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Extraction from CABI CPC 2002 Edn prepared by Alan MacLeod, CSL, UK

Selected texts for Listroderes costirostris CABI CPC 2002 Edn



NAMES AND TAXONOMY

Preferred Name


Listroderes costirostris Schönherr
Taxonomic Position

Domain: Eukaryota

Kingdom: Metazoa

Phylum: Arthropoda

Class: Insecta

Order: Coleoptera

Family: Curculionidae
Other Names Used

Desiantha nociva French

Listroderes difficilis Germain

Listroderes hypocritus Hustache

Listroderes lugubris Germain

Listroderes obliquus Klug

Listroderes paranensis Hustache

Listroderes vicinus Hustache


BAYER CODE: LISTCO

Common Names


English:

Australian tomato weevil

brown vegetable weevil

buff-colored tomato weevil

carrot weevil

dirt-colored weevil

turnip weevil

vegetable weevil

weevil, Australian tomato

weevil, buff-colored tomato

weevil, vegetable


NOTES ON TAXONOMY AND NOMENCLATURE


Listroderes is a Neotropical genus comprising about 66 described species (Wibmer and O'Brien, 1986).
L. costirostris was first described from Rio de Janeiro, Brazil. Morrone (1993) redescribed this species and provided a description of a male, indicating that this weevil is not always parthenogenetic.
There is some confusion concerning synonyms of L. costirostris. L. obliquus Klug has long been considered a synonym (CIE, 1964). However, according to Wibmer and O'Brien (1986), L. obliquus Klug is considered a valid species. May (1994) recognized L. costirostris and L. obliquus as synonyms of L. difficilis Germain, whilst Wibmer and O'Brien (1986) recognized L. difficilis as a valid species. However, O'Brien and Wibmer (1982) listed L. obliquus and L. difficilis as synonyms of L. costirostris.
Morrone (1993) listed the full synonymy for L. costirostris provided in the tabular data, where L. obliquus and L. difficilis are recognized as synonyms of L. costirostris. However, he stated that there will remain some doubt as to what the synonyms of L. costirostris truly represent until biological and chromosomal data have been assembled.


HOST RANGE


L. costirostris is a polyphagous pest, primarily affecting vegetables, but can also feed on flowers and many wild hosts (High, 1939).
In Western Australia, potatoes, tomatoes and root crops are the preferred food plants; although most vegetables are attacked during the winter; peas, beans and pumpkins are relatively immune (Jenkins, 1944).
In Uruguay, Parker (1950) reported that adults fed on young potato crops, whilst the larvae fed principally on chickweed, Stellaria media (Parker et al., 1950).
In Queensland, L. costirostris is a pest of tobacco in seed beds (Broadley, 1975).
L. costirostris was found attacking garland chrysanthemum (Chrysanthemum coronarium) and other winter vegetables, including Gynura bicolor and various brassica crops in Taiwan (Hsu and Chiang, 1983).
Principal weeds hosts include capeweed (Cryptostemma calendulacea and Arctotheca calendula), chickweed, dandelion, milkthistle (Sonchus), wild radish, wild parsnip and marsh mallow (Malva rotundifolia).

Primary hosts: polifago (polyphagous), Allium cepa (onion), Allium sativum (garlic), Arachis hypogaea (groundnut), Beta vulgaris (beetroot), Brassica napus var. napus (rape), Brassica oleracea (cabbages, cauliflowers), Chrysanthemum coronarium, Cichorium endivia (endives), Capsicum annuum (bell pepper), Daucus carota (carrot), Lactuca sativa (lettuce), Lycopersicon esculentum (tomato), Nicotiana tabacum (tobacco), Petroselinum crispum (parsley), Pastinaca sativa (parsnip), Raphanus raphanistrum (charlock), Solanum melongena (aubergine), Solanum tuberosum (potato), Spinacia oleracea (spinach).


