TY - JOUR
AU - Edde, Peter, A
AB - Abstract The cigarette beetle, Lasioderma serricorne(F.) (Coleoptera: Anobiidae), is an important pest in the food and tobacco industry in many regions of the world. Despite a great deal of research, control of this pest still relies on the use of phosphine fumigation, which is becoming less effective as the insect develops resistance to this compound. In addition, series of other nonchemical methods used to control the insect have given mixed and irregular results. This review summarizes and discusses information on important aspects of the biology and ecology of the cigarette beetle, and its control. The topics covered include a taxonomic discussion of the cigarette beetle, which includes a discussion of other anobiid species of economic importance. The mating behavior of the insect and conditions favorable for pest development were described. The review also includes a discussion of the life stages of the insect, its feeding habit, and economic damage. Important aspects of its chemical ecology and a discussion on the association between this species and its microorganisms, and major natural enemies, were presented. A summary of its flight behavior, including the factors governing flight initiation and temporal and seasonal flight activity were reviewed. Finally, the control methods currently used in the management of the insect were described. The review also identifies potential areas of further research on L. serricorne and gives an analysis of the control methods worthy of further investigation in the search for practical and sustainable methods for the management of this pest. cigarette beetle, tobacco beetle, stored-product insect The cigarette beetle, Lasioderma serricorne (F.) (Coleoptera: Anobiidae), is a pest of considerable economic importance in several countries of the world. The species is the most destructive of the insect pests of unprocessed and processed tobacco and extremely large variety of other materials of both plant and animal origin (Runner 1919, Howe 1957, Hagstrum et al. 2013). Important work on L. serricorne biology has been carried out by many entomologists. Prominent studies include those by Jones (1913), Runner (1919), Howe (1957), and Sivik et al. (1957), who shed light on the biology of the insect under different environmental conditions. The effect of different temperatures and humidities on the duration of various life stages of the insect was examined by Powell (1931) and Rayner (1951). Powell (1931) also published interesting information on the effect of food quality on the development of the insect under constant conditions of temperatures and humidities. Aspects of the nutritional ecology of the insect and associated symbionts were intensely studied by Fraenkel and Blewett (1943), Pant and Fraenkel (1950), and later by Dowd (1989). Moreover, the most important original contribution on the embryology of the species was made by Canzanelli (1935). Finally, an elucidation of the chemical ecology of the insect, especially of its female-produced pheromone, was made by Chuman et al. (1979, 1981, 1985). The most comprehensive publication on the biology of L. serricorne, which has yet been written, at the time of this writing, was by Ashworth (1993). The work by Ashworth (1993), however, omitted information on the management of this economically important insect. Ryan (1995) and da Silva et al. (2018) reviewed the biology and management of L. serricorne. Hence, the current review presents a summary of relevant literature on the general biology and ecology of L. serricorne, reviews published information on its control measures, and identifies areas where knowledge is inadequate or lacking with the aim of encouraging future research on L. serricorne. General Classification Taxonomy and General Notations L. serricorne is placed in the tribe Lasiodermini, further in Xyletininae, a subfamily of the beetle family Anobiidae. In addition to L. serricorne, other economically well-known members of the family include Xestobium rufovillosum (De Geer) (the deathwatch beetle), Stegobium paniceum (L.) (the drugstore beetle), and Anobium punctatum De Geer (the furniture beetle). The nomenclature of this taxon, however, appears to be in flux. For example, Lawrence and Newton (1995) pointed out the priority of the coleopteran family Ptinidae over Anobiidae, making the former a senior synonym. The Ptinidae and Anobiidae were probably grouped because of their similarity in size and coloring, and their similar habit. Ptinidae consists of over 2,200 species in about 230 genera worldwide (Lawrence 1991), of which about 464 anobiid species in about 63 genera occurs in North America and Mexico. A key to the genera of Anobiidae found in North America and Mexico was published by Philips (2002) and White (1971). Boving (1927) published detailed descriptions and classifications of the family Anobiidae according to larval morphology. Parkin (1933) reported on the external morphology of eight species of larvae belonging to the family Anobiidae and constructed keys for their identification. The adult of the anobiids are cylindrical to oval or globular, 1.1−9.0 mm long (usually less than 5 mm; Borror et al. 1992). Their heads are strongly deflexed under a shield-shaped pronotum. The shape of the antenna is filiform. The antennae consist of 11 segments, of which the last three segments are longer, and sometimes broader than the other segments. The antennae are inserted widely apart, with the distance between them more than the length of the first antennal segment. While in Ptinidae, the antennal insertions are narrowly separated, the distance between them is less than the length of the first antennal segment (Fall 1905). Their prosternum is reduced and often excavated to receive the head. Their tarsal formula is 5-5-5. The adult of anobiid beetles are short lived and rarely feed. Anobiid larvae are wood borers, but some species have become adapted to feed on various stored-products. The genus Lasioderma was established by Stephens in 1832. The author placed the single species testaceum Duftschmid in the genus. Currently, L. testaceum is considered as a synonym of L. serricorne. Lasioderma includes five species in North America and over 50 species are recognized worldwide (Arango and Young 2012). Two of the generic synonyms for Lasioderma are Hypora (Mulsant & Rey 1864) and Pseudochina (Jacquelin du Val 1860). There is no general agreement as to the original home of L. serricorne. The insect was originally described in 1792 from a specimen of “dry plants” collected from the Americas (Fabricius 1792). However, Alfieri (1932) mentioned having found the insect in a sealed amphora of the tomb of the Egyptian pharaoh Tutankhamun, suggesting its presence in Egypt about 1,500 B.C.E. The author observed that the beetle has scarcely altered morphologically in the 3,500 yr that have elapsed. The first record of the association of L. serricorne with cured tobacco in Europe was made in Paris in 1848 (Runner 1919, Canzanelli, 1935) and in the United States around 1885–1886 (Fullaway 1914, Runner 1919). At present, L. serricorne occurs throughout the tropical and subtropical regions, and in heated buildings in temperate regions. The distribution of the species among geographical regions usually occurs through being transported, at all life stages, in infested commodities (Blanc et al. 2006). The most suitable climate for its development is in the tropical and subtropical zone. The only limitation to its distribution appears to be exerted by low temperature and to a lesser extent by low humidity (Rayner 1951). Synonyms L. serricorne was first described as Ptinus serricornis by Fabricius in 1792. Thereafter, it was referred to, by various authors, as Lasioderma breve (Wollaston, 1861), Lasioderma castaneum (Melsheimer, 1845), Lasioderma flavescens (Dahlbom, 1837), Lasioderma fuscum (Rey, 1892), Lasioderma rufescens (Sturm, 1826), Lasioderma testaceum (Duftschmid, 1825), Lasioderma torquatum (Chevrolat, 1859), Pseudochina serricornis (Fabricius, 1792), Ptinus serricornis (Fabricius, 1792), Ptilinus testaceus (Duftschmid, 1825), Xyletinus brevis (Wollaston, 1861), and Xyletinus torquatum (Chevrolat, 1859). It was Le Conte, who in 1865 described the beetle as L. serricorne. In 1892, Rey listed the insect as Pseudochina fuscum, but many authorities continue to regard L. serricorne as the valid name of the insect. In addition to “tobacco beetle” or “cigarette beetle”, other common names recorded for the beetle include “tobacco bug”, “tobacco borer”, “tobacco weevil”, and “cheroot beetle” (Maxwell-Lefroy 1906, Runner 1919, Rayner 1951). Another common name is the “tow bug” derived from the feeding habit of the insect on upholstery fillings such as tow, flax, and hemp (Hartnack 1939). Runner (1919) noted that L. serricorne was erroneously referred to in older literatures as “tobacco flea” or “tobacco flea beetle” from confusion of the species with Epitrix parvula (F.) (Coleoptera: Chrysomelidae), which attacks growing tobacco. Life Stages Egg Jones (1913), LeCato and Flaherty (1974), Kucerová and Stejskal (2010), and Gautam et al. (2014) have given detailed descriptions of the external morphology of L. serricorne eggs. Fletcher (1977a) described methods for the collection of large numbers of L. serricorne eggs. The eggs are opaque pearl white when freshly laid but becoming yellowish shortly before hatching. They are oval or oblong in shape, and varying from 0.29 to 0.50 mm long, and 0.18–0.25 mm in diameter. The average length and diameter of the egg is 0.35 and 0.20 mm, respectively (Jones 1913, Canzanelli 1935). Each egg weighs about 8.0–9.0 μg. Rayner (1951) reported that oviposition in L. serricorne occur within 12 h after mating, Canzanelli (1935) mentions 2 d, while Kurup and Parkhe (1962) reported that the pre-oviposition period ranged from 1 to 5 d. The eggs are laid in the early evening and during the night directly on the dried food material (Sivik et al. 1957). During summer months the oviposition and post oviposition period is about 7 d each (Sivik et al. 1957). The total number of eggs laid by a single female in a lifetime varied ranging from 10 eggs to 100 eggs or more (Runner 1919, Sivik et al. 1957, Kurup and Parkhe 1962, Yu 2008). The average ranged from 40 to 76. Punzo (1975) noted that the greatest rate of oviposition in stored-product insects occur at temperatures approaching the upper limit for reproduction and decreases sharply at higher temperatures and more slowly at lower temperatures. The number of eggs laid by the beetle tends to vary among different food substrates. For example, under identical laboratory conditions (28–32°C and 72.5–80.5% relative humidity [RH]), the females were found to lay 14, 19, 25, 37, and 40 eggs on rice, cowpea, groundnut, maize, and wheat, respectively (Allotey and Unanaowo 1993). At 26.7°C and 70% RH, most of the eggs are laid within the first 9–11 d of the life of the insect, with most being deposited during the first 3–4 d after mating (Jones 1913, Rayner 1951, Yu 2008). On flue-cured tobacco, usual oviposition rate is 2–5 eggs per day for each mated female and the greatest number of eggs that can be laid during a 24-h period is about 25–30. Crowding may cause female beetles to deposit most of their mature eggs quickly (Reed et al. 1935), but overcrowding will reduce the overall number of eggs laid per female (Kurup and Parkhe 1962). At 20°C the embryonal development of L. serricorne eggs takes about 10 d (Canzanelli 1935). About 90–95% of the eggs hatched in 6–8 d (range, 4–11 d) at 27–30°C and 65–70% RH. Similarly, Tenhet and Bare (1951) reported the average incubation period to be 7 d during summer months. Rayner (1951) found that egg viability is about 100% between 21.0 and 32.2°C at 44% or 75% RH. Larva During hatching, the larva chews its way through the chorion at the posterior margin of the papillated region, so that it lifts like a cap enabling the larva to emerge through the opening (Rayner 1951). Anobiid larvae, including those in the genera Lasioderma, have been described in detail (Boving 1927, Lawrence 1991). The newly hatched L. serricorne larva is semitransparent, but gradually assumes a whitish or yellowish color. If the larva feeds on colored food substances, such as tobacco, the food may be seen in the digestive tract through the transparent larval skin. The newly emerge larva may eat the whole egg shell and other unhatched eggs if no other food is available (Rayner 1951, Howe 1957). On emergence, the larva is slender and fully stretched, very active and is capable of crawling considerable distances in search of food (Rayner 1951, Kurup and Parkhe 1962). Kurup and Parkhe (1962) found that unfed larvae that were 4–5 d old can travel up to 195 cm. Thus, infestation may easily spread from infested to uninfested materials. The young larvae are negatively phototrophic, that is, would retreat from light (Runner 1919, Rayner 1951). They will enter small holes in search of food and can penetrate packaged commodities. According to Runner (1919), newly hatched larvae are capable of surviving for 5–10 d without food, but other workers (e.g., Kurup and Parkhe 1962) suggested a maximum of 2 d. To molt, a median dorsal split first occurs in the head capsule of the old integument, allowing the capsule to slip forward over the larval head and to push down with the rest of the exuviae to the posterior region of the body where the whole of the skin is cast off (Runner 1919, Canzanelli 1935, Rayner 1951). During the second molt, the larvae assume the characteristic scarabaeiform shape. Older larvae are less active but are still capable of considerable wandering. They tend to penetrate deeply into commodities packed loosely but staying in the peripheral in tightly packed containers. When fully grown, larvae become turgid, immobile, and stops feeding. It then forms a thin cell or cocoon made of food waste material cemented together by secretion produced by the midgut. The cocoons are made wherever the larvae can find a spot that would give firm foundation for the cocoon. For example, in milled food substrates, the insect may build the cocoon loosely inside the rearing media. The cocoons are typically ovoid but varies in the shape, measuring about 4.5 mm long and 3 mm wide (Runner 1919). L. serricorne typically undergoes four growth periods or instars before pupation. However, up to six larval instars have been reported depending on the temperature in which the insect is reared, with the frequency of larval molting increasing as temperatures decrease (Rayner 1951, Niiho 1984). For