selective herbicides and cultivation pass and it will recover in a few hours. Sahhar 1978), Mexico (Ranhotra et al. 1998) is often unsatisfactory (Blackshaw 1996), Plant Protection Quarterly Vol.24(1) 2009 7 facilitating its spread once initial infesta- et al. 1987, Norton et al. 1989, Gardener Food and Agriculture 35, 408-14. tions become established. Malva parvifl ora et al. 1993a,b, Squella and Carter 1996), Al-Nsour, A., Mansour, A., Al-Musa, A. is naturally tolerant to glyphosate (Group with hardseeded small seeds most likely and Salem, N. (1998). Distribution and M) (Wu and Dasgheib 2001). Glyphosate to survive digestion. The hardseededness incidence of faba bean necrotic yel- is a non-selective, foliar applied herbicide of Malva species, as well as small size, allows virus in Jordan. Plant Pathology 47, which has been extensively used around low a substantial proportion of seeds to 510-5. the world for the past 20 years. It gives survive mastication and digestion. Most APVMA (2008). Australian Pesticides and only partial control of Malva species (>92%) physically dormant seeds placed Veterinary Medicines Authority. www. (Dastgheib and Frampton 2000, Wu and into a sheep rumen survive up to 48 hours apvma.gov.au/; accessed 1 February Dasgheib 2001, Chorbadjian and Kogan of digestion, whilst only 1.4% of scarifi ed 2008. 2002). It is well known that the activity non-physically dormant seeds survive di- Asar, M.A. (1972). Effect of feeding some of glyphosate can be antagonized by the gestion for 12 hours (Michael et al. 2006c). weeds to growing chicks. Alexandria presence of polyvalent cations (Stahlman The hard, impermeable seed coat of M. Journal of Agricultural Research 20, 14. and Phillips 1979, Nalewaja and Matysiak parvifl ora protects the embryo from chemi- Auld, B.A. and Medd, R.W. (1992). ‘Weeds: 1991) that lead to the formation of poorly cal and enzymatic scarifi cation (Sumner an illustrated botanical guide to weeds soluble glyphosate salts that are less likely and Cobb 1967), which enables seed sur- of Australia’. (Inkata Press, Mel- to be taken up by the plant. Microanalysis vival during rumen digestion. bourne). of the leaf surface of Abutilon theophrasti, However, when dormant seeds were Australian Weeds Committee (2008). Nox- from a closely related genus and a member fed to sheep and faeces collected, the ious weed list for Australian States and of the Malveae group, indicates the pres- large majority (82%) were killed through Territories (Database). http://www. ence of two antagonistic elements, magne- mastication, digestion and gut passage weeds.org.au/noxious.htm; accessed 1 sium and calcium (Hall et al. 2000). Further (Michael et al. 2006c). The proportion of March 2008. analysis led to the discovery of specialized hardseededness within any M. parvifl ora Badawi, M.A. and El-Sahhar, K.F. (1978). trichomes (chalk glands) that secrete these weed population depends largely on plant Effect of gibberellin on some vegetative cations. Therefore, the glyphosate toler- and seed development stage (Michael et al. characters of Malva parvifl ora. Research ance of M. parvifl ora may be due to spe- 2007) so the risk of spreading viable seeds Bulletin Faculty of Agriculture Ain Shams cialized structures, such as trichomes, and is very low if sheep graze only on imma- University, 16. further investigation is required. ture plants and seeds. The palatability of Barker, W.R. (1977). The Species of Malva This tolerance to glyphosate can be M. parvifl ora to sheep at different plant and L. and Lavatera L. (Malvaceae) natural- reduced by the addition of certain herbi- seed development stages is unknown and ized in South Australia. Journal of Ad- cides, which when mixed together, pro- research is needed. elaide’s Botany Garden 1, 107-14. duce a synergistic interaction that increas- Baskin, C.C. and Baskin, J.M. (1998). ‘Seeds: es the effi ciency of glyphosate or main- Natural enemies ecology, biogeography and evolution of tains its optimum activity at lower rates. Whilst there has been little research on dormancy and germination’. (Academ- Such herbicides include lactofen (Wells natural enemies of M. parvifl ora and thus ic Press, Lexington, Kentucky). and Appleby 1992), a diphenyl ether her- the prospect of using biological control in Baskin, J.M. and Baskin, C.C. (2004). A bicide that inhibits the production of pro- Australia is unknown, the bioherbicide classifi cation system for seed dorman- toporphyringogen oxidase (Group G), and BioMal™ (derived from the fungi Colle- cy. Seed Science Research 14, 1-16. fl uroxypyr (Chorbadjian and Kogan 2002), totrichum gloeosporioides Penz.) is success- Bautista, R.C., Mau, R.F.L., Cho, J.J. and which disrupts plant cell growth (Group fully used to control M. pusilla in Canada Custer, D.M. (1995). Potential of tomato I). Directed spray applications of undi- (Zimdahl 1999). spotted wilt tospovirus plant nests in luted ammonium nitrogen fertilizer cause Hawaii as virus reservoirs for transmis- 99% control at the 1–4 leaf stage and 77% References sion by Frankliniella occidentalis (Thysa- at the 5–7 leaf stage (Agamalian 1991). Abdel-Hafez, S.I.I. and El Naggar, S.M. noptera, Thripidae). Phytopathology 85, (2001). Fungal biodiversity, pollen mor- 953-8. Other treatments phology and leaf surface of some na- Bentham, G. (1863). Malvaceae. Flora Aus- Malva parvifl ora plants have a substantial tive species in Egypt. Feddes Repertorium traliensis I, 197-221. taproot and they are thus diffi cult to kill. 112, 127-40. Berrie, L.C., Rybicki, E.P., Rey, M.E.C. Plants are able to regrow after being cut Abousabaa, K.E. (2001). Nutritional as- (2001). Complete nucleotide sequence down or eaten to the crown area, though pects of wild plants, nutritional compo- and host range of South African cassava cultivation can kill M. parvifl ora if the tap- sition of Malva parvifl ora and Sisybrium mosaic virus: further evidence for re- root is severed below the crown (Chor- irio. CIHEAM-IAMC 6, 241. combinations amongst begomoviruses. badjian and Kogan 2002). Mowing and Abusteit, E.O. and Shehata, S.A. (1993). Journal of General Virology 82, 53-8. grazing of M. pusilla will delay growth for Critical period of weed competition in Bidack, L. and Brandham, P.E. (1995). In- a short time, followed by rapid recovery potatoes (Solanum tuberosum L.) sum- traspecifi c uniformity of chromosome and increased branching below the injured mer plantation. 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