Secondary hosts: Gynura bicolor, Papaver (poppies), Petunia hybrida, Phlox, Viola wittrockiana hybrids.
Wild hosts: Buddleia (Butterflybush), Cirsium spp., Eupatorium, Geranium carolinianum (Carolina geranium), Helenium (sneezeweeds), Linaria (Toadflax), Malva (mallow), Morus alba (mora), Oxalis (wood sorrels), Plantago (Plantain), Rumex (Dock), Scutellaria (Skullcap), Sonchus (Milkthistle), Verbena, Veronica peregrina (American speedwell), Stellaria media (chickweed).
Affected Plant Stages: Seedling stage, and vegetative growing stage.
Affected Plant Parts: Leaves, roots, stems, and whole plant.


GEOGRAPHIC DISTRIBUTION

Notes on distribution


L. costirostris is a native of South America and was first discovered in the USA (Mississippi) in 1922, and in California during 1925 (High, 1939). It is the most widespread species of the genus Listroderes, and has been accidentally introduced into Australia, New Zealand, Spain, South Africa, USA, Japan and Easter Island (Morrone, 1993).
In the USA, it now occurs in the Gulf and southern states and in Oklahoma, Arizona, and California. In North Carolina, the vegetable weevil occurs throughout the state but is generally more common in the southern Coastal Plain.
L. costirostris has probably been established in New Zealand since the 1930s as Muggeridge (1933) reported it being present in widely separated localities. It was found in Australia for the first time in Victoria in 1905, and is now widespread and intermittently injurious in all the States, especially Queensland and New South Wales (Wilson and Wearne, 1962).
The distribution map includes records based on specimens of L. costirostris from the collection in the Natural History Museum (London, UK): dates of collection are noted in the List of countries (NHM, various dates).


List of countries


Europe

France: present, few occurrences (CABI/EPPO, 2000; EPPO, 2002)

Portugal: present, no further details (Balachowsky, 1963; CABI/EPPO, 2000; EPPO, 2002)

Spain: restricted distribution (Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Balearic Islands: present, no further details (Balachowsky, 1963; CABI/EPPO, 2000; EPPO, 2002)

Canary Islands: present, no further details (Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)


Asia

[China]


Taiwan: present, no further details (Hsu & Chiang, 1983; CABI/EPPO, 2000; EPPO, 2002)

Japan: present, no further details (Matsumoto, 1962; Morrone, 1993; CABI/EPPO, 2000)

Honshu: present, no further details (Shiraki, 1952; Morimoto, 1992; CABI/EPPO, 2000; EPPO, 2002)

Kyushu: present, no further details (Yokomizi & Nagano, 1979; Morimoto, 1992; CABI/EPPO, 2000; EPPO, 2002)

Ryukyu Archipelago: present, no further details (Morimoto, 1992; CABI/EPPO, 2000; EPPO, 2002)

Shikoku: present, no further details (Morimoto, 1992; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Korea, Republic of: present, no further details (Morimoto, 1992; CABI/EPPO, 2000; EPPO, 2002)
Africa

Morocco: present, no further details (Balachowsky, 1963; CABI/EPPO, 2000; EPPO, 2002)

South Africa: present, no further details (Brain, 1929; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)
Western Hemisphere

Argentina: widespread (Bosq, 1942; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Bolivia: present, no further details (Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Brazil: present, no further details (Buchanan, 1937; Morrone, 1993; CABI/EPPO, 2000)

Bahia: present, no further details (NHM, 1905; CABI/EPPO, 2000; EPPO, 2002)

Parana: present, no further details (Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Santa Catarina: present, no further details (Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Chile: widespread (Olalquiaga, 1961; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Easter Island: present, no further details (Morrone, 1993; CABI/EPPO, 2000)

Paraguay: present, no further details (Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

USA: restricted distribution (CABI/EPPO, 2000)

Alabama: widespread (High, 1939; CABI/EPPO, 2000; EPPO, 2002)

Arizona: restricted distribution (O'Brien & Wibmer, 1982; CABI/EPPO, 2000; EPPO, 2002)