example, Howe (1957) obtained four larval instars at 30°C and 70% RH, while Niiho (1984) found that the number of molt increased from four instars between 22.5 and 30.0°C to five instars at 20°C. Larvae are quiescent and stop feeding about 24 h before each molt. Larvae increase in mass including length through the larval instars. The average body length of the first, second, third, and fourth larval stage to be 0.6, 1.8, 2.8, and 5.0 mm, respectively. The average head capsule width of the larva is about 0.11, 0.18, 0.30, and 0.49 mm for the first through fourth larval instar, respectively. Descriptions of the matured L. serricorne larvae are provided by Rayner (1951) and Howe (1957). Prepupae and Pupae The matured larvae do not pupate at once after the cocoon had been constructed (Sivik et al. 1957). They lie quiescent in the cocoon while it undergoes structural changes preparatory to pupation, a stage that Runner (1919) referred to as the “prepupal”. At 30°C, the matured larva cast its skin after 2–4 d in the cocoon (Howe 1957). Ordinarily the insect pupates in the cocoon; however, the presence of a cocoon is dispensable as some larvae can transform into pupae and later adults without having made a cocoon (Canzanelli 1935). In this case, it may be difficult to discern the onset of the prepupal stage. Even with cocoon formation, Rayner (1951) found that the prepupal stage is not distinguishable from the matured larva by any other distinct external characteristics. Before transforming to pupa, the matured larva or prepupa undergoes a slight contraction in length losing it scarabaeiform shape to become perfectly straightened, with the body becoming more deeply wrinkled. The duration of the prepupal stage is about 2 d but may extend up to 5 d (Jones 1913). Pupae are uniformly white with a slight greenish tinge when they are newly formed, but gradually assume a reddish-brown color, which darkens with age (Runner 1919, Canzanelli 1935). They are inactive, with movement limited to wriggling of the abdominal segments. The apex of the elytra of pupae elytra often reaches the fourth abdominal segment. Their metathoracic legs lie beneath and do not reach the tips of the inner wings. The head is drawn back into the thorax and bent under the pronotum. The average length of the pupa is 3.5 mm (range, 2.5–3.75 mm) and average width 1.7 mm (range, 1.1–2.0 mm; Canzanelli 1935). Female pupae are often slightly bigger than those of males reared on the same media (Rayner 1951). Sexual dimorphism is present at the tip of the abdomen, which can be seen after the molted skin is removed. The terminal segment of the female pupa has a pair of lateral lobes, which are very distinct and are absent in the male (Rayner (1951). Genitalia are globular and not projecting in the males but are three-segmented and divergent in the females (Halstead 1963). In Italy, the pupal stage lasts 11–12 d between April and August, and from September to October the pupal development varies from 20 to 24 d (Canzanelli 1935). However, working in the United States, Sivik et al. (1957) quoted 3–8 d (average 5 d) during summer months, and Tenhet and Bare (1951) reported 7 d in the summer, but 10–14 d in cooler weather. Adult The adult stays in the pupal cells from 4 to 6 d before they issue, sexually matured, and with fully developed coloration. To emerge, the pupal skin splits on the dorsal side of the head to the abdomen and the adult emerges aided by a wriggling motion of the entire body. Both sexes of the adult are similar in external appearance. A general description of the adult has been given above under the discussion on classification and descriptions of the anobiids. Moreover, detailed description of the adult has also been given by Fabricius (1792), Fall (1905), Runner (1919), and Howe (1957). In general, the body is elongate-oval, moderately convex, pubescence moderate, and sub-recumbent. There are 11 segments on the antennae. To these may be added the fact that the second and third antennal segments in the species are smaller than the first, the third distinctly triangular, and the fourth to tenth about as wide as long, the eleventh oval, slightly longer than the tenth. The general body color is reddish-brown. But a black body color form of the species may be found (Coffelt and Vick 1973). Unlike the reddish-brown color of the normal adult, the exoskeleton of the adult mutant is black in homozygotes and mahogany brown in heterozygotes. The mutation is also visible during the latter part of each larval and the pupal stadium. However, unlike in the black mutant of Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae) (Onyearu and Graham 1967), the development and reproductive fitness of the black-bodied beetles are parallel to that of the reddish-brown color of the normal adult (Coffelt and Vick 1973). Adult females weigh more than males, with this difference in weight first occurring during the third larval instar (Lefkovitch and Currie 1963). The average size of the adult female is 2.6 mm long and 1.4 mm wide, while the male beetle measures on an average 2.1 mm in length and 1.2 mm in breadth (Bhalodia and Chari 1976). However, because beetle size does overlap, it may not be a reliable criterion of’ sex determination. The sex ratio of the beetle is about 1:1 throughout the active flight season, but there are periods during the summer when either male or female may be found in greater numbers than the other (Sivik et al. 1957). Adult longevity is 2–7 wk (Runner 1919, Rayner 1951, Howe 1957, Allotey and Unanaowo 1993, Yu 2008). However, several factors may affect the longevity of the adult beetles. For example, Yu (2008) found that unmated males and females lived for 29–35 d and mated adults lived for 17–23 d, but this was shorter that the 60–90 d reported by Powell (1931) for unmated females. The feeding status of the adult may also affect its longevity. Shinoda and Fujisaki (2001) found beetles provided with both pollen and water at the same time lived significantly longer than unfed beetles. Larval food may also influence adult longevity. Adults whose larvae were reared on rice, cowpea, groundnut, maize, and wheat at 28–32°C and 72.5–80.5% RH lived from 27 to 33 d, 23 to 27 d, 24 to 32 d, 26 to 31 d, and 24 to 28 d, respectively (Allotey and Unanaowo 1993). Sex Differences Several methods have been used to morphologically distinguish between sexes. For example, Reed et al. (1935) and Rayner (1951) cleared the adult by soaking them for about 24 h in a 10% solution of potassium hydroxide. The cleared insects were then placed in a small glass dish holding glycerin to reveal the genitalia when examined under a binocular microscope. In the method of Papadopoulou and Buchelos (2002a), adult beetles were placed in 70% alcohol or benzene for about 5 min. The solvent treatment of the beetles resulted in urosternite transparency, making the V-shaped apodeme visible by simple observation of the abdomen under the binocular microscope. Runner (1919) separated the sexes by abdominal dissection of adult beetles to examine the reproductive structures. While these methods are accurate for determining the sexes of the adult beetle, they entail the destruction of the insect, thus inadequate to use where live insects are needed to gather important biological or bionomical data. As suggested for R. dominica (F.) (Coleoptera: Bostrichidae) live or freshly killed adult can be reliably sexed by gently squeezing the last two to three ventral abdominal segments until the tips of the genitalia appear to reveal the transparent female oviduct or sclerotized male clasper (Edde 2012). To reduce mortality caused by squeezing, the insect can be sexed at the immature stage using the sexual dimorphism described above for the pupal stage. Three major factors, however, may prevent sexing L. serricorne pupae in this way. Firstly, the last larval integument may adhere to the terminal segment of the pupa and may be impossible to remove the integument without injury to the pupa. However, pupal sexing is rarely hampered using this method. Secondly, it may prove difficult to sex old pupae because with age the lateral lobes tend to disappear, and the terminal segments would have started to invaginate (Rayner 1951). Thirdly, without some protection against loss of water, pupae may dry up easily. Mating Behavior and Condition for Development Mating Behavior Although L. serricorne can mate as soon as individuals emerge from the pupal cell (Rayner 1951), beetles usually begin to mate on the second or third days after adult emergence (Runner 1919, Levinson and Levinson 1987, Papadopoulou 2006a). Mating may take place both inside and outside commodity packages (Sivik et al. 1957). Courtship and mating in L. serricorne is aided by both pheromonal and tactile stimuli provided by the female, which are perceived by respective sensilla of the antennae and palpi of the male (Coffelt and Burkholder 1972, 1973; Levinson and Levinson 1987). However, males often try to mate with other males, especially near the pheromone source (Chuman et al. 1982). A brief description of the female-produced pheromone is given later in the review. Coffelt and Burkholder (1973) found that both the physiological (ovarian) and behavioral (mating receptivity) maturity in the species coincides at 4–6 d after adult emergence. This age also coincides with the period when female sex pheromone production and mating activity is at maximum (Coffelt and Burkholder 1973). Rayner (1951), Tobin and Smith (1971), Levinson and Levinson (1987), and Papadopoulou (2006) have published reports on the mating behavior of L. serricorne. Contrary to observations in many other stored-product coleopteran species (e.g., Benelli et al. 2017), during its copulation the male cigarette beetle does not stay on top of the female, but in an opposed way, such that only the abdominal ends of both male and female touch one another. Both sexes of the beetle are capable of mating more than once in their life cycle (Bhalodia and Chari 1976, Levinson and Levinson 1987). But the second mating usually takes place about 6 h after the first mating (Bhalodia and Chari 1976). The greatest number of multiple mating observed for a female was 3, and up to 10 (mean, 7.2) for males (Coffelt 1975). Males showed no loss of fertility at advanced ages and after multiple mating (Coffelt 1975). Unmated females do not lay eggs (Runner 1919, Dick 1937, Rayner 1951), but some mated females may lay infertile eggs (Runner 1919, Fletcher and Long 1976). Female L. serricorne usually mate after each oviposition (Rayner 1951, Tobin and Smith 1971). Repeated mating has been observed in other major stored-product beetle species as well. For instance, in the confused flour beetle Tribolium confusum Duval (Coleoptera: Tenebrionidae), a second mating, even in females that are still laying fertile eggs, has a marked stimulatory effect (Dick 1937). Similarly, the adult female of yellow mealworm, Tenebrio molitor L. (Tenebrionidae) may delay or reduce oviposition or be incapable of achieving maximal fecundity until they have gained the material and/or genetic benefits of mating multiple times (Worden and Parker 2001). However, multiple mating has no effect in the case of L. serricorne. For example, Coffelt (1975) found that the number of offspring produced by females that have mated 1, 2, or 3 times were similar. Coffelt (1975) also noted the occurrence of heterozygous offspring among progeny of multiple mated females, which suggest that, rather than any effect on fecundity, multiple mating in the species serves as a mechanism to increase diversity. Coffelt (1975) opined that overall fecundity is a function of female weight (i.e., larval nutrition) and the age at which the adult begins to oviposit. L. serricorne females not only mate several times, but will also mate with multiple males, a phenomenon referred to as polyandry. Female insects that mate with more than one male may derive reproductive fitness, genetic benefits, or both (Ridley 1988, Worden and Parker 2001, Gay et al. 2009, Okada et al. 2017, Rafter et al. 2017). For example, Rafter et al. (2017) found most females T. castaneum or R. dominica captured in flight traps had mated with more than one male and those females that had mated several times had the potential to produce offspring of multiple genotypes, showing that polyandry is beneficial in these species. Likewise, Worden and Parker (2001) found that T. molitor females that mated with four different males had higher oviposition rates than females that mated an identical number of times, but with the same male. However, polyandry carries fitness costs but not benefits to females in L. serricorne (Okada et al. 2017). Female L. serricorne that were courted by two males had reduced lifespans compared with females that mated only once, showing a direct fitness cost to females due to contact with two males (Okada et al. 2017). The direct cost to the female tend to be greater when the second mate was an aggressive male, due to increased male sexual harassment (Gay et al. 2009, Okada et al. 2017), and from the damage caused by male genitalia, which bear spines that puncture the female’s reproductive tract, and/or toxic elements in the ejaculate (Gay et al. 2009). Conditions for Development Numerous studies on the lifecycle of the insect under controlled environmental conditions have been published (Powell 1931, Canzanelli 1935, Rayner 1951, Howe 1957, Manoto et al. 1976, Niiho 1984, Mahroof and Phillips 2008, Yu 2008, Tamang 2015, Suneethamma 2016). The developmental rates of both sexes are similar (Tamang 2015). Data on the relative duration of the different life stages of the insect are summarized in Table 1. The amount of damage likely to be caused by a pest depends on how rapidly the population increases (Howe 1965). The most favorable conditions for the rapid development of L. serricorne larvae include the following: 1) suitable food sources in compact or concentrated form, 2) high and uniform temperature, 3) high humidity, 4) protection from bright light, and 5) protection from rapid evaporation (Runner 1919). Table 1. Shape, measurement, and days needed for the development of the various stages of L. serricorne Stadia Shape/Type Average measurement (mm)a,b Average length of the stadia (days)c Head Length Width 28°Cd 29°Ce 32°Cf 35°Ce Egg Ovoid-elliptical — 0.39 0.18 8.1 5.5 4 5 1st instar Campodeiform 0.12 1.21 0.20 4.7 — — — 2nd instar Scarabaeiform 0.17 1.95 0.57 4.5 — — — 3rd instar Scarabaeiform 0.37 2.47 0.73 4.7 — — — 4th instar Scarabaeiform 0.49 4.46 1.28 11.8 — — — Larva total — — — 25.6 39 16 34 Prepupa — — — — 