weevil

The milfoil weevil (Euhrychiopsis lecontei) is a small, herbivorous aquatic beetle, belonging to the family Curculionidae, that is native to North America. It is a watermilfoil (Myriophyllum spp.) specialist, meaning that it feeds and develops only on plants in this genus. Its original host before the introduction of Eurasian watermilfoil (Myriophyllum spicatum) was the native northern watermilfoil (M. sibiricum). The weevil completes all life stages fully submersed, feeding and developing on milfoil, and the larvae are stem miners. These characteristics make it unique, as specialist herbivores are very rare among aquatic insects. These characteristics are why the milfoil weevil has shown the most promise as a potential biocontrol agent for Eurasian watermilfoil and why it has been the subject of much research. Several other herbivores, such as the lepdidopterans Acentria ephemerella and Parapoynx spp., and the midge Cricotopus myriophylli, can damage and control milfoil and may be important in some locations, but have not shown as much potential as the milfoil weevil.

Milfoil weevils affect Eurasian watermilfoil mainly by boring through the stem and consuming the cortex. This results in reduced plant buoyancy and accumulated carbohydrate stores, which may ultimately decrease overwinter survival and competitive ability. Because milfoil declines have often occurred over winter and in early summer or have persisted over several years, it is likely that longer term effects, such as reduced overwinter survival or reduced competitive abilities are important to sustained control of Eurasian watermilfoil. Our observations suggest that plant community response, i.e., the ability of other species to occupy space left by damaged milfoil or the stress imposed by competition with other plants, is also important to successful biological control.

Research on the operational use of the milfoil weevil is continuing and incomplete. Although we know milfoil weevils can control milfoil, we are currently unable to predict when and in what lakes this will occur. More work is needed to determine what controls milfoil weevil and other milfoil herbivore populations and what role plant competition plays in successful control.


Where they're found

The milfoil weevil appears widespread across the northern U.S. and southern Canada and occurs in many lakes in Minnesota and Wisconsin. It is found in lakes that have Eurasian watermilfoil and lakes with the native northern watermilfoil that have not yet been infested with Eurasian watermilfoil. Recent surveys in Wisconsin indicate that the milfoil weevil likely occurs in most lakes that have northern or Eurasian watermilfoil.

In the Midwest, milfoil declines have been associated with the milfoil weevil in Fish Lake, Wisconsin and a less well-documented decline in McCullom Lake, Illinois. In Minnesota, documented declines have been associated with the weevil in sites at four lakes (Auburn, Cenaiko, Otter and Smith’s Bay of Lake Minnetonka); in some cases the declines persisted and in other cases  suppression only lasted a few years.

In several lakes of the Minneapolis Chain of Lakes, milfoil weevil densities are very low and milfoil is not controlled. Experimental evidence suggests that a density of 100-200 weevils per square meter of bottom may be required to affect a decline. At many sites in Minnesota, weevil densities have failed to reach such levels, although 25-75 weevils per square meter (or less) may be adequate to cause declines if they persist through the summer. We are currently investigating factors that influence weevil density; fish predation appears to be an important factor at sites with low weevil populations. Mechanical harvesting of milfoil over large areas can reduce weevil abundance, but harvesting was not occurring at many sites we have studied.

Life cycle

Like many insects, the milfoil weevil develops through 4 stages: egg, larva, pupa and adult.  The complete life cycle of the milfoil weevil takes 21-30 days at 20-25 degrees C, with survival ranging from 20-70%; there is a positive linear relationship between development rate and temperature.

Egg  weevil

The milfoil weevil lays its eggs on milfoil plants, usually on apical meristems within a few feet of the water surface. Female weevils lay single eggs that they stick to the plant, and generally lay one to several eggs on a meristem. However, up to 29 eggs have been laid on a single apical meristem when no other options were available. Eggs are a yellow-cream color, elliptical, and approximately 0.5 mm long. The egg stage lasts about 3-6 days at typical mid-summer temperatures of 20-25 degrees C. Hatching success has been reported to be between 65-100%.