Arkansas: widespread (High, 1939; O'Brien & Wibmer, 1982; CABI/EPPO, 2000; EPPO, 2002)

California: restricted distribution (High, 1939; O'Brien & Wibmer, 1982; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Florida: restricted distribution (High, 1939; Tappan, 1974; O'Brien & Wibmer, 1982; CABI/EPPO, 2000; EPPO, 2002)

Georgia (USA): widespread (High, 1939; O'Brien & Wibmer, 1982; CABI/EPPO, 2000; EPPO, 2002)

Hawaii: present, no further details (Buchanan, 1937; Pemberton, 1947; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Kentucky: restricted distribution (CIE, 1964; CABI/EPPO, 2000; EPPO, 2002)

Louisiana: widespread (High, 1939; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Maryland: restricted distribution (CIE, 1964; CABI/EPPO, 2000; EPPO, 2002)

Mississippi: widespread (High 1939; O'Brien & Wibmer, 1982; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Missouri: restricted distribution (CIE, 1964; CABI/EPPO, 2000; EPPO, 2002)

New Jersey: present, no further details (Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

North Carolina: widespread (O'Brien & Wibmer, 1982; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Oklahoma: restricted distribution (CIE, 1964; CABI/EPPO, 2000; EPPO, 2002)

Oregon: present, no further details (Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

South Carolina: widespread (High, 1939; O'Brien & Wibmer, 1982; CABI/EPPO, 2000; EPPO, 2002)

Tennessee: widespread (High, 1939; CABI/EPPO, 2000; EPPO, 2002)

Texas: restricted distribution (High, 1939; O'Brien & Wibmer, 1982; CABI/EPPO, 2000; EPPO, 2002)

Virginia: restricted distribution (CIE, 1964; CABI/EPPO, 2000; EPPO, 2002)

Uruguay: present, no further details (Ruffinelli & Carbonell, 1954; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Venezuela: present, no further details (CIE, 1964; CABI/EPPO, 2000; EPPO, 2002)
Oceania

Australia: widespread (Wilson & Wearne, 1962; CABI/EPPO, 2000; EPPO, 2002)

Australian Northern Territory: present, no further details (Wilson & Wearne, 1962; CABI/EPPO, 2000; EPPO, 2002)

New South Wales: widespread (Gurney, 1932; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Queensland: present, no further details (Wilson & Wearne, 1962; Cunningham, 1969; Broadley, 1975; Allsopp & Lloyd, 1982; CABI/EPPO, 2000; EPPO, 2002)

South Australia: present, no further details (Wilson & Wearne, 1962; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Tasmania: present, no further details (Miller, 1945, 1979; CABI/EPPO, 2000; EPPO, 2002)

Victoria: present, no further details (Prescott, 1934; Anon., 1970; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

Western Australia: present, no further details (Jenkins, 1944; Button & Hardy, 1971; Morrone, 1993; CABI/EPPO, 2000; EPPO, 2002)

New Caledonia: present, no further details (Cohic, 1956; CABI/EPPO, 2000; EPPO, 2002)

New Zealand: present, no further details (Muggeridge, 1933; Ferguson & Hartley, 1978; Morrone, 1993; May, 1994; CABI/EPPO, 2000; EPPO, 2002)

Norfolk Island: present, no further details (Hawkins, 1942; CABI/EPPO, 2000; EPPO, 2002)


[footnotes not shown]