3.1 — 2.6 Pupa Exarate 2.75 1.32 4.6 4 9 3.6 Egg to adult — — — — 38.8 56 29 49 Stadia Shape/Type Average measurement (mm)a,b Average length of the stadia (days)c Head Length Width 28°Cd 29°Ce 32°Cf 35°Ce Egg Ovoid-elliptical — 0.39 0.18 8.1 5.5 4 5 1st instar Campodeiform 0.12 1.21 0.20 4.7 — — — 2nd instar Scarabaeiform 0.17 1.95 0.57 4.5 — — — 3rd instar Scarabaeiform 0.37 2.47 0.73 4.7 — — — 4th instar Scarabaeiform 0.49 4.46 1.28 11.8 — — — Larva total — — — 25.6 39 16 34 Prepupa — — — — 3.1 — 2.6 Pupa Exarate 2.75 1.32 4.6 4 9 3.6 Egg to adult — — — — 38.8 56 29 49 aTamang (2015). bFigure obtained from insect reared on turmeric powder at temperature varying from 24.20 to 32°C (average, 29.8°C) and humidity from 50 to 73% RH (average, 68.8% RH). cFigures obtained under constant conditions of humidity 65% RH (Yu, 2008), or 75% RH (Rayner 1951, Powell 1931) turmeric powder (Tamang 2015), poultry feed (broiler grower feed; Yu, 2008), yeast cake (Powell 1931), or tobacco (Rayner 1951) used as food. dYu (2008). eRayner (1951). fPowell (1931). Open in new tab Table 1. Shape, measurement, and days needed for the development of the various stages of L. serricorne Stadia Shape/Type Average measurement (mm)a,b Average length of the stadia (days)c Head Length Width 28°Cd 29°Ce 32°Cf 35°Ce Egg Ovoid-elliptical — 0.39 0.18 8.1 5.5 4 5 1st instar Campodeiform 0.12 1.21 0.20 4.7 — — — 2nd instar Scarabaeiform 0.17 1.95 0.57 4.5 — — — 3rd instar Scarabaeiform 0.37 2.47 0.73 4.7 — — — 4th instar Scarabaeiform 0.49 4.46 1.28 11.8 — — — Larva total — — — 25.6 39 16 34 Prepupa — — — — 3.1 — 2.6 Pupa Exarate 2.75 1.32 4.6 4 9 3.6 Egg to adult — — — — 38.8 56 29 49 Stadia Shape/Type Average measurement (mm)a,b Average length of the stadia (days)c Head Length Width 28°Cd 29°Ce 32°Cf 35°Ce Egg Ovoid-elliptical — 0.39 0.18 8.1 5.5 4 5 1st instar Campodeiform 0.12 1.21 0.20 4.7 — — — 2nd instar Scarabaeiform 0.17 1.95 0.57 4.5 — — — 3rd instar Scarabaeiform 0.37 2.47 0.73 4.7 — — — 4th instar Scarabaeiform 0.49 4.46 1.28 11.8 — — — Larva total — — — 25.6 39 16 34 Prepupa — — — — 3.1 — 2.6 Pupa Exarate 2.75 1.32 4.6 4 9 3.6 Egg to adult — — — — 38.8 56 29 49 aTamang (2015). bFigure obtained from insect reared on turmeric powder at temperature varying from 24.20 to 32°C (average, 29.8°C) and humidity from 50 to 73% RH (average, 68.8% RH). cFigures obtained under constant conditions of humidity 65% RH (Yu, 2008), or 75% RH (Rayner 1951, Powell 1931) turmeric powder (Tamang 2015), poultry feed (broiler grower feed; Yu, 2008), yeast cake (Powell 1931), or tobacco (Rayner 1951) used as food. dYu (2008). eRayner (1951). fPowell (1931). Open in new tab The best RH for L. serricorne development ranged from 65 to 75%. Published reports on the minimum humidity levels required for the development of the species varied among authors. Canzanelli (1935) and Howe (1965) reported the lower limit to be 30%, but Powell (1931) states 45%, while Rayner (1951) quoted 23% for beetles reared on milled corn. High humidity does not prevent the growth of L. serricorne unless the food is destroyed by mold growth (Powell 1931, Rayner 1951, Howe 1965). The effects of humidity on the duration of L. serricorne life cycle are greatest in the egg and larval stage than on the pupal stage because while both the incubation period of the egg and larval development are lengthened in low humidities, pupal development is affected very little by humidity (Powell 1931). Insects are sensitive to fluctuations in temperature because they do not have the ability to regulate their body temperature. At 75% RH, the best temperature range for rapid development of the cigarette beetle is 29–35°C (Rayner 1951). The species appears to be one of the most heat tolerant species among storage insect pests. Larvae can survive for about a week at 40°C provided it is returned to best temperature range for development of the insect, and this ability increases with larval age (Howe 1957). The upper temperature limit for development of newly hatched larvae is between 35 and 37.5°C (Rayner 1951, Howe 1957). There are conflicting reports in literature concerning the life stage of L. serricorne that is most tolerant to elevated temperatures. Runner (1919) reported that fourth-instar larvae and pupae (cocoons) are most tolerant to heat, with the pupal stages being the most tolerant, probably because their cells act as barrier to heat penetration and/or water loss. Other workers (Powell 1931, Subramanyam 2009, Yu et al. 2011), however, found the egg stage to be the most tolerant. Yu et al. (2011) even suggested that egg hatchability can serve as a good criterion to decide the susceptibility of L. serricorne to elevated temperatures between 46 and 54°C. At 46°C the LT99 for egg hatchability and egg-to-adult emergence was 10.08 and 9.97 h, respectively (Yu et al. 2011). At 50°C, it decreased to 3.17 and 2.77 h, and 0.65 and 0.63 h at 54°C. Interestingly, Adler (2002a) found the eggs to be the most tolerant life stage at 50°C, but this was reversed at 45°C (Adler 2002b). Acclimation to sublethal elevated temperatures can significantly improve heat tolerance in the species, thus increasing their chance of survival when subjected to temperatures that were hitherto lethal for the insect. For example, Meng et al. (2018) found the LT50 of adults, pupae, larvae, and eggs of L. serricorne that were acclimated to 42°C for 20 h were 2.2, 2.2, 3.4, and 4.8 times higher than that of insects without acclimation, respectively, when exposed to lethal elevated temperature of 50°C. Understanding the temperature/time/ mortality relationships of insects is important to know least temperature and time to which the insects should be exposed for effective control (Subramanyam 2009). The lower temperature limits needed for L. serricorne larval development range from 15 to 19°C (Runner 1919, Howe 1957, Niiho 1984). However, other authors (e.g., Rayner 1951, Papadopoulou 2004) found the lower critical temperatures for larval or pupa development to lie between 13 and 15°C. In general, the susceptibility of L. serricorne to low temperatures is dependent on the developmental stage of the insect. The fourth-instar larval and adults are the most resistant to low temperature (Crumb and Chamberlin 1934; Childs et al. 1968, 1970; Imai and Harada 2006). Fletcher and Long (1976) found that beetles can lay eggs at temperatures below 15°C, but the embryonic processes only occur at the best temperature range for development of the insect (Canzanelli 1935). At temperatures below 15°C, L. serricorne eggs do not hatch and embryos do not develop even when the eggs are removed to optimum temperature range (Crumb and Chamberlin 1934). A good proportion of L. serricorne eggs may hatch at 18.3°C (Fletcher and Long 1976), but at this temperature, the development of the insect will continue, but at a slower rate (Crumb and Chamberlin 1934). The effects of larval food on the growth and development of L. serricorne populations have been investigated by several authors (Powell 1931, Howe 1957, Bharathi et al. 2001, Mahroof and Phillips 2008). At 32°C and 75% RH, Powell (1931) found that 18–20 more days are needed for the insect to develop from the egg stage to adult emergence on tobacco than in yeast (Table 2). At 25−27°C, a stable population of L. serricorne reared on wheat feed can increase by 60% every week (Howe 1957). In addition, assuming a stable age distribution, the population of the insect will increase 7.123 times every 3 wk for insects reared on finely sieved wheat feed or broad English bran at 30°C and 60% RH (Lefkovitch and Currie 1967). The effects of food type on the development of L. serricorne reared under similar environmental conditions of temperatures and humidity are summarized in Table 2. In addition to food type, the life history parameters of the species are affected by the nutritional levels of the larval diet, being lower on less nutritive foods. As shown in Table 3, important life parameters (e.g., net reproductive rate, intrinsic rate of increase) are highest on the tobacco type (flue-cured) than on the less nutritive type (cigar wrapper). Cigar wrappers are usually made from burley or reconstituted leaf sheet (RL). A description of the different tobacco types, including flue-cured has been provided by Edde (2018). RL sheet is a paper-like material made from recycled tobacco fines, tobacco stems, and other tobacco materials. Table 2. Development of the immature stages, and total developmental period, of L. serricorne reared on different food sources Temp. (°C) RH (%) Developmental period (d) Rearing media Reference Egg Larva Prepupa Pupa Egg-to-adult 25 67 6.15 17.9 2.8 3.8 30.7 Cornmeal-yeast mixture (18:1) Manoto et al. (1976) 25 70 9.8 65.1 — 6.4 81.4 Bread crumbs Niiho (1984) 26 60–90 6.7 28.90 — 12.6 54.7 Fennel Suneethamma (2016) 26 60–90 6.3 27.83 — 12.3 53.1 Coriander Suneethamma (2016) 27.5 70 7.0 51.3 — 4.6 62.9 Bread crumbs Niiho (1984) 28 65 8.1 25.6 — 4.6 38.3 Poultry feed Yu (2008) 28 60 4.8 38.0 — 4.6 46.0 Wheat flour Mahroof and Phillips (2008) 28 60 5.2 53.0 — 8.2 66.4 Leaf tobacco Mahroof and Phillips (2008) 28 60 6.6 63.0 — 7.6 77.2 Paprika Mahroof and Phillips (2008) 28 60 6.6 75.3 — 5.7 87.7 Cayenne pepper Mahroof and Phillips (2008) 28 60 5.0 73.0 — 18.3 95.3 Chili Mahroof and Phillips (2008) 28 60 4.8 92.0 — 10.4 107.4 Cigar tobacco Mahroof and Phillips (2008) 29 75 5.5 38.0 3.1 4.4 57.0 Tobacco leaves Rayner (1951) 29 75 5.5 27.0 2.9 4.2 45.0 Corn meal Rayner (1951) 29.8 68.8 9.0 27.0 5.0 7.0 48.0 Turmeric powder Tamang (2015) 30 70 6 20 — 4 30 Wheat feed Howe (1957) 32 75 — — — — 28 Magic yeast Powell (1931) 32 75 — — — — 38–48 Tobacco Powell (1931) Temp. (°C) RH (%) Developmental period (d) Rearing media Reference Egg Larva Prepupa Pupa Egg-to-adult 25 67 6.15 17.9 2.8 3.8 30.7 Cornmeal-yeast mixture (18:1) Manoto et al. (1976) 25 70 9.8 65.1 — 6.4 81.4 Bread crumbs Niiho (1984) 26 60–90 6.7 28.90 — 12.6 54.7 Fennel Suneethamma (2016) 26 60–90 6.3 27.83 — 12.3 53.1 Coriander Suneethamma (2016) 27.5 70 7.0 51.3 — 4.6 62.9 Bread crumbs Niiho (1984) 28 65 8.1 25.6 — 4.6 38.3 Poultry feed Yu (2008) 28 60 4.8 38.0 — 4.6 46.0 Wheat flour Mahroof and Phillips (2008) 28 60 5.2 53.0 — 8.2 66.4 Leaf tobacco Mahroof and Phillips (2008) 28 60 6.6 63.0 — 7.6 77.2 Paprika Mahroof and Phillips (2008) 28 60 6.6 75.3 — 5.7 87.7 Cayenne pepper Mahroof and Phillips (2008) 28 60 5.0 73.0 — 18.3 95.3 Chili Mahroof and Phillips (2008) 28 60 4.8 92.0 — 10.4 107.4 Cigar tobacco Mahroof and Phillips (2008) 29 75 5.5 38.0 3.1 4.4 57.0 Tobacco leaves Rayner (1951) 29 75 5.5 27.0 2.9 4.2 45.0 Corn meal Rayner (1951) 29.8 68.8 9.0 27.0 5.0 7.0 48.0 Turmeric powder Tamang (2015) 30 70 6 20 — 4 30 Wheat feed Howe (1957) 32 75 — — — — 28 Magic yeast Powell (1931) 32 75 — — — — 38–48 Tobacco Powell (1931) Open in new tab Table 2. Development of the immature stages, and total developmental period, of L. serricorne reared on different food sources Temp. (°C) RH (%) Developmental period (d) Rearing media Reference Egg Larva Prepupa Pupa Egg-to-adult 25 67 6.15 17.9 2.8 3.8 30.7 Cornmeal-yeast mixture (18:1) Manoto et al. (1976) 25 70 9.8 65.1 — 6.4 81.4 Bread crumbs Niiho (1984) 26 60–90 6.7 28.90 — 12.6 54.7 Fennel Suneethamma (2016) 26 60–90 6.3 27.83 — 12.3 53.1 Coriander Suneethamma (2016) 27.5 70 7.0 51.3 — 4.6 62.9 Bread crumbs Niiho (1984) 28 65 8.1 25.6 — 4.6 38.3 Poultry feed Yu (2008) 28 60 4.8 38.0 — 4.6 46.0 Wheat flour Mahroof and Phillips (2008) 28 60 5.2 53.0 — 8.2 66.4 Leaf tobacco Mahroof and Phillips (2008) 28 60 6.6 63.0 — 7.6 77.2 Paprika Mahroof and Phillips (2008) 28 60 6.6 75.3 — 5.7 87.7 Cayenne pepper Mahroof and Phillips (2008) 28 60 5.0 73.0 — 18.3 95.3 Chili Mahroof and Phillips (2008) 28 60 4.8 92.0 — 10.4 107.4 Cigar tobacco Mahroof and Phillips (2008) 29 75 5.5 38.0 3.1 4.4 57.0 Tobacco leaves Rayner (1951) 29 75 5.5 27.0 2.9 4.2 45.0 Corn meal Rayner (1951) 29.8 68.8 9.0 27.0 5.0 7.0 48.0 Turmeric powder Tamang (2015) 30 70 6 20 — 4 30 Wheat feed Howe (1957) 32 75 — — — — 28 Magic yeast Powell (1931) 32 75 — — — — 38–48 Tobacco Powell (1931) Temp. (°C) RH (%) Developmental period (d) Rearing media Reference Egg Larva Prepupa Pupa Egg-to-adult 25 67 6.15 17.9 2.8 3.8 30.7 Cornmeal-yeast mixture (18:1) Manoto et al. (1976) 25 70 9.8 65.1 — 6.4 81.4 Bread crumbs Niiho (1984) 26 60–90 6.7 28.90 — 12.6 54.7 Fennel Suneethamma (2016) 26 60–90 6.3 27.83 — 12.3 53.1 Coriander Suneethamma (2016) 27.5 70 7.0 51.3 — 4.6 62.9 Bread crumbs Niiho (1984) 28 65 8.1 25.6 — 4.6 38.3 Poultry feed Yu (2008) 28 60 4.8 38.0 — 4.6 46.0 Wheat flour Mahroof and Phillips (2008) 28 60 5.2 53.0 — 8.2 66.4 Leaf tobacco Mahroof and Phillips (2008) 28 60 6.6 63.0 — 7.6 77.2 Paprika Mahroof and Phillips (2008) 28 60 6.6 75.3 — 5.7 87.7 Cayenne pepper Mahroof and Phillips (2008) 28 60 5.0 73.0 — 18.3 95.3 Chili Mahroof and Phillips (2008) 28 60 4.8 92.0 — 10.4 107.4 Cigar tobacco Mahroof and Phillips (2008) 29 75 5.5 38.0 3.1 4.4 57.0 Tobacco leaves Rayner (1951) 29 75 5.5 27.0 2.9 4.2 45.0 Corn meal Rayner (1951) 29.8 68.8 9.0 27.0 5.0 7.0 48.0 Turmeric powder Tamang (2015) 30 70 6 20 — 4 30 Wheat feed Howe (1957) 32 75 — — — — 28 Magic yeast Powell (1931) 32 75 — — — — 38–48 Tobacco Powell (1931) Open in new tab Table 3. Life parameters of L. serricorne on three tobacco typesa,b Life parameterc Net reproductive rate Intrinsic rate of increase (Females/female/d) Mean generation time (d) Finite rate of increase (females/female/d) Doubling time Annual rate of increase Tobacco type  Flue cured 6.5017 0.045 42.1 1.0454 15.63 1.073 × 107  Burley 5.3963 0.0306 55.49 1.0308 22.82 6.537 × 104  Cigar wrapper 3.6791 0.0286 45.55 1.028 24.41 3.169 × 104 Life parameterc Net reproductive rate Intrinsic rate of increase (Females/female/d) Mean generation time (d) Finite rate of increase (females/female/d) Doubling time Annual rate of increase Tobacco type  Flue cured 6.5017 0.045 42.1 1.0454 15.63 1.073 × 107  Burley 5.3963 0.0306 55.49 1.0308 22.82 6.537 × 104  Cigar wrapper 3.6791 0.0286 45.55 1.028 24.41 3.169 × 104 aModified from Bharathi et al. (2001). bNutritional quality of the tobacco types are in the decreasing order: Flue-cured > Burley > Cigar wrapper. Nutritional quality was determined based on the chemical composition of nicotine, reducing sugar, and chloride contents of the leaves. cThe rearing was carried out at 27 ± 2°C and 65 ± 5% RH. Open in new tab Table 3. Life parameters of L. serricorne on three tobacco typesa,b Life parameterc Net reproductive rate Intrinsic rate of increase (Females/female/d) Mean generation time (d) Finite rate of increase (females/female/d) Doubling