Larva weevil

After hatching, first instar larvae feed on meristem tissues for 3 to 5 days, while older larvae spend the majority of their time inside the stem where they feed on cortical and vascular tissues. Larvae are generally found in the top 1 m of the plant. Stems that have been hollowed-out by weevil larvae appear darkened, are less buoyant due to gas leakage, and are weaker than undamaged stems. Occasionally, larvae will bore out of the stem, travel up or down the stem in a spiral path, and bore back into the stem. This behavior is most common when a larva reaches the end of an internode. Late instar larvae reach a length of about 4.5 mm. Development time through the larval stage ranges from 7-10 days at 20-25 degrees C and larvae damage about 15cm of stem from their feeding. Survival through the larval stage has been recorded at 78-90%.

Pupa weevil

The milfoil weevil pupates (metamorphoses) inside of milfoil stems in a pupal chamber. Generally, they are found further down the stem than larvae (> 0.5 m), possibly because a larger diameter stem is preferred for pupation. However, successful pupal development has been recorded on stems as narrow as 1 mm. Typical development times through the pupal stage are 8-12 days at mid-summer water temperatures (20-25 degrees C). Survival through the pupal stage has been recorded at 69-80%.

Adult weevil

Adult milfoil weevils are usually located on the upper 1 m of milfoil plants. They are weak swimmers, and will usually remain on a plant even after it has been disturbed. Adult weevils primarily eat milfoil leaves, but will also consume stem tissues. This is the only stage of the weevil that can exit the water. However, they appear to only leave the water in fall when they migrate to shore and over winter terrestrially in leaf litter near the shoreline. It remains unclear if the weevil swims or flies on this short migration to shore, but many raft in on milfoil fragments. They possess wings, but weevils are rarely observed in flight, which appears restricted to the terrestrial phase. Adult milfoil weevils are approximately 2-3 mm in length and have lived as long as 162 days in captivity. Females lay an average of 1 to 5 eggs per day (3/day over lifetime), and total egg production by captive females ranged up to 562 eggs.

Water temperature, host plant, and host plant quality have been shown to affect development time and success. This life cycle period allows for three or more generations per summer. Adults leave the shore in spring and return to the water after ice out, between mid-April and mid-May in Minnesota; they do fly at this time and may disperse to other lakes in the spring.

Host plant choice

The milfoil weevil is highly specific to milfoil plants (Myriophyllum spp.) Because the weevil is endemic to North America, and Eurasian watermilfoil probably was not established in North America until the 1940s, it is evident that the original host was northern watermilfoil on which it is commonly found. However, with the introduction and spread of Eurasian watermilfoil across much of North America, the milfoil weevil was exposed to a novel plant that is closely-related to its natural host. When both plants are present, weevil densities are usually higher on Eurasian watermilfoil and host choice tests indicate a strong preference for Eurasian watermilfoil and performance (survival, time to complete development, and size) is generally better on Eurasian than northern watermilfoil.  It is clear the milfoil weevil has undergone a host range expansion to include and prefer Eurasian watermilfoil.

Biocontrol potential

As indicated above, Eurasian watermilfoil declines have been associated with the occurrence of the milfoil weevil and other herbivorous insects at numerous locations. Declines of Eurasian watermilfoil associated with these insects have been documented in several lakes in Vermont, New York and Massachusetts, Fish Lake Wisconsin, a lake in Washington state, several sites in Minnesota, and several sites in Illinois.

Smaller scale experimental manipulations, in field enclosures and outdoor tanks have shown substantial and significant effects of these herbivores on Eurasian watermilfoil. These observations all indicate that the herbivores can control Eurasian watermilfoil, however, research has yet to determine when, where and how these herbivores will be effective or where their densities will be sufficient to control the plant.

Researchers have predicted that densities of 100-200 weevils per square meter (or approximately 0.5-2 weevils per stem) may be needed to effectively control Eurasian watermilfoil. More recent observations suggest that control may occur at lower densities (<0.5/m2), however, control clearly does not consistently occur at these lower densities unless they persist throughout the summer. Several issues appear to be important but need more investigation. First, factors that limit weevil density and abundance must be determined and ameliorated. Second, a positive plant community response after milfoil suppression appears important. Lastly, the mechanisms by which long term control occurs is not clear.