BIOLOGY AND ECOLOGY


Most of these notes are taken from High (1939) who described the life history of the pest in southern USA. L. costirostris deposits its eggs on plants or in nearby soil. The crowns of host plants are the preferred locations when ample moisture is present. After depositing the eggs on the leaves and stems, the weevil attaches them to the leaves by a glutinous secretion exuded at the time of oviposition. In the soil, the weevil constructs a small pocket with its ovipositor before depositing the eggs. The eggs are usually deposited singly, although masses of two to eight or more eggs are sometimes found. Weevils depositing only unfertilized eggs often oviposit in masses.
On the Gulf of Mexico coast, USA, egg deposition usually starts in late September and continues until the end of April, but this period varies greatly according to temperature. If oviposition begins in early autumn, as a result of some adults emerging from aestivation by temperatures that are low enough to stimulate activity, egg deposition by these individuals will be completed before cold weather. However, if high temperatures postpone emergence from aestivation until late autumn, such adults will not deposit their complement of eggs until the following spring. Egg incubation period ranged from 15 to 33 days when the average daily temperature was 12.8-24.4°C. During the spring and autumn most of the eggs under observation hatched within 15-20 days after deposition.
Immediately after hatching, the larvae begin feeding on the buds of the host plants, or on the undersides of leaves close to the buds. Later in their development they feed on all the foliage parts, and on root crops, such as turnips and carrots, they often feed on the roots, causing severe damage. The larval period under laboratory conditions varies greatly according to temperature and moisture, averaging approximately 35 days. Larvae are found in the field from the last week of October until mid-May of the following year, occurring in greatest abundance during December, January and February.
After the last moult the larva enters the soil and forms an earthen cell, within which it undergoes a resting or prepupal stage. This may last only a day or two, but where soil moisture is high or when temperatures are low, the resting stage is prolonged. Larvae entering the soil in late autumn and winter are much slower in transforming to pupae than are those entering the soil in the spring. Mature larvae, if disturbed after constructing their cells, will immediately begin repairing the cells before transforming into pupae.
The pupae are found in the soil near the host plant, at depths ranging from 12 to 50 mm, depending on soil moisture and texture. The pupal period ranges from 13 to 41 days. Pupae are present in the field during a normal season from mid-November until the middle of June, the peak abundance occurring in January.
As the pupa transforms into the adult weevil, the wings change colour from creamy white to pale brown. After transformation, the adult remains in the pupal cell for a few days before emergence until its body is sufficiently hardened. At emergence, the adult is pale brown and conspicuously marked with a pale grey 'V'-shaped mark on the elytra. Both these features are obscured during the adult stage, since the characteristic dull greyish-brown colour soon becomes predominant. Emergence of adults usually begins in early December and terminates in mid-June, peak emergence occurring between December and May.
The newly-emerged weevils are voracious feeders and seek food immediately after emergence. In the summer months the adults usually become inactive, except for occasional periods of slight feeding activity. They usually feed on the foliage of their host plants, but under some circumstances they feed on the roots of vegetables. Feeding occurs principally during the night, although weevils have been observed feeding during the day when heavy foliage gives them protection against direct sunlight. During the day, weevils hide under leaves, clods of earth, or other objects close to the soil surface.
As temperatures rise early in the summer, the adults aestivate and may be collected in large numbers under old straw and rubbish around the edges of gardens and fields. Adults may also shelter under loose bark, or in finely-pulverized soil during the early part of summer, but this may be only a temporary retreat, since in late summer it is difficult to locate weevils in the soil. After temperatures fall at the end of the summer, the adults leave their hiding places and resume their feeding activities on various host plants. The adults occur in the field throughout the year, but are present in greatest abundance from the end of December until mid-April.
L. costirostris may be disseminated by natural and artificial means. In the southern States of the USA it has spread northward, eastward and westward at a rate of approximately 50 miles a year. Dissemination has been more rapid in open, cultivated sections than in the wooded sections, suggesting that heavy forest growth may retard the normal rate of dispersal. Numerous field observations suggest that flight may be the principal means of dispersal. Inspections during the summer in some fields that were heavily infested the previous spring failed to reveal the presence of the weevil, whereas in fields that were free of infestation during the spring, large populations of aestivating weevils were found. Adults crawl rather slowly and it is doubtful whether they disperse significantly in this manner.
Under laboratory conditions in Mississippi, the time required for a complete life cycle during the spring and early autumn ranged from 48 to 111 days. In the Gulf of Mexico coast area, USA, there is only one generation annually (High, 1939).
In New Zealand, larvae are active during the autumn, winter and spring. The adults begin to emerge during September to October and aestivate in the hotter months of summer and early autumn (Muggeridge, 1933).
In Victoria, Australia, adults oviposit from mid-March to mid-September, depositing 1 to 30 eggs per day, representing some 300 to 1500 eggs per adult in a season. Reproduction is parthenogenetic, since there are no males. Larvae usually hatch in 11-24 days and by mid-winter, plants can be infested with 5-12 larvae. The pupal stage lasts for 14-16 days (Prescott, 1934).
In Uruguay, oviposition and larval development occurs throughout the winter from April to November, and the summer is passed in the adult stage (Parker et al., 1950).
In the Taipei area of Taiwan, adults appeared in November and only females were found. Larval populations reached a peak in late December. Feeding took place mainly at night, and pupation occurred in the soil. Adults emerged in February; they fed for 1-2 weeks, migrated from vegetable fields during March-April, and aestivated until November. In the laboratory at 16-22°C, the larval stage lasted 4-6 weeks and the pupal stage 4-5 weeks (Hsu and Chiang, 1983).
According to Morrone (1993), eggs are laid singly and lightly stuck to stems, low foliage and crowns of host plants, or pushed into the soil. Larvae feed on the aerial parts of plants, whereas mature larvae burrow into the soil by day and feed at night. Larvae moult to pupate in cells beneath the soil surface. The duration of the stages varies with temperature: eggs 14-17 days; larva 30-46 days; pupa 10-16 days; and adult 12 months.
In Japan, L. costirostris hibernates in a non-diapause state during the winter and can survive in northern areas despite low temperatures. Studies determining cold tolerance of larvae and adults indicated that the supercooling points of the adults and final-instar larvae were -9.0 and -6.7°C, respectively. All adults and half of the larvae tested could not tolerate -10°C for 3 hours. As air temperatures decreased, adults and larvae moved beneath the bottom leaves in contact with the ground surface or around the stalks where field temperatures were higher than in the aerial parts (Tsumuki et al., 1993).