time Annual rate of increase Tobacco type  Flue cured 6.5017 0.045 42.1 1.0454 15.63 1.073 × 107  Burley 5.3963 0.0306 55.49 1.0308 22.82 6.537 × 104  Cigar wrapper 3.6791 0.0286 45.55 1.028 24.41 3.169 × 104 Life parameterc Net reproductive rate Intrinsic rate of increase (Females/female/d) Mean generation time (d) Finite rate of increase (females/female/d) Doubling time Annual rate of increase Tobacco type  Flue cured 6.5017 0.045 42.1 1.0454 15.63 1.073 × 107  Burley 5.3963 0.0306 55.49 1.0308 22.82 6.537 × 104  Cigar wrapper 3.6791 0.0286 45.55 1.028 24.41 3.169 × 104 aModified from Bharathi et al. (2001). bNutritional quality of the tobacco types are in the decreasing order: Flue-cured > Burley > Cigar wrapper. Nutritional quality was determined based on the chemical composition of nicotine, reducing sugar, and chloride contents of the leaves. cThe rearing was carried out at 27 ± 2°C and 65 ± 5% RH. Open in new tab However, dietary factors other than nutritional contents, including moisture contents may exert significant effects on the rate of development of the species. Depending on the rearing diet, the best moisture contents for L. serricorne development are between 10 and 14% at temperatures of 29–35°C. The incubation period of L. serricorne eggs or the growth rate of larvae kept under similar environmental conditions of food, temperatures, and humidity tends to vary, resulting in overlapping of generations among populations of the species (Runner 1919, Rayner 1951). Much of the mortality in L. serricorne occurs in the early instar stages, particularly in the first instar (Bharathi et al. 2001). Feeding Habit and Economic Damage Food L. serricorne belong to a group of insects which infest “stored food products”, and feeds and lives in many kinds of dried vegetable products and upon a few dried animal substances. Its common names “tobacco beetle” and “cigarette beetle” are of a misnomer since the species has a wide range of hosts. L. serricorne may have the most varied taste in food of most storage insects just behind T. castaneum (Howe 1957, Hagstrum et al. 2013). Substrates that have been reported as breeding materials or food for L. serricorne have been compiled by different authors (e.g., Runner 1919, Howe 1957, Hagstrum et al. 2013). Howe (1957) listed 38 different plant materials, six types of animal matter, and an assortment of miscellaneous material such as cloth, paper, books, upholstery, and furniture stuffing that have been reported damaged by L. serricorne. Summarizing the commodities that have been found infested by this species, Hagstrum et al. (2013) listed 222 dried plant and animal products and cites studies on the ability of L. serricorne to breed on 49 of these commodities. The occurrence of the beetle in many of the reported substances may have been accidental, or the larva may not have been able to complete its developmental cycle (Runner 1919, Hori et al. 2011, Hagstrum et al. 2013). Therefore, more breeding experiments are required to help determine the true range of materials liable to infestation. Example of such studies is that by Mahroof and Phillips (2008). Nevertheless, the fact that L. serricorne is an important pest of tobacco could be mostly attributed to the absence of competition with other stored product species, which cannot develop in this commodity. A biomarker that distinguishes different hosts may also be useful to identify the food use pattern of the insect. Such biomarkers have been found reliable to predict host use pattern or larval food for insects that do not feed as adults (Mahroof 2013). Both manufactured tobacco products and the unprocessed tobacco leaves are infested. However, all tobaccos are not attacked indiscriminately or with equal intensity. L. serricorne preference for cured tobacco is about the following order: Flue-cured type, Turkish types, Cuban and Puerto Rico types, Connecticut Broadleaf and Havana types, Pennsylvania Seedleaf, Dark-fired types, and Burley. Flue-cured and Turkish/Oriental tobacco types have some of the highest sugar and lowest nicotine content while Dark-fired tobacco has some of the highest nicotine and little to no sugar at all. This ranking suggests that the cured tobacco insect prefer the sweeter grade of tobacco as these are low nicotine and high sugar content (Milne 1961). Nevertheless, higher nicotine levels and lower sugars may not alone retard growth and reproductive fitness of the insect because other nutritional factors are present, which may act alone or synergistically, to affect the toxicity of nicotine to the beetles (Yamamoto and Fraenkel 1960). For example, artificial diets having up to 10% nicotine were not lethal to the beetle when yeast was added (Yamamoto and Fraenkel 1960). Economic Importance Apart from tobacco, information is limited about the nature of damages caused by L. serricorne on its economically important food hosts. Adults of the beetle will cut holes to penetrate or escape from packaged commodities leaving a neat round hole, which classifies this species to the category of “true penetrators” of food packages (Highland 1983, Riudavets et al. 2007, Athanassiou et al. 2011). Adults rarely feed, but may feed on pollen readily, and drink water or aqueous solutions of saccharose or honey (Runner 1919, Shinoda and Fujisaki 2001). Larval feeding causes most of the damage observed in infested commodities. In tobacco leaves, larvae tunnel out long cylindrical galleries especially near the midrib, while in ground food, eggs are laid on the surface, and resulting larvae feed down into the flour in all directions. The injury caused by the insect is often severe, owing to its habit of spending its entire life stages inside its food, feeding on them and causing the following damages: 1) weight losses due to amount of raw materials actually consumed by the larvae; 2) reduced market value of manufactured products by production of excrement, cast-off-skins, dead bodies, gnawed particles, and other products; and 3) loss of goodwill as consumer may turn away after buying infested products (Reed and Vinzant 1942). However, the greatest dollar loss caused by L. serricorne occurs in leaf during storage. At 28°C and 70% RH, about 0.012–0.15 g of food (Turkish tobacco) is required to feed one newly hatched cigarette beetle larva to adulthood. The weight of frass resulting from the feeding activity averaged 0.10 ± 0.02 g per beetle. In the tobacco industry, the damage and economic loss that occurs with L serricorne infestations are estimated to be about 0.7–1% of the total warehoused tobacco commodity (USDA 1971, Farnham et al. 2007). The degree of infestation, however, depends upon the temperature, RH, leaf type, and other conditions in storage. Manufactured products are also damaged by larvae, hence the common name “cigarette beetle”. Like other packaged agricultural products, consumer complaints involving L. serricorne infestation in the finished product is a common occurrence in the tobacco industry, particularly in hot humid climates during summer months. The period of greatest danger of factory or retail infestation in the United States is typically from June to October (Edde, unpublished data), but also in other countries (e.g., Papadopoulou and Buchelos 2002b). The actual amount of product consumed usually is of far less importance than is the presence of frass, dust, dead larvae, and adults, which render the manufactured product unsalable. Parallel to some species of the beetle family Dermestidae and Carabidae, L. serricorne has been associated the enteric infection known as canthariasis, an important insectal disease in humans caused by the adult or their larvae (Sun et al. 2016). Rare cases of intestinal canthariasis in children attributed to L. serricorne have been reported in China (Sun et al. 2016) and Malaysia (Mokhtar et al. 2016). The symptoms include general discomfort, fever, irritation of the eyes, and gastrointestinal disorders (Mokhtar et al. 2016, Sun et al. 2016). Victims may also pass out larvae in their stool. Canthariasis is caused by accidentally feeding on food materials infested by immature stages of insects, which can result in the infection of gastrointestinal tract, urogenital system, nasal sinuses, ears, and faces of mammals. Moreover, just like other stored-product beetle species, the enteric bacteria of L. serricorne include species of the genus Enterococcus, which possess antibiotic resistance genes that can be spread by horizontal transfer to more serious human pathogens (Yezerski et al. 2005, Miller et al. 2014). Chemical Ecology Female-Produced Pheromone Female L. serricorne release a sex pheromone which elicits a very strong attraction in conspecific males and excites them to copulate. (4S,6S,7S)-4,6-Di-methyl-7-hydroxynonan-3-one (serricornin or 4S,6S,7S-serricornin) is the main component of the female-produced pheromone (Chuman et al. 1979, 1985). The pheromone is produced in a specific gland located in the second abdominal sternite of the female, ventrally attached to the narrow part of a V-shaped apodeme (Levinson et al. 1983, Levinson and Levinson 1987). The gland is connected to a duct which leads to an orifice beneath the genital pore. Other compounds found in the pheromone gland of female L. serricorne are presented in Table 4. These compounds are regarded as minor components, produced in lower quantities, and are less attractive to male beetles (Chuman et al. 1985). For example, the attractiveness of anhydroserricornin, the second most abundant compound in the pheromone, to the male is less than 1/103 of that SSS-serricornin (Mochizuki et al. 1984), and the compound showed no synergistic or inhibitory effect when combined with serricornin. However, a stereoisomer of the natural pheromone 4S,6S,7S-serricornin, 4S,6S,7R-serricornin displayed a strong inhibitory action against the pheromonal activity of SSS-serricornin (Chuman et al. 1981, Mochizuki et al. 1984), thus could prove helpful in integrated management of the species (Okada et al. 1992). The primary compound in most commercial pheromone lures for L. serricorne is serricornin. Still, there are published surveys in which anhydroserricornin was used in monitoring studies with satisfactory results (Buchelos and Levinson 1993; Papadopoulou and Buchelos 2002b,c). Table 4. Components of the sex pheromone of L. serricorne (Chuman et al., 1985) Common name Structural formula Serricornin (4S,6S,7S)-4,6-Di-methyl-7-hydroxynonan-3-one Anhydroserricornin 2,6-Diethyl-3,5-dimethyl-3,4-dihydro-2H-pyran NA 4,6-Dimethylnonan-3,7-dione NA 4,6-Dimethylnonan-3,7-diol NA 4,6-Dimethyl-7-hydroxy-4-nonen-3-one Serricorone (2S,3R)-2,3-Dihydro-3,5-dimethyl-2-ethyl-6-(l-methyl-2-oxobutyl)-4H-pyran-4-one Serricorole (2S,3R)-2,3-Dihydro-3,5-dimethyl-2-ethyl-6-(1-methyl-2-hydroxybutyl)-4H-pyran-4-one Common name Structural formula Serricornin (4S,6S,7S)-4,6-Di-methyl-7-hydroxynonan-3-one Anhydroserricornin 2,6-Diethyl-3,5-dimethyl-3,4-dihydro-2H-pyran NA 4,6-Dimethylnonan-3,7-dione NA 4,6-Dimethylnonan-3,7-diol NA 4,6-Dimethyl-7-hydroxy-4-nonen-3-one Serricorone (2S,3R)-2,3-Dihydro-3,5-dimethyl-2-ethyl-6-(l-methyl-2-oxobutyl)-4H-pyran-4-one Serricorole (2S,3R)-2,3-Dihydro-3,5-dimethyl-2-ethyl-6-(1-methyl-2-hydroxybutyl)-4H-pyran-4-one NA (not available). Open in new tab Table 4. Components of the sex pheromone of L. serricorne (Chuman et al., 1985) Common name Structural formula Serricornin (4S,6S,7S)-4,6-Di-methyl-7-hydroxynonan-3-one Anhydroserricornin 2,6-Diethyl-3,5-dimethyl-3,4-dihydro-2H-pyran NA 4,6-Dimethylnonan-3,7-dione NA 4,6-Dimethylnonan-3,7-diol NA 4,6-Dimethyl-7-hydroxy-4-nonen-3-one Serricorone (2S,3R)-2,3-Dihydro-3,5-dimethyl-2-ethyl-6-(l-methyl-2-oxobutyl)-4H-pyran-4-one Serricorole (2S,3R)-2,3-Dihydro-3,5-dimethyl-2-ethyl-6-(1-methyl-2-hydroxybutyl)-4H-pyran-4-one Common name Structural formula Serricornin (4S,6S,7S)-4,6-Di-methyl-7-hydroxynonan-3-one Anhydroserricornin 2,6-Diethyl-3,5-dimethyl-3,4-dihydro-2H-pyran NA 4,6-Dimethylnonan-3,7-dione NA 4,6-Dimethylnonan-3,7-diol NA 4,6-Dimethyl-7-hydroxy-4-nonen-3-one Serricorone (2S,3R)-2,3-Dihydro-3,5-dimethyl-2-ethyl-6-(l-methyl-2-oxobutyl)-4H-pyran-4-one Serricorole (2S,3R)-2,3-Dihydro-3,5-dimethyl-2-ethyl-6-(1-methyl-2-hydroxybutyl)-4H-pyran-4-one NA (not available). Open in new tab Very little is known on the factors which affect pheromone production in the species. Coffelt and Burkholder (1972) and Coffelt and Burkholder (1973) reported that pheromone production is higher during the first few days of adult life, reaching maximum at 4–5 d, and gradually declines thereafter until senescence. Similarly, the responsiveness to the main pheromone component increases with the age of unmated males from the first to the fourth week of adult life (Levinson and Levinson 1987). Synchrony between reproductive maturation and pheromone emission has been demonstrated in a related beetle, S. paniceum (Barratta 1977). Pheromone and other stimuli provided by female L. serricorne are perceived by the male sensory receptors located on all segments of the antennal flagellum (segments 3–11; Levinson and Levinson 1987). After a positive attraction to serricornin, unmated males lower their head and prothorax and vibrate their antennae, extend their legs, and rapidly walk around, followed by mounting of the female (Coffelt and Burkholder 1972, Levinson and Levinson 1987). Antennal elevation and leg extension may occasionally occur in the absence of the locomotory response but invariably preceded it in all positive responses (Coffelt and Burkholder 1972). Like observations in some phytophagous insect (e.g., Giga and Smith 1985, Anbutsu and Togashi 2000), female L. serricorne mark their oviposition sites with certain semiochemicals when they oviposit, so that other females can recognize the site and refrain from oviposition on the same site. The oviposition deterring compounds was identified as (2S,3R,1′S)-2,3-dihydro-3,5-dimethyl-2-ethyl-6-(l′-methyl-2′-oxobutyl)-4H-pyran-4-one (α-serricorone) and its l′ R-epimer (β-serricorone; Imai et al. 1990). As showed in Table 4, serricorone is one of the minor sex pheromone components of the insect, which suggest a bi-functional nature for the compound. Each of the two compounds has the same level of oviposition deterring activity (Imai et al. 1990). The use of oviposition deterring compounds would ensure reasonable dispersal of the species and thereby give adequate availability of space and nutrition for the larval forms when hatched (Howlader and Ambadkar 1995). Plant Volatiles Several studies (e.g., Kohno et al. 1983, Mahroof and Phillips 2007, Hori et al. 2011, Phoonan et al. 2014) have