Keep in mind that successful weed biocontrol aims to reduce pest populations to tolerable levels and that low densities of the pest are needed to maintain control agent populations. Successful biocontrol of Eurasian watermilfoil would reduce its abundance, eliminate surface matting and promote healthy native plant communities but would not eliminate the plant (or its native relative) from the lake.



Do you have milfoil weevils in your lake?

Many people have asked how to get milfoil weevils (Euhrychiopsis lecontei) or if they have milfoil weevils in their lake. EnviroScience of Stow, Ohio marketed milfoil weevils and a control process (Milfoil Solution) for over 15 years but ceased providing this service in 2015. We are not aware of any other similar services although some local and state governments have supported weevil stocking and may continue.  There are legitimate concerns about moving weevils and milfoil within and between states. Currently our best advice is to determine if you have weevils in your lake and take steps to conserve or promote weevil populations and native plant communities. If, for example, sunfish are limiting your herbivore populations, stocked herbivores will likely be eaten before they can develop sufficient populations. This is what you are looking for:

weevil

General search approach:

To search for weevils you need to be able to locate watermilfoil plants in your lake; northern and Eurasian are the two most common species in the upper Midwest:

ewm
     

Once you have located a milfoil bed the simplest things to look for are signs of weevil damage - weevil damage is more quickly spotted than adult weevils. Searching is easiest by snorkeling, but can also be accomplished by examining stems while wading or least efficiently from a boat. Milfoil weevils live in the top three feet, more typically top foot, of healthy milfoil plants - avoid flowering plants and look for plants that are submersed, several inches to a foot below the surface. Search in 3-8 feet of water. Examine the top foot of plants under the water or grab clumps of milfoil a foot or two beneath the surface for examination.

Weevil-damaged plants will have darkened and hollowed stems contrasting, as seen below, with healthy golden or green stems.

ewm

Occasional small holes should also be present in the darkened stem. The larvae will mine from the meristem (growing plant tip) downward; depending on the extent of damage, only meristems may be missing, a few inches may be mined and darkened, or several feet of a stem may be damaged. Weevils almost always cause darkening of the entire stem circumference - damage by other insects such as midges (Chironomidae) usually only results in darkening of one side of the stem.

ewmThis plant shows a missing meristem (dense green clump of leaves at the tip of the plant) and extensive mining damage (mining hollows out the stem resulting in darkening and lack of structural integrity).

Note that the mining is not continuous. Also note the darkened circular areas lower on the stem. These are pupation sites - finding these sites are fairly certain signs of milfoil weevil activity.

Pupae and pupal holes will typically be located 2-3 feet down the milfoil stem and generally are several inches or more below the closest stem mining damage. The pupa pictured is metamorphosing from a larva to adult.

Larvae are small and non-descript without visible legs - occasionally they venture out of the stem, but mining damage, pupae or adults are better to search for in the field.

The meristems should also be examined for eggs. Eggs are about 0.5mm in diameter, yellowish and most adhered to or embedded within the meristem.

If you spot damage or eggs in an area you can then search for adults. Adults can be positively identified to species and either adults or very advanced pupae are required for positive identification. As indicated in the above photos, adults are typically near the tops of submersed milfoil and are small. All weevils have a characteristic snout, which in the milfoil weevil looks like an elephant trunk. Milfoil weevils spend the summer submersed; if you find weevils on the shore or on emergent plants or emergent milfoil flow spikes they are almost certainly not milfoil weevils. 

If you find weevils in your lake and wish to be certain they are the milfoil weevil, you should preserve several adults in alcohol (70% or stronger rubbing alcohol will suffice) and include a paper label with your name, lake name and county, nearest city and date. The specimens can then later be verified by an expert.

The milfoil weevil undergoes a complete life cycle, egg, larva, pupa and adult, underwater during the summer. In Minnesota, weevils will lay eggs from late May to early September; in October adults migrate to shore where they spend the winter in leaf litter at dry sites right along shore. Populations seem to go in pulses - late June through early August seems to be the best time to locate adult weevils. If you spot damage but no adults, return to the same site in two weeks and check again; repeat throughout the summer if necessary.
For more information, please contact Dr. Ray Newman.