NATURAL ENEMIES

Notes on natural enemies

L. costirostris is preyed upon by several species of spiders and ants, although the predatory fire ants described here are also pests. The fire ant, Solenopsis xyloni, has been found to destroy large numbers of larvae, pupae and adults in insectary cages, as well as adult weevils that had been made inactive by poison or natural causes in the field. The ants have also been observed to attack eggs and newly-hatched larvae repeatedly in the field. On several occasions, S. pergandei and the thief ant, S. molesta, have been observed attacking larvae. Along the Gulf of Mexico coast, USA, these ants are abundant and assist in destroying large numbers of juvenile weevils during the autumn, winter and spring (High, 1939).


Several parasitoids have been isolated from L. costirostris larvae in Argentina and Uruguay, although their importance in the field is unknown. These include: Tersilochus parkeri [Stethantyx parkeri], an unnamed Triaspis species, Epiplagiops littoralis, Stethantyx argentinensis, an unidentified species of Myiophasia (Pseudoclista). Euoestrophasia aperta, and unnamed species of the genera Microctonus and Clistromorpha (Hyalomyodes) have also been isolated from adults in Argentina. Parker et al. (1950) recorded parasitism of L. costirostris by representatives of Tersilochus, Epiplagiops and an unidentified nematode from Argentina. The Tersilochus and Epiplagiops parasitoids together achieved parasitism rates ranging from 6 to 74% in Argentina during 1942-1945. Clancy (1969) also recorded larval parasitism rates of 4.8-9.2% in California by an unidentified species of Tersilochus.
The entomophagous fungi Zoophthora phytonomi (as Entomophthora sphaerosperma) and Beauveria globulifera were isolated from larvae in Argentina and Uruguay.
According to Wilson and Wearne (1962), an unidentified nematode attacked larvae and adults in the field in Brisbane, New South Wales, Australia. Nematodes belonging to the genus Diplogaster, were found within the dead bodies of adults by High (1939).
Field observations in the southern USA have demonstrated that guineafowls, chickens and other domestic poultry are of considerable local value in reducing the damage caused by L. costirostris. Some growers have used flocks of young guineafowls and chickens to clean up heavy pest infestations after the crops were so badly damaged that they had no marketable value. In some fields this reduced the weevil population to such an extent as to prevent severe infestations to crops planted the following season (High, 1939).
High (1939) reported sparrows, meadowlarks, mockingbirds and blackbirds repeatedly feeding on larvae, although no stomach examinations of these birds were made to authenticate these observations.