demonstrated the ability by L. serricorne to orient to plant volatiles. Attractants, that is, chemicals that cause an insect to make oriented movements towards an odor source, have also been investigated in plant derivatives and promising candidates such as β-ionone (Phoonan et al. 2014) and catechol (Nagasawa et al. 2014) have been reported. Others, including benzaldehyde, α-terpineol, linalool, S-perillaldehyde, S-limonene, β-caryophyllene, a-humulene (Hori 2004a), citronellal, and citral (Lü and Liu 2016) appears to have dual functions, i.e., they could attractant or repel the beetle depending on the concentrations of the compounds. For example, Hori (2004a) found that α-terpineol, linalool, and S-perillaldehyde strongly repelled L. serricorne at a dose of 1 μl but tend to attract the beetles at very low dose. Four other compounds, benzaldehyde, α-humulene, β-caryophyllene, and S-limonene were also attractive to the beetles at a dose of 1 μl. Repellents for L. serricorne have also been examined in plant extracts and promising chemical compound such as β-thujaplicine (hinokitiol; Hori 2004b) and rutin (Lü and Liu 2016) have been identified. Female L. serricorne respond more sensitively to plant volatiles than males (Mahroof and Phillips 2007, Hori et al. 2011). One explanation for this behavior is that female L. serricorne are likely the first-arriving or considered ‘pioneer’ beetles, thus would need to employ primary attraction (attraction without pheromone) for host finding. Upon arrival at a suitable food source, the female releases their pheromone for secondary attraction, this time, to recruit conspecific males for mating and food colonization. However, the beetles have been observed to be strongly attracted to and readily deposit their eggs on materials that are not suitable for larval food (Hori et al. 2011, Nagasawa et al. 2014). It is possible that, like other phytophagous insects (e.g., Edde and Phillips 2006) female L. serricorne may not be able to discriminate between suitable and less suitable host plants solely on the basis of plant volatiles. Alternatively, there are potential differences in the factors that govern host choice for oviposition and larval growth in the species (Nagasawa et al. 2014). Detailed understanding of the role of plant volatiles in plant–insect interactions would be beneficial in the development of tools for the enhancement of environmentally solutions for managing the pest (Nagasawa et al. 2014). Most of the pheromone traps currently in use for cigarette beetle monitoring employed the female-produced serricornin, which only attract the male beetle. To enable enhanced monitoring or control of the insect, future trap design should include the methods for reducing the population of both sexes of the beetle, thus reducing the need for chemical control. Chemical compounds that are attractive to both sexes could play a key role here. Gas chromatographic-electroantennographic (GC-EAD) studies and field studies are required to help identify and verify those chemical compounds that are attractive to both sexes of the beetle that could be used in lure development for the beetle. Runner (1919), Fletcher and Long (1976), and Fletcher (1977b) found that host plant volatiles provide ovipositional cues for mated L. serricorne females, but oviposition stimulants contained in stored food-products have been poorly examined. Nagasawa et al. (2014) reported that catechol (pyrocatechol or 1,2-dihydroxybenzene), an important constituent of coffee beans, has strong oviposition stimulant for female L. serricorne. Host plant or their volatiles may also influence pheromone production by L. serricorne or may synergistically enhance the responsiveness of the males to pheromone-baited traps (Cox 2004, Edde and Phillips 2006, Mahroof and Phillips 2007). Pheromone traps supplemented with plant-derived lures may be effective in reducing overall pest populations because such traps will capture both sexes of the beetle. Interactions with Other Organisms Microorganisms Fungi Insects need but cannot synthesize sterol and vitamins of the B group, which are indispensable for its normal functioning and development. For example, vitamins of the B group function as co-enzymes in various required enzymatic reactions in insects (Douglas 2017). In 1944, Blewett and Fraenkel demonstrated the presence of intracellular symbionts in L. serricorne larvae. The symbiont, Symbiotaphrina kochii Jurzitza (Taphrinaceae), is housed in specialized tissues (mycetomes) at the junction of the foregut and midgut of the insect (Jurzitza 1969, Noda and Kodama 1996). S. kochii helps L. serricorne to synthesize the required vitamins in considerable quantities, thus making it possible for the insect to develop on foodstuffs that are deficient in them (Fraenkel and Blewett 1943, Pant and Fraenkel 1950, Jurzitza 1969, Nasir and Noda 2003). S. kochii is transmitted to the next generation superficially on the eggs, which the larvae acquire when they eat the eggshell upon hatching (Blewett and Fraenkel 1944). L. serricorne can survive on food hosts having various harmful xenobiotics, such scopolamine, polyphenolics, and flavonoid glycosides, many of which may be harmful to other insects (Milne 1961, Dowd 1989, Shen and Dowd 1991). The species can detoxify these chemicals in plants in order to use the plants for food and reproduction. S. kochii helps in this regard by enzymatic detoxification (Dowd 1989, Shen and Dowd 1991). Several detoxifying enzymes, including aromatic ester hydrolase, glucosidase, phosphatase, and glutathione transferase have been associated with S. kochii (Shen and Dowd 1991). Other factors are also involved in the ability of L. serricorne to overcome harmful alkaloids that may be present in its food. For example, instead of biologically induced detoxification, certain alkaloids are simply secreted with little or no degradation by the insect (Blum 1983, Farnham et al. 2007). Similar mechanism has been used to explain the ability of the tobacco hornworm, Manduca sexta (L.) Lepidoptera: Sphingidae), which feeds on fresh tobacco leaves, to survive on the plant (Self et al. 1964a,b). In this regard, larvae of M. sexta are believed to readily excrete the ingested nicotine before a toxic dose could accumulate in the body of the insect, and where a large quantities of nicotine have been accumulated, the insect is able to metabolize the nicotine to less harmful alkaloids (e.g., cotinine), which the insect then excretes (Self et al. 1964a,b). The process of an absorption barrier in the digestive tract may also prevent intoxication by these toxins (Hsiao 1986). Several other fungi species have been isolated from L. serricorne (e.g., Kawakami et al. 2002, 2016; Kawakami and Takahashi 2007; Nakagawa et al. 2008). Kawakami et al. (2002), for example, isolated 17 species of molds and about 5 species of yeasts from the body surface of beetles captured in pheromone traps in Japan. Species in the genera Aspergillus, Eurotium, Penicillium, and Arthrinium were the predominant isolates. Majority of the fungal spores appears on the body surfaces and digestive tracts of the beetles. Some of these fungal pathogens are known to cause human health concerns (Kawakami and Takahashi 2007, Nakagawa et al. 2008, Kawakami et al. 2016), but the ecological function of the relations between L. serricorne and association with the respective fungus is not well understood. However, an Aspergillus sp. and several other mold complexes inoculated on tobacco disks were found to have attractant properties for L. serricorne compared with nonmold control. Moreover, penicillic and oxalic acids were nontoxic for L. serricorne, and citrinin and rubratoxin B inhibited its growth (Wright et al. 1980). Bacteria Bacillus cereus (Frankland & Frankland) (Bacillaceae) has been isolated from L. serricorne and can cause significant mortality of larvae (Thompson and Fletcher 1972). In addition to B. cereus, Yaman et al. (2008) isolated another 14 bacteria species from L. serricorne that were collected from tobacco stores in Turkey. The identified bacteria are B. megaterium, Bacillus thuringiensis, B. subtilis, B. pumilus, B. atrophaeus, B. badius, B. claussi, B. parabrevis, Brevibacterium liquefaciens, Brevibacillus parabrevis, Micrococcus luteus, Pseudomonas syringae, Staphylococcus gallinarum, and Salmonella typhimurium. Little information is available on the relationship between these bacteria and L. serricorne, while most data are focused on the gut bacterial fauna. Other Insects The cigarette beetle has several natural enemies. The most important are the parasitoids Anisopteromalus calandrae (Howard) and Lariophagus distinguendus (Foerster) (Hymenoptera: Pteromalidae) (Bare 1942, Papadopoulou and Athanassiou 2004). Both species are solitary ectoparasitoids that attack the late larval, prepupal, and pupal stages of their host. Bare (1942) published a brief description of the biology of A. calandrae, while the biology of L. distinguendus has been described by Steidle (1998). Other hymenopteran wasps parasitic on L. serricorne larvae or pupae of the cigarette beetle include Brachophagus sp. (Eurytomidae), Cephalonomia gallicola (Ashmead), Israelius carthami Richards (both Bethylidae), Theocolax elegans (Westwood), Pteromalus cerealellae (Ashmead), Norbanus sp. (Pteromalidae), and Ericydnus sipylus (Walker) (Encyrtidae) (Jones 1913, Banks 1917, Bare 1942, Rayner 1951, Richards 1952, Lim et al. 2007, Bilal et al. 2011). The following have also been mentioned as enemies of the cigarette beetles: Acari—Predatory mites, including Acarophenax lacunatus (Cross & Krantz) (Acarophenacidae), Pyemotes ventricosus (Newport), P. tritici (Lagreze-Fossat & Montane) (Pyemotidae), Tyrophagus putrescentiae (Schrank), Thyreophagus angusta (Banks) (Acaridae), Blomia gracilipes (Banks) (Echimyopodidae), Seiulus sp., Typhlodromips swirskii (Athias-Henriot) (Phytoseiidae), Cheyletus sp. (Cheyletidae), and Rhagidia sp. (Rhagidiidae), which feeds principally on immature stages of a wide range of stored product insect pest, including L. serricorne (Jones 1913, Runner 1919, Bare 1942, Rayner 1951, Papadopoulou 2006, Canevari et al. 2012, Saleh 2012). Blattisocius keegani Fox (Blattisociidae) prey on L. serricorne eggs (Rao et al. 2002). Coleoptera—The adults and larvae of Thaneroclerus buqueti (Lefebvre) and T. girodi Chevrolat (Cleridae) prey especially on L. serricorne larvae and pupae (Canzanelli 1935, Cotton and Good 1937, Rayner 1951). The Cadelle beetle, Tenebroides mauritanicus (L.) and Tribolium sometimes prey on L. serricorne pupae (Canzanelli 1935, Rayner 1951). Psocoptera—Liposcelis divinatorius (Miiller) (Liposcelididae) has been reported to attack L. serricorne eggs (Rao et al. 2002). Flight and Flight Behavior Flight Initiation and Temporal Flight Activity Papadopoulou and Buchelos (2002b) reported that adult cigarette beetles start flying 3–5 d after they emerge from the cocoon. Unlike R. dominica (Edde 2012), L. serricorne appears to require higher temperature thresholds before the species can start flying. For example, Tenhet (1961) found little or no L. serricorne activity when commodity temperatures were below 18°C. Similarly, Fardisi and Mason (2013) reported that the minimum temperature for flight initiation in the species was between 19 and 22°C, which is similar to the temperature threshold (21–23°C) needed for flight initiation in the anobiid Ptilinus ruficornis Say (Acciavatti 1972). The study by Fardisi and Mason (2013) demonstrated that flight initiation in L. serricorne is highly dependent on environmental temperature, sex, age, and mating status of individuals. For example, the minimum temperature at which virgin young, virgin old, mated young, and mated old females were observed to initiate flight was 25, 22.5, 25, and 27.5°C, respectively, while all of males, irrespective of age and mating status, would start flying was 22.5°C. Thus, males, especially young virgin males, and older virgin females of the cigarette beetles are likely to start flying earlier than other beetles of their gender and thus are more likely to be detected. L. serricorne flight activity was at the highest when the average maximum temperature was 31°C (Reed et al. 1935). Adult L. serricorne are attracted to moderate light but are repelled by sunlight or strong electric light (Reed et al. 1935). Canzanelli (1935) reported that the insect is quite active in full daylight and in the evening. Other authors (Runner 1919, Reed et al 1935, Back 1939, Howe 1957) concluded that the insect would normally take to flight toward sunset or on cloudy days, when intensity of light is low, and during darkness. According to Reed et al. (1935), peak flight activity was from about 5 p.m. until darkness. Less than 10% of the flight activity was reported to occur between 7 a.m. and 5 p.m. (USDA 1971). Like other stored-product insects, L. serricorne prefers colors of shortest wavelength or ultra violet light (UV light) (Katsuki et al. 2013, Miyatake et al. 2016). The beetle would move toward blue or blue-violet and showed the least preference for red light (Runner 1919, Katsuki et al. 2013). Male L. serricorne exhibited greater preference for UV lights than female beetles (Cuyler and Tenhet 1963). However, Katsuki et al. (2013) found that UV (375 nm) and blue (470 nm) LEDs effectively trapped both sexes of L. serricorne, irrespective of mating condition. Miyatake et al. (2016) showed the feasibility and benefits of combining UV-LED light trap with a sex pheromone for monitoring and mass trapping of L. serricorne under laboratory and field conditions. den Doop (1919) speculated that the maximum distance L. serricorne can fly is not more than 1.2 km, while Tenhet (1961) opined beetles readily fly 0.8 km, and under favorable conditions, may fly more than 1.6 km. From the flight experiments conducted by Cuyler and Tenhet (1963), beetles would fly up to 0.6 km away from the point of release, which was the farthest distance investigated by the researchers. It is likely, therefore, that the study by Cuyler and Tenhet (1963) may not reflect the maximum flight capability of this beetle. L. serricorne are fairly strong fliers and migrant beetles form an important source of L. serricorne infestation in different agricultural landscapes. Despite the interest in the distance that the cigarette beetle can fly, this important aspect of the insect behavior has not been studied in detail in either the field or laboratory perhaps because most studies have considered infestations only within buildings where adults could walk to new oviposition sites (Acciavatti 1972). Therefore, it