List of natural enemies


Parasitoids:

- Epiplagiops littoralis, attacking: larvae, in Argentina, Uruguay

- Euoestrophasia aperta, attacking: adults, in Argentina

- Myiophasia atra, attacking: larvae, in Argentina

- Stethantyx argentinensis, attacking: larvae, in Argentina

- Stethantyx parkeri, attacking: larvae, in Argentina, Uruguay


Pathogens:

- Beauveria globulifera, attacking: larvae, in Argentina, Uruguay



- Zoophthora phytonomi, attacking: larvae, in Argentina, Uruguay


ECONOMIC IMPACT


The primary injury by larvae is caused when they devour the buds of the host plants, stunting them, followed by foliage attack, with the exception of the main stems and larger veins. The loss of the leaves not only interferes with the normal functioning of the plants, but affects the market value of the crops, for example, in southern USA, the leaves of turnips have a greater market value than the roots. Even when the infestation is not severe enough to cause noticeable injury, the mere presence of the larvae lowers the market value of the crops (High, 1939).
The monetary loss caused by L. costirostris in the USA is difficult to estimate as not only commercial vegetable growers are affected, but also a large number of home gardeners. The weevil attacks field crops, but also frequently causes serious injury to crops grown in cold-frames, heated beds and seedbeds. The degree of weevil infestation and crop loss varies considerably from season to season and also on different crops (High, 1939).
High (1939) reported serious economic damage to vegetable crops in the ten USA States where the pest was established in 1930. Growers in Mississippi, USA, reported losses as high as 90%, with common losses ranging from 40 to 70%. During 1933, the estimated crop losses for tomatoes alone ranged from 5 to 70% of the total crop value. Entire plantings of turnips, carrots, cabbages, mustard and spinach can be destroyed or seriously injured during the early stages of growth by this weevil. Practically all vegetable growers in the region adjacent to the Gulf of Mexico, USA, have suffered crop losses due to L. costirostris damage (High, 1939).
L. costirostris is intermittently injurious in all states of Australia, but especially Queensland and New South Wales (Wilson and Wearne, 1962).


PHYTOSANITARY RISK


As L. costirostris reproduces parthenogenetically, a single individual is capable of establishing the species in any new area where climatic conditions are favourable (Lovell, 1932). During the height of the breeding season, the weevil can be transported easily from place to place through the shipment of green vegetables and root crops, either on the plants or secluded in shipping containers. Repeated observations have demonstrated that the pest may be accidentally transported by persons or vehicles (High, 1939), although L. costirostris is not included on any quarantine lists for any country.


SYMPTOMS


L. costrirostris stunts the growth of numerous vegetables by destroying the tender young crown leaves; carrots and turnips being preferred by the larvae (Muggeridge, 1933).
The holes made in the leaves of host plants by small larvae are round, whereas those made by larger larvae are irregularly shaped. When the larvae are numerous, representing approximately 10-25 per plant, the leaves are skeletonized, with only the main stems and larger veins remaining. When attacking onions, the larvae often tunnel inside the leaves (High, 1939).
Adults often cause extensive damage by feeding on the leaves of small tomato and potato plants and by cutting off the stems of plants at ground level, the injury resembling that of cutworms (Noctuidae). Weevils also attack buds and leaves later in development (High, 1939).
Descriptors: Leaves: external feeding. Roots: external feeding. Stems: external feeding. Whole plant: cut at stem base; external feeding.