is important to establish, flight capacity, and behavior during dispersal to determine the true significance of the insects to overall pest infestation. An important feature of the flight behavior of L. serricorne is that the insect tends to fly slow, but upward (Rayner 1951, Sivik et al. 1957, Retief and Nicholas 1988). For example, Rayner (1951) observed that of the number of cigarette beetles released by the worker, several flew over a 5.5-m roof. This behavior makes it easy for the insect to be picked up in moderate winds and wind drift, which could significantly aid in transporting them to greater distances. It is not clear if there are differences in the flight activity pattern and ability of the male and female L. serricorne. However, studies by Cuyler and Tenhet (1963) showed that more males were recaptured more than females in traps placed at the maximum distance (0.6 km) investigated. Seasonal Flight Activity The seasonal occurrence of L. serricorne varies with location and the conditions under which host commodities are stored. In heated buildings the insect is apt to be present in all stages at almost any time of the year throughout the temperate and subtropical regions, but development may be slowed during the winter (Tenhet and Bare 1951). In the tropics, L. serricorne flight activity occurs throughout the year because there is no dormant period, and the insect may have six or more generations a year (Runner 1919). For example, Jones (1913) recorded that the beetle is active and breeds continuously in the Philippines (12.8797° N, 121.7740° E), with March and April being the months of the greatest abundance of adults. Similarly, Kurup and Parkhe (1962) observed six generations of the beetle per year in India (20.5937° N, 78.9629° E). Distinct broods are rarely observed in the tropics, breeding overlap and all stages of development may constantly be present. In subtropical zones there are one to three complete generations of the beetle a year (Tenhet 1961). But unlike in a tropical climate, there are distinct broods or peaks of adult flight activity. Data obtained by Tenhet and Bare (1951) and Tenhet (1961) from observations in unheated tobacco warehouses in Richmond, VI (37.5407° N, 77.4360° W), indicated that the beetle was active from May to November. The insect passes the winter in the larval stage. Larvae begin to pupate in late spring, and the first adult of the brood appears in late May. The generation overlaps, but the spring peak is reached about the middle of June, repeating again in June/July and in early October. The population size of the first and third generations are usually smaller than the second is, but the second generation may be large in mild weather. Many of the second- and third-generation larvae constitute the overwintering populations. The length of active season for L. serricorne in the United States varies from 27 to 40 wk, depending on location. In Florida (27.6648° N, 81.5158° W) beetles may be active all year. Three generation per year has also been reported from Thessaloniki, Greece (40.6401° N, 22.9444° E; Papadopoulou and Buchelos, 2002b). The onset and cessation of the seasonal activity of the insect in the United States was similar to observation in Thessaloniki, Greece, where adult active flight activity also started in late April or mid-May, and ends in late October or later, when the ambient temperature was about 13°C (Papadopoulou and Buchelos 2002b). The population density of L. serricorne fluctuates widely. In tobacco storages the most important factors governing the density are types, grades, age, and quality of tobacco present; temperature; RH; and insect mortality from natural enemies. Control Parasites, predators, and pathogens have been found to be associated with L. serricorne in tobacco warehouses (Kearns 1934, Livingstone and Reed 1936, Bare 1942, Tenhet and Bare 1951, Tenhet 1961, Kaelin et al. 1994, Eliopoulos et al. 2002, Papadopoulou and Athanassiou 2004, Bilal et al. 2011). However, the effectiveness of natural enemies in the management of L. serricorne has mainly been quantified in the laboratory for pathogens (Kaelin et al. 1999), parasites (Gredilha et al. 2006, Cheong and Yoon 2016), and predators (Rao et al. 2002, Papadopoulou 2006b). Papadopoulou (2006b) found that the percentage of L. serricorne killed by naturally occurring T. putrescentiae in tobacco warehouses was about 20%. The most crucial point in preventing insect attack at the postharvest stages of agricultural commodities is sanitation (Tenhet 1961). Scrupulous cleanliness in the factory, wholesale, or retail establishment, including the prompt destruction or treatment of all waste material, damaged stock, etc., in which the beetles may breed, are essential components for controlling this pest. Because most of the insects attacking stored-products, including L. serricorne, live outdoor and many are great fliers, thereby enabling infestation to spread easily by insects moving in and out of facilities, screens may be installed on doors, windows, and other openings in the warehouses and manufacturing facilities to check insect movement into and out buildings and to increase the effectiveness of other control measures. Screens with openings less than 1.0 mm in width have been found adequate to prevent the passage of L. serricorne (Vinzant and Reed 1941) and other stored-product insect pests of similar body size or larger. Screens incorporated with contact insecticides could provide added advantage (Rumbos et al. 2018). Insecticide-treated nets not only act as a physical barrier, which keeps away the unwanted insect pests, they act chemically by repelling or killing the insects that come in contact with the net (Rumbos et al. 2018). Commodities should be disinfested by using proper pest elimination protocols (e.g., fumigation, extreme temperatures), before they are at once placed under the nets, under a “start clean–stay clean” concept. Managers rely on information obtained through an effective monitoring program to help to respond promptly to infestation problems. Conventional monitoring devises for stored-product insect pests include light traps, physical devices (e.g., electric grid, fly tapes), food or chemical attractants. Traps baited with a synthetic formulation of the female-produced pheromone can be used to satisfactorily monitor changes in L. serricorne population dynamics within the target area (Faustini et al. 1990). Nevertheless, other traps have also been evaluated with success (Papadopoulou and Buchelos 2002c). Pheromone traps placed 1.5–2.0 m above the floor is considered best positioning for monitoring this species in most agricultural landscapes. Pheromone-baited traps are often specific, thus help to eliminate the need to sift through the captured insects to obtain the target insect. They could also be used to pinpoint infestations, thereby requiring less time searching for cigarette beetle harborages. However, male cigarette beetles are the primary sex trapped in serricornin-baited traps, accounting for 90% or more of the population of beetles captured. Therefore, for decision making, the interpretation of cigarette beetle pheromone-baited counts should consider the fact that only one sex of the insect is trapped. The effective method of preventing L. serricorne attack when commodities must be held for considerable periods of time is to store them at low temperatures (Swingle 1938, Childs et al. 1970, Imai 2010). Imai (2010) reported the numbers of cigarette beetles captured in pheromone-baited traps decreased by 98% when tobacco leaf warehouses were moved from a warmer part of Japan to regions of the country where the daily mean temperature is 5°C for over 70 consecutive days in a year. L. serricorne life stages vary in their susceptibility to low temperatures. Childs et al. (1970) observed complete mortality of the third- and fourth-instar larvae held at 4.4°C for 3 wk. Swingle (1938) demonstrated that a temperature of 2.2°C for 2 or more consecutive weeks will kill all stages of L. serricorne infesting packaged tobacco leaves, and that exposure of infested products at −4°C for 7 d will also produce complete mortality of all life stages of the beetle. In some subtropical regions, the temperature during the winter may drop below 2.2°C for several consecutive weeks. This fact may be helpful to forecast the relative abundance of L. serricorne to be expected during the following summer by careful monitoring the temperature within the commodity during the preceding winter months. At atmospheric pressure, tobacco leaves treated with steam at about 8.7 bars would raise the temperature to 115.6°C, which quickly kills all stages of the cigarette beetle. In case of established infestations of L. serricorne within facilities, control can be achieved through heat sterilization (Hansen et al 2011, Yu et al. 2011). Typically, an enclosed area is heated to 54–60°C for 18–30 h to kill all life stages of the target pest. The actual numbers of hours the insects are exposed to lethal temperatures under this recommendation often exceed 40 h. This is because countdown to 30 h does not begin until most of the temperature sensors are at 54°C. Roesli et al. (2003) reported that trap captures of L. serricorne was significantly reduced after the heat treatment of a feed mill in Kansas, United States. The most popular and effective methods of controlling L. serricorne, and most other stored-product insects is by fumigation. The entire or part of a warehouse or manufacturing plant can be sealed up and fumigated. Fumigation of bulk commodities and structures are discussed at length in earlier publications (e.g., Bond 1984, Davis and Harein 1985, CORESTA 2009, Reichmuth 2010). When properly performed, phosphine fumigation will destroy all insect life stages in most types of packaging, while it enables a residue-free treatment. Reliance on phosphine is likely to continue for the near future because of international regulatory and market acceptance of this material and the lack of practical alternatives (Jagadeesan et al. 2012). However, evidence for genetically inherited resistance to phosphine in several storage insect pests, including L. serricorne, is accumulating in many countries due to heavy use (Rajendran and Narasimhan 1994, Zettler and Keever 1994, Jagadeesan et al. 2012, Sağlam et al. 2015). Similarly, differences have been observed in the levels of phosphine resistance among L. serricorne populations between countries and within the same country likely due to genetic variability among and within populations (Saglam et al. 2015, Coelho-Bortolo et al. 2016). Thus, the movement of infested commodities in commerce may contribute to the dissemination of resistant L. serricorne individuals, facilitating resistant gene flow (Coelho-Bortolo et al. 2016). The processes involved in the selection for phosphine resistance in L. serricorne have not been studied in detail. However, similar investigations in other stored-product insects such as R. dominica (Schlipalius et al. 2002), T. castaneum (Jagadeesan et al. 2012), and C. ferrugineus (Jagadeesan et al. 2016) have identified two major autosomal genes that are linked to the expression of phosphine resistance, but that are incompletely recessive (Collins et al. 2002). The foundation of any effective resistance management strategy should, therefore, include resistance testing system as a requirement for a good fumigation practice. This will help to recognize resistance at the early stage and limit its spread between and within geographical regions, thus prolong effectiveness of phosphine as a curative tool. Detrimental effects of long-term use of phosphine in large-scale fumigation include the fact that the compound is highly corrosive to copper, the major conductor used in many categories of electrical wiring in buildings and electronic circuitry. For example, rate of the reaction of phosphine with copper exceed that of all other metals by one order of magnitude or more (Bond et al. 1984). Direct application of contact or residual insecticides can also be used to manage L. serricorne infestation where the use of fumigation or extreme temperature is impractical. A commercial formulation of (S)-methoprene having 33.6% a.i. has been developed and gained approval of the United States Environmental Protection Agency for crack and crevice treatments in warehouses and processing facilities and is widely used in the country (EPA Reg. No: 2724-427). The lethal effect of methoprene on L. serricorne occurs during the late stage in the development of the embryo and works as an analogue to juvenile hormone, preventing the development of the pupa into the adult (Marzke et al. 1977). However, resistance to methoprene has been detected in L. serricorne populations in the United States (Benezet and Helms 1994) and other stored-product insect pests (Wijayaratne 2011, Edde 2012). A contact insecticide (e.g., pyrethrins or pyrethroids) applied in the form of aerosols or fogs of enough strength will kill the beetles that are hit by the spray particles. Pyrethroids/pyrethrins act on the insect nerve membrane at the synapses by affecting sodium ion channels (Eberhardt 1997). However, this type of treatment is only effective against insects in the airspace and will not penetrate package commodity or cracks and crevices (Tenhet 1961). Space spray treatments for cigarette beetle control should be undertaken around dusk, a time of day that coincides with the greatest period of flight activity of the insects. The effectiveness of space treatment depends upon systematic applications to kill the beetles before the eggs are deposited. Other environmental friendly technologies or compounds that have been used, tested or with potential for use for the control of L. serricorne are given below, each of which is discussed separately. Increased research efforts on these novel control measures will help to improve efficacy and overcome some of the significant barriers to their further development as control agents for stored-product insects. Entomopathogenic Viruses Viruses in the family Baculoviridae are important biological control agents of pests in agriculture and forestry. Baculoviruses tend to be species or genus specific, thus offer a viable alternative to conventional pesticides compared with other biological control agents or synthetic plant protectants. However, there is a dearth of information on the use of nuclear polyhedrosis virus (NPV) for the control of stored-product insect pests. NPVs isolated from the Indian meal moth Plodia interpunctella (Hübner) and Ephestia cautella (Walker) (Lepidoptera: Pyralidae) and have been used to control these pests in the laboratory and field (Hunter and Hoffman 1972, McGaughey 1975). The Indian Meal Moth Granulosis Virus (IMMGV) is currently approved by the regulatory agencies in the United States as a bioinsecticides for use in processing, packaging and storage areas of dried nuts, fruits, and various other commodities (EPA 2004). Polyhedrosis viruses with specific pathogenic activity against larvae of several stored-product beetle pests, including L. serricorne have also been isolated (Alfazairy et al. 2003). However, several constraints currently restrict the use of baculoviruses, and other biopesticides described below, as insecticides. These include: 1) biopesticides contain living material; thus products often have reduced shelf lives, 2) requires from days to weeks to kill the insects, 3) genetic resistance to disease infection in the host may also occur, and 4) availability of high-cost commercial formulations (Boots and Begon 1993, Bonning and Hammock 1996, Lacey et al. 2015). Endotoxins from Bacillus spp. B. thuringiensis is the most widely used microbial insecticide marketed worldwide. It is effective against many economically important forestry and field crop insect pests (Navon 2000). The bacterium is also effective against some stored-product insect pests, including L. serricorne (Thompson and Fletcher 1972, McGaughey 1994, Blanc et al. 2002, Tsuchiya et al. 2002). Another bacterium, Bacillus pulvifaciens (Nakamura), has been shown to have insecticidal effects against larvae of L. serricorne (Jackson and Long 1965). Thompson and Fletcher (1972) reported that L. serricorne was successfully controlled by B. cereus and B. thuringiensis var. thuringiensis. The required LD50 was 4.29 × 106 spores per gram of medium for B. cereus. Tsuchiya et al. (2002) have also associated 28 B.t. isolates from Japan with strong insecticidal activities against L. serricorne larvae. Kaelin et al. (1999) isolated three B. thuringiensis strains from stored tobacco residues, with high homology to B. thuringiensis subsp. tenebrionis. The 65-kDa crystal toxin present in all three strains was found to be encoded by genes identical to the cry IIIA gene responsible for the insecticidal activity of B. thuringiensis subsp. tenebrionis. The workers reported the responses of L. serricorne larvae to spore/crystal suspensions from the strains to range from 60 to 80% mortality after 7 d of ingestion. Several workers have described and written on the mode of action of Bt (e.g., Thompson and Fletcher 1972, Blanc et al. 2002, Tsuchiya et al. 2002). Endotoxins from Bacillus spp are considered ideal for pest management because of their specificity to pests and because of its lack of toxicity to humans or nontarget organisms of many crop pests (Navon 2000, Blanc et al. 2002). B. thuringiensis has been approved for application against stored-product insect pests in the United States, but little research work has been conducted with the cigarette beetle. In parallel with findings with field crop pests (reviewed in Tabashnik et al. 2013), some workers have alerted on the inevitability of B.t. resistance among stored-product insect pests (e.g., McGaughey and Beeman 1988). Entomopathogenic Fungi Entomopathogenic fungi occur in nature and belong mostly to the phyla Chytridiomycota, Zygomycota, Oomycota, Ascomycota, and Deuteromycota. However, most work related to stored-product insects is limited to Beauveria bassiana and Metarhizium anisopliae (both Ascomycota) (Rumbos and Athanassiou 2017a, Saeed et al. 2017). Both fungal species have a wide host range and their efficacy against a wide range of stored-product insect pests has been proven under laboratory conditions (Wakil et al. 2014, Rumbos and Athanassiou 2017a, Saeed et al. 2017). Two other species of Ascomycota, Purpureocillium lilacinum and Lecanicillium attenuatum, have been recovered from the cadavers of several coleopteran stored-product insect pests in Punjab, Pakistan (Wakil et al. 2014), and could have potential for insect control. The modes of action of most entomopathogenic fungi are similar and have been described in detail by several authors, including Aw and Hue (2017). Unlike B.t. or viruses, ingestion is not essential for a fungal infection to occur. The pathogens are easily transmitted from insect to insect by contact. Entomopathogens are environmentally friendly, have a broad spectrum of hosts, and most formulations can be applied with the same technical means as conventional contact insecticides. Other opportunities and future challenges for the use of entomopathogenic fungi against stored-product insects have been highlighted by Rumbos and Athanassiou (2017a). Still, there is inadequate information on the efficacy of these compounds against L. serricorne, which is an area that merits additional investigation. Nematodes Certain entomopathogenic nematodes, including Heterorhabditis bacteriophora Poinar (Nematoda: Heterorhabditidae), Heterorhabditis megidis Poinar, Jackson and Klein (Nematoda: Heterorhabditidae) and two strains of Steinernema carpocapsae Weiser (Nematoda: Steinernematidae) and Steinernema feltiae Filipjev (Nematoda: Steinernematidae) have been applied for the control of L. serricorne, with limited results (Rumbos and Athanassiou 2012). Further experimental work is required to optimize the conditions under which these entomopathogenic nematodes could be used for the effective control of stored-product pests. An important concern on the applicability of these agents in the stored-product environment is that these agents require some certain moisture levels to act (Rumbos and Athanassiou 2017b), which may not be conducive for the dry environment encountered in storage. Spinosad This is a chemical class of insecticide, consisting of a mixture of two active components, spinosyn A and spinosyn D derived from a rare naturally occurring soil-dwelling actinomycete bacterium called Saccharopolyspora spinosa (Kirst et al. 1992). The insecticide has both contact and systemic effects and causes excitation of the insect nervous system, leading to involuntary muscle contractions, prostration with tremors, and finally paralysis (Salgado 1998). The efficacy of spinosad has been demonstrated against important storage insect pests, including L. serricorne, and E. elutella (Blanc et al. 2004, Hertlein et al. 2011). The material has been registered in the United States since 2005 for use on stored commodities (Anonymous 2005) and in several countries (reviewed in Hertlein et al. 2011). In general, L. serricorne is considered moderately susceptible to spinosad, as it can survive dose rates that are effective in the case of other major stored-product insects (Getchell and Subramanyam 2007, Hertlein et al. 2011). However, spinosad in currently not widely used in stored-product insect pest management (IPM) in the United States. Commercial formulation of the compound, Sensat, was made available in the United States in May 2018. Moreover, Sensat is currently not registered for use on tobacco. Diatomaceous Earth Diatomaceous earths (DEs) are natural silica powder based on fossilized skeletons of diatoms, having over 90% amorphous SiO2. Several formulations of the material have been registered in many countries, including the United States, for use as grain protectants, crack and crevice treatments, and structural treatments (Athanassiou et al. 2006, Korunić 2013, Shah and Khan 2014). When DE is picked up by an arthropod, the sharp-edged silica destroys the wax layers of the cuticle and material quickly absorb the epicuticular lipids and body fluids, leading to desiccation and death of the arthropod (Stanley et al. 1962, Korunić 2013). According to Korunić (2013), the most sensitive storage insect pests to DEs are the beetle species in the genus Cryptolestes (Laemophloeidae), and the most tolerant are those of the genus Prostephanus (Bostrychidae). The efficacy of DE is affected by several factors, including its source, temperature, humidity, and characteristics of target pests and substrate (Shah and Khan 2014). To reduce bulkiness and improve efficacy, DEs are often enhanced with other insecticidal materials to give control at much lower doses (Korunić 2013, Shah and Khan 2014). Examples of these enhanced formulations are two DE formulations with the synergistic mode of physical and chemical actions; one with the addition of the soil bacteria metabolite abamectin (DEA) and the other with the addition of the plant extract bitterbarkomycin (DEBBM; Athanassiou et al. 2006). Working at 24 to 26°C and 90–95% RH, Korunić (2010) found that a commercial formulation of an enhanced DE used at 500 ppm cause 100% mortality of L. serricorne populations within 5 d exposure period. A potential tolerance or resistance development should, however, be considered when choosing DE as to protect commodities due to the possibility of genetic resistance developing in the target organisms (Korunić, 2013). This cautionary note is based on the findings by Fields (2003), who successfully increased the tolerance of three stored-product insect pests to a DE formulation in the laboratory over a 3-yr period. Still, the application of DE and other inert materials in tobacco ecosystems has not been evaluated in detail and needs added studies. Irradiation Insect exposure to ionizing radiation may cause sterilization through induction of dominant lethal mutation in the genetic material (Knipling 1955). The doses needed to prevent the reproduction of stored-product pests range from 50 Gy for T. molitor to 450 Gy for the Angoumois moth, Sitotroga cereallela (Olivier) (Lepidoptera: Gelechiidae) (Hallman 2013). For L. serricorne, the dose reported varies from 100 Gy (Imai et al. 2006) to 300 Gy (Kongratarpon et al. (2003). Irradiation shortens the life span of insects by accelerating senescence and may also cause protein denaturation and impair enzymatic activity (Salem et al. 2014). According to Kongratarpon et al. (2003), the LD50 and LD99 for L. serricorne eggs are 327 Gy (at 7 d exposure period) and 834 Gy (at 3 d exposure period), respectively; LD50 and LD99 for the last instar larvae 505 Gy (at 7 d exposure period) and 1,547 Gy (at 12 d exposure period); and for the adults are 931 Gy (at 3 d exposure period) and 1,746 Gy (at 12 d exposure period), respectively. In general, the adults of most stored-product insects (e.g., Tilton et al. 1966) are the most tolerant to the effects of gamma radiation, followed by the pupae, the larvae, and the eggs. And among adults of the same species, females appear more sensitive to sterilizing effects of irradiation than males (Tilton and Brower 1987). In addition, the dose needed to produce 100% mortality in several life stages is depended entirely on the insect’s age at the time of treatment (Runner 1916, Manoto et al. 1976). Cornwell and Bull (1960), Tilton and Brower (1987), and Hallman (2013) published detailed review papers that summarizes and discusses information on various aspects of the use of ionizing radiation for the control of insect infestation in grains and grain product. Although the technical feasibility of using radiation for commodities to cause insect mortality without affecting product quality is now proven, widespread adoption of the technology is currently being hampered by certain factors, including the ease of bulk handling of commodities and integration of irradiation treatment into established product handling procedures (Cornwell and Bull 1960). Controlled Atmosphere Storage insects are aerobic organisms requiring oxygen for their survival. Oxygen concentrations within an enclosure can be altered to within a few percent of setpoints through the addition of N2 or scrubbing of CO2 to kill the arthropods. Controlled atmospheres that are insecticidal typically have ≥20% CO2 and/or ≤1% O2, with the rest of the atmosphere composed of N2 gas (Mitcham et al. 2006). These atmospheres also prevent mold growth and maintain product quality. The mechanism used by insects to respond to low O2 environments has been proposed by Zhou et al. (2000) and Mitcham et al. (2006). Several authors (e.g., Harein and Press 1968, Jay et al. 1971, Mbata and Phillips 2001, Bera et al. 2007, Yang et al. 2008) have also reported on the mortality of stored-product insect pests under various controlled atmosphere treatments. The efficacy of control atmosphere as a pest management tool is dependent on several factors, including temperature, RH, and life stage of the insect, and efficiency in the delivery and maintenance of the concentration of gas needed in the chamber (Jay 1971, Mbata and Phillips 2001, Mitcham et al. 2006). For example, for best results, treatment should be conducted when temperatures exceed 21°C due to enhanced respiratory demand (Jay et al. 1971). Studies conducted at Altria Inc. (unpublished data) showed that the fourth-instar larva and prepupal of L. serricorne life stages appear to be most tolerant to CO2. Carbon dioxide increases the toxicity of a number of fumigants to insects, thus controlled atmosphere and fumigation may be used in a complementary way to increase the effectiveness of treatment (Bond 1989). Although the exact parameters for control and kill of the different life stages of L. serricorne and the tobacco moth Ephestia elutella Hübner (Lepidoptera: Pyralidae) in bulk tobacco has been defined (CORESTA 2013), the technology has not been widely adopted by the tobacco industry despite their efficacy. Capital and running costs of the equipment are among the major limiting factors. Molecular Genetics and Genomics Approach Molecular data can help to illuminate the genetic architecture of a number of phenotypic traits that enable an insect to continue to be an agricultural pest and can also be used to identify new gene targets for developing control measures (Leptinotarsa Sequencing Consortium 2018, Tribolium Sequencing Consortium 2008, Park and Beeman 2008). The transcriptome of L. serricorne is currently being worked out. But the mitochondrial genome of the beetle has been sequenced by Yang et al. (2017). According to these workers, the mitochondrial genome is about 15,958 bp and composed of 13 protein-coding genes (PCGs), two rRNA genes, 22 tRNA genes, and a control region. Sequence-specific RNAi holds great promises in stored-product insect pests management, especially for their use for knocking down the expression of genes to determine their functions. In T. castaneum, for example, efforts have been made to investigate genes involved in cuticle breakdown and synthesis during molting, a very vulnerable stage for insects, making it a target for development