MORPHOLOGY

Eggs
The egg is spherical or slightly elliptical and is approximately 1 mm or less in diameter. When first deposited it is creamy-white, which rapidly changes to a dark yellow until just prior to hatching, when it is black. The surface of the chorion is smooth and shiny (High, 1939).

Larvae


The larva has been described by Barrett (1930), Peterson (1951), described and figured by High (1939), Anderson (1987), Lee and Morimoto (1988) and May (1994). It was separated from soil-inhabiting larvae of other species in keys produced by May (1966, 1977).
Body slender, slightly curved, eusterna bilobed, dorsally strongly convex, flattened ventrally, greenish cream to bright green according to host; setae minute, flattened apically; maximum body length 11.5-15.0 mm, maximum width 3.5 mm. Head: free, broader than long, width 1.4-1.75 mm, pale to medium brown with clearly defined, darker maculate pattern; two pairs of ocelli present, each convex, distinct; endocarina absent; mandibles with median and submedian acute supplementary teeth. Pronotum paler in colour than head, dusky with darker maculae. Abdominal segments 1-7 with three dorsal folds; segments 2-6 with proleg-like ampullae; anal proleg present. Anus ventral, trilobate. There are four instars.

Pupae


The pupa was described and figured by High (1939) and May (1994). The pupa is white, the legs and wing pads are yellow and the abdominal segments are pale green. The rostrum is broad and bent back along the body to the first pair of legs. Body length is 7.5-10.0 mm, over half as wide as long; pronotal width 2.5 mm. Cuticle glabrous; setae short and dark, spiniform on small tubercles. Urogomphi are pale, darker basally with three small, associated setae.

Adults


Integument brown, except for antennae and tarsi which are reddish brown, head and rostrum without scales, densely pubescent; pronotum with indistinct, median stripe of pale grey-brown scales; elytra bearing regular rows of slender, erect, white and brown scales, two to three times longer than diameter of adjacent scales, which are rounded, overlapping, decumbent, grey, golden-brown and dark brown, and forming more or less distinct, 'V'-shaped fascia behind the middle. Antennae with funicular segments 1 and 2 elongate, the former slightly longer than the latter. Rostrum tricarinate, gradually broadening from base to apex, bearing lateral, downward pointing flange anterior to the eyes on which the scrobe terminates. Ventral carina of scrobe toothed. Frons not narrower than rostrum at base in dorsal view. Eyes oval, clearly separated, acuminate ventrally. Pronotal disc slightly convex, with well-developed, broad, shallow, arc-shaped impression anteriorly; ocular lobes distinct, with straight flanks. Prothorax with two yellowish lines. Scutellum small, densely pubescent. Elytra with shoulders distinct, more or less angular, apex of interstice 5 with conical tubercle; elytral interstices slightly convex; setae short. Metatibiae with one spur. Protarsi with segments 1, 2 and 3 of increasing width, all tarsi with third segment bilobed, metatarsi elongate; tarsal claws paired and free. Venter setose. Metasternum shorter than transverse diameter of mesocoxae. Aedeagus robust in lateral view. Body length 6.4-10.0 mm.
A key to the Listroderes species of Argentina, including L. costirostris, was provided by Hustache (1926). Morrone (1993) provided a key to the adults of the L. costirostris species group and figured the apical lobe of the male genitalia.


SIMILARITIES TO OTHER SPECIES


According to High (1939), the banded vegetable weevil, L. apicalis, is often found attacking vegetables in association with the vegetable weevil. However, both sexes of L. apicalis are usually present, whereas L. costirostris is mostly parthenogenetic. L. costirostris is slightly larger and darker than L. apicalis and possesses the characteristic pale-coloured 'V'-shaped figure on the elytra, whereas L. apicalis is a paler shade of brown with a pale-coloured bar extending across the rear of the elytra. The posterior elytral tubercles are more prominent in L. apicalis.