of new insect control tools (Throne 2010). Kirk et al. (2013) and Mamta and Rajam (2017) have published detailed reviews on the potential of new molecular genetics and genomic techniques for effective management of insect pests. The review also discusses the important parameters, which must be considered for genetic control of insect pests without producing any effect on nontarget organisms and environment. A recent review of Colorado potato beetle, Leptinotarsa decemlineata Say (Coleoptera: Chrysomelidae) outline steps required for commercialization of RNAi-based insecticide for its control (Palli 2014). The Behavioral Approaches e.g., Mating Disruption It is possible to improve the insect captures in pheromone traps to the point where they go beyond their convectional use for monitoring functions. One of these novel ideas is the application of mating disruption; in which large amount of synthetic sex pheromone is released into a space to create an atmosphere that is difficult for males to find females for mating, thereby causing population suppression or extinction. A mating disruption product consisted of a dispenser that holds copious quantities of the sex pheromone. These dispensers are placed in the area to be treated where they release strong plumes of pheromones (Mahroof and Phillips 2014). Mating disruption for stored-product moths such as Plodia and Ephestia species is commercially available as a component of an integrated pest management-based control strategy in the food industry. Nevertheless, it seems that the control of L. serricorne through mating disruption is likely to occur much more gradually than that in the case of stored-product moths, for which the “shut down” of trap captures and suppression of progeny production is more rapid (reviewed in Trematerra et al. 2011, Mahroof and Phillips 2014). At least one mating disruption formulation is now commercially available for L. serricorne, and further testing will evaluate the feasibility of using the method in “real world” applications. Summary L. serricorne, a stored-product insect, is cosmopolitan. The insect attacks a variety of dried substances, but the most common food is cured tobacco leaves. Infestation by this insect is more prevalent in agricultural commodity warehouses and manufacturing facilities. Details of the bioecology of the cigarette beetle were presented in this review. This information is essential because a whole-system approach to IPM requires a thorough knowledge of the target insect. Integration and effective use of various control components (IPM) are often enhanced when the target organism is correctly identified, and its biology and ecology are known. At present, control of the cigarette beetle still relies on the use of phosphine fumigation and contact insecticides. However, both materials are becoming less effective as the insect develops resistance to the compounds. The goal should be to minimize the selection for resistance to any type of pesticide, or to help regain susceptibility in insect pest populations in which resistance has already developed. IPM is therefore important in keeping the efficacy of valuable insecticides. Non-chemical methods of control are also used in the management of the beetle, but these too are prone to mixed or irregular results. Other ecologically sound options with potentials for the management of L. serricorne, and related stored-product insect pests, are reviewed. Recommendations for Future Research Of more immediate concern is the development of tools for effective monitoring of the beetle. Most of the pheromone traps currently in use for cigarette beetle monitoring employed the female-produced serricornin, which only attract the male beetle. To enable enhanced monitoring or control of the insect, future trap design should include the methods for reducing the population of both sexes of the beetle, thus reducing the need for chemical control. Chemical compounds that are attractive to both sexes could play a key role here. Several basic and applied questions concerning nutritional ecology and ability of L. serricorne to adapt to different food sources need answers to aid the development of sustainable management strategies. For example, breeding experiments are required to help determine the true range of materials liable to infestation. A biomarker that distinguishes different hosts would also be useful to identify the food use pattern of L. serricorne, and other stored-product insects. Despite the interest in the distance that the cigarette beetle can fly, this important aspect of the insect behavior has not been studied in detail in either the field or the laboratory. It is also not clear if there are differences in the flight activity pattern and ability of the male and female L. serricorne. Therefore, it is important to establish, flight ability, and behavior during dispersal to determine the true significance of the insects to overall pest infestation. While most data are focused on the gut bacterial fauna, a variety of bacteria and fungi have been isolated from the body surface of L. serricorne. Still the ecological function of the relations between L. serricorne and association with the respective microorganism is not well understood. Reliance on phosphine is likely to continue for the near future due to lack of practical alternatives, among other factors. Thus, it is essential to design technologies that would prolong effectiveness of the compound as a curative tool. The processes involved in the selection for phosphine resistance in L. serricorne also need to be studied. In addition, other insecticides with potential for use for the control of L. serricorne need to be further investigated and registered. Acknowledgments I thank Drs Christos Athanassiou, Greg Daglish, and Pali Subba for their comments and suggestions on this manuscript. References Cited Acciavatti R. E . 1972 . The ecology of Anobiidae (Coleoptera) associated with northern hardwood forests in Central New York, with special reference to Ptilinus ruficornis Say . Ph.D. dissertation, State University College of Forestry, Syracuse University , Syracuse . Google Preview WorldCat COPAC Adler C . 2002a . Efficacy of high temperatures to control Lasioderma serricorne and Rhyzopertha dominica. Integr . Prot. Stored Prod. IOBC Bull 25 : 187 – 191 . Adler C . 2002b . Efficacy of heat treatments against the tobacco beetle Lasioderma serricorne F. (Col., Anobiidae) and the lesser grain borer Rhyzopertha dominica F. (Col., Bostrichidae), pp. 617 – 621 . In P. F. Credland , D. M. Armitage , C. H. Bell , P. M. Cogan , and E. Highley (eds.), Proceedings of the 8th International Working Conference on Stored Product Protection , 22–26 July 2002 , York, UK . CAB International , Wallingford . Alfazairy A. A. , A. A. Hendi A. M. El-Minshawy , and H. H. Karam . 2003 . Entomopathogenic agents isolated from 19 coleopteran insect pests in Egypt . Egypt. J. Biol. Pest Co . 13 : 125 . WorldCat Alfieri A . 1932 . Les insectes de la tombe de Toutankhamon . B. So Entomol d’Egypte 24 : 188 – 189 . WorldCat Allotey J. , and I. E. Unanaowo . 1993 . Aspects of the biology of Lasioderma serricorne (F.) on selected food media under tropical conditions . Int. J. Trop. Insect Sci . 14 : 595 – 601 . Google Scholar Crossref Search ADS WorldCat Anbutsu H. and K. Togashi . 2000 . Deterred oviposition response of Monochamus alternatus (Coleoptera: Cerambycidae) to oviposition scars occupied by eggs . Agric. For. Entomol . 2 : 217 – 223 . Google Scholar Crossref Search ADS WorldCat Anonymous . 2005 . Spinosad; pesticide tolerance . Fed. Regist. 70 : 1349 – 1357 . Arango R. A. and D. K. Young , 2012 . Death-watch and spider beetles of Wisconsin—Coleoptera: Ptinidae . Forest Service, Forest Products Laboratory, General Technical Report FPL-GTR-209. USDA , Beltsville, MD . Google Preview WorldCat COPAC Ashworth J. R . 1993 . The biology of Lasioderma serricorne . J. Stored Prod. Res . 29 : 291 – 303 . Google Scholar Crossref Search ADS WorldCat Athanassiou C. G. , Z. Korunic , N. G. Kavallieratos , G. G. Peteinatos , M. C. Boukouvala , and N. H. Mikeli . 2006 . New trends in the use of diatomaceous earth against stored-grain insects, pp. 730 – 740 . In I. Lorini , B. Bacaltchuk , H. Beckel , E. Deckers , E. Sundfeld , J. P. dos Santos , J. D. Biagi , J. C. Celaro , L. R. D’A. Faroni , L. de O. F. Bortolini , M. R. Sartori , M. C. Elias , R. N. C. Guedes , R. G. da Fonseca , and V. M. Scussel (eds.), Proceedings of the 9th International Working Conference on Stored-Product Protection , 15–18 October 2006 , Campinas, Sao Paulo, Brazil . ABRAPOS , Passo Fundo, Rio Grande do Sul, Brazil . Athanassiou C. G. , J. Riudavets , and N. G. Kavallieratos . 2011 . Preventing stored product insect infestations in packaged-food products . Stewart Postharvest Rev . 3 : 8 . WorldCat Aw K. M. S. , and S. M. Hue . 2017 . Mode of infection of Metarhizium spp. fungus and their potential as biological control agents . J Fungi (Basel). 3 . doi: WorldCat Crossref Back E. A . 1939 . The cigarette beetle as a pest of cottonseed meal . J. Econ. Entomol . 32 : 739 – 749 . Google Scholar Crossref Search ADS WorldCat Banks N . 1917 . New mites, mostly economic (Arach., Acar.) . Entomol. News . 28 : 193 – 199 . WorldCat Bare C. O . 1942 . Some natural enemies of stored-tobacco insects, with biological notes . J. Econ. Entomol . 35 : 185 – 189 . Google Scholar Crossref Search ADS WorldCat Barratta B. I. P . 1977 . Sex pheromone emission by female Stegobium paniceum (L.) (Coleoptera: Anobiidae) in relation to reproductive maturation and oviposition . Bull. Entomol. Res . 67 : 491 – 499 . Google Scholar Crossref Search ADS WorldCat Benelli G. , D. Romano , N. Kavallieratos , G. Conte , C. Stefanini , M. Mele , C. Athanassiou , and A. Canale . 2017 . Multiple behavioural assymetries impact male mating success in the khapra beetle, Trogoderma granarium . J. Pest Sci . 90 : 901 – 909 . Google Scholar Crossref Search ADS WorldCat Benezet H. J. , and C. W. Helms . 1994 . Methoprene resistance in the cigarette beetle, Lasioderma serricorne (F.) (Coleoptera: Anobiidae) from tobacco storages in the southeastern United States . Resist. Pest Manage. Newslett . 6 : 12 – 14 WorldCat Bera A. S. N. Sinha A. Gaur , and C. Srivastava . 2007 . Effect of carbon dioxide rich atmosphere on storage insects and fungi . Indian J. Agric. Sci . 77 : 756 – 761 . WorldCat Bhalodia N. K. , and M. S. Chari . 1976 . Bionomics of cigarette beetle Lasioderma serricorne F. (Anobiidae: Coleoptera) . Gujarat Agric. Univ. Res. J . 2 : 5 – 14 . WorldCat Bharathi J. L. , U. Srieedhar , B. Kishore , and J. V. Prasad . 2001 . Life table studies of cigarette beetle, Lasioderma serricorne Fab. on FCV, burley and cigar wrapper tobacco . Tob. Res . 27 : 147 – 156 . WorldCat Bilal H. , A. M. Basheer , and A. T. Saleh . 2011 . Survey and abundance of common parasitoid and predatory species of the cigarette Beetle, Lasioderma serricorne (F.) (Coleoptera: Anobiidae) in tobacco storage warehouses in Syria . Egypt. J. Biol. Pest Cont . 21 : 151 – 153 . WorldCat Blanc M. , P. Kaelin , and F. Gadani . 2002 . Bacillus thuringiensis (Bt) for the control of insect pests in stored tobacco: a review . Beitr Tabakforsch Int. /Contrib. Tob. Res . 20 : 15 – 21 . WorldCat Blanc M. P., C. Panighini F. Gadani , and L. Rossi . 2004 . Activity of spinosad on stored-tobacco insects and persistence on cured tobacco stripst . Pest Manag. Sci . 60 : 1091 – 1098 . Google Scholar Crossref Search ADS PubMed WorldCat Blanc M. P., N. Lugon-Moulin C. Panighini H. Pijnenburg , and L. Rossi . 2006 . Structure of worldwide populations of Lasioderma serricorne (Coleoptera: Anobiidae) as revealed by amplified fragment length polymorphism profiles . Bull. Entomol. Res . 96 : 111 – 116 . Google Scholar Crossref Search ADS PubMed WorldCat Blewett M. and G. Fraenkel . 1944 . Intracellular symbiosis and vitamin requirements of two insects, Lasioderma serricorne and Sitodrepa panicea . Proc. Royal Soc. B: Biol. Sci . 132 : 212 – 221 . WorldCat Blum M. S . 1983 . Detoxication, deactivation, and utilization of plant compounds by insect, pp. 265 – 275 . In P. A. Hedin (ed.), ACS Symposium Series Vol. 208. America Chemical Society , Washington, DC . doi: 10.1021/bk-1983-0208.ch015 Google Preview WorldCat COPAC Bond E. J . 1989 . Manual of fumigation for insect control . FAO Plant Production and Protection Papers No. 54, Food and Agriculture Organization of the United Nations (FAO) , Rome, Italy . Google Preview WorldCat COPAC Bond E. J. , T. Dumas , and S. Hobbs . 1984 . Corrosion of metals by the fumigant phosphine . J. Stored Prod. Res . 20 : 57 – 63 . Google Scholar Crossref Search ADS WorldCat Bonning B. C. , and B. D. Hammock . 1996 . Development of recombinant baculoviruses for insect control . Ann. Rev. Entomol . 41 : 1 91 – 210 . Google Scholar Crossref Search ADS WorldCat Boots M. , and M. Begon . 1993 . Trade-off with resistance to a granulosis virus in the Indian meal moth, examined by a laboratory evolution experiment . Funct. Ecol . 7 : 528 – 534 . Google Scholar Crossref Search ADS WorldCat Borror D. J. , C. A. Triplehorn , and N. F. Johnson . 1992 . An introduction to the study of insects . Harcourt Brace College Publishers , New York, NY . Google Preview WorldCat COPAC Boving A. G . 1927 . The larva of Nevermannia dorcatomoides Fisher, with comments on the classification of the Anobiidae according to their larvae (Coleoptera: Anobiidae) . Proc. Entomol. Soc. Wash . 29 : 51 – 72 . WorldCat Buchelos C. Th. , and A. R. Levinson . 1993 . Efficacy of multisurface traps and Lasiotraps with and without pheromone addition, for monitoring and mass-trapping of Lasioderma serricorne F. (Col., Anobiidae) in insecticide-free tobacco stores . J. Appl. Entomol . 116 : 440 – 448 . Google Scholar Crossref Search ADS WorldCat Canevari G. D. C. , F. Rezende , R. B. Da Silva , L. R. D’Antonino Faroni , J. C. Zanuncio , S. Papadopoulou , and J. E. Serrão . 2012 . Potential of Tyrophagus putrescentiae (Schrank) (Astigmata: Acaridae) for the biological control of Lasioderma serricorne (F.) (Coleoptera: Anobiidae) . Braz. Arch. Biol. Tech . 55 : 299 – 303 . Google Scholar Crossref Search ADS WorldCat Canzanelli A . 1935 . Contributo alla embriologia e biologia deltarlo del tabacco (Lasioderma serricorne Fabricius) . Boll. Lab. Zool. Agr. Bachic . 4 : 81 – 116 . WorldCat Cheong S. W. , and C. S. Yoon ; Changwon National University. Industry-Academy Cooperation Foundation . 2016 . Method for controlling Lasioderma serricorne using Anisopteromalus apiovorus . U.S. patent 14/491,975. Google Preview WorldCat COPAC Childs D. P