DETECTION AND INSPECTION METHODS


Examine the buds and undersides of young crown leaves of susceptible plants for small, greenish-cream to bright greenish larvae up to 15.0 mm in length and dorsally convex in shape. Also look for the damage caused by larvae which consists of either small round holes or large irregular holes eaten from the underside, with the main veins left intact.
Examine the undersides of leaves at night for greyish-brown adult weevils, up to 10 mm in length, except during the summer months. During the summer months search for adults under straw and other plant debris lying on the ground at the edges of gardens or fields, and also under loose bark of trees. In addition, look for signs of adult damage to young tomato and potato plants, which are cut off at ground level. Such damage resembles that caused by cutworms (Lepidoptera: Noctuidae).

CONTROL

Cultural Control

Since large numbers of immature weevils are present in the soil at various depths during the winter and early spring, frequent ploughing followed by harrowing will destroy many of them directly, or indirectly by exposing them to predators (High, 1939). Early tomato and potato crops that have to be replanted in autumn owing to the damage caused by the weevil can be protected by cleaning up the ground thoroughly (Gurney, 1932).


In Australia, Prescott (1934) suggested harrowing during July to October to break up the pupal cells; rotation of crops (peas, beans, cucurbits or cereals are not attacked) and keeping fields clear of wild hosts, particularly capeweed (Arctotheca calendula), chickweed (Stellaria media) and marsh mallow (Malva rotundifolia).
During the summer when the weevils are aestivating along fence rows and ditch banks or beneath dead grass and rubbish in, or close to infested fields, large numbers of weevils can be destroyed by thoroughly cleaning up such places by burning or by otherwise disposing of all available shelter. This practice should also include the removal of loose bark from nearby trees, a common shelter for adult weevils (High, 1939).

Biological Control


Four species of parasitoid that have so far been introduced from Argentina and Uruguay to California, USA, are the tachinid, Epiplagiops littoralis (Blanchard, 1943), the ichneumonids, Stethantyx parkeri and S. argentinensis (Blanchard, 1945) and a braconid of the genus Triaspis (Annand, 1945). According to Parker et al. (1950) these parasitoids are not known to have become established. Difficulty was experienced in breaking the aestivating diapause of the Stethantyx adults to adapt them to the seasonal rhythm of the northern hemisphere, and E. littoralis required an alternative host during the summer.
Wilson and Wearne (1962) describe the introduction of parasitoids from Uruguay and Argentina during 1957-1958 to New South Wales, Australia. The species imported were the tachinid Stomatomyia littoralis, S. argentinensis and two strains of S. parkeri. Of 646 adults of S. littoralis released in November, no recoveries were made. This tachinid does not aestivate, and the likelihood of its becoming established is reduced by the need for an alternative summer host. Several attempts at releases of natural enemies have occurred in Australia, none of them have resulted in the establishment of these parasites (Clausen, 1978).
High (1939) stated that it appeared feasible to use domesticated poultry, such as guineafowls and chickens, as a control measure.

Chemical Control


Treatment should begin when 5% or more of small, newly set plants (within 3 weeks after transplanting) are killed or injured. Vegetable weevil larvae in tobacco plant beds can be controlled using parathion, disulfoton or acephate insecticides. In the field, adult weevils can be controlled using parathion-methyl or parathion dust.
Field trials undertaken in Florida during 1973-1974 to study the chemical control of L. costirostris on tobacco showed that dust or spray treatments with acephate and methomyl were satisfactory substitutes for DDT, which was banned for use on tobacco in 1969 (Tappan, 1974).
Foliar applications of carbofuran or quinalphos in New Zealand provided greater control of larval infestations on cabbage during August-September 1973 in adverse weather conditions than the standard treatment with parathion. Two applications of azinphos-ethyl, carbofuran, methamidophos, parathion or quinalphos gave similar and acceptable reductions in larval numbers. After a single application of DDT, an intolerable number of live larvae remained on the plants. A marked improvement in plant growth could be seen after the application of insecticides (Ferguson and Hartley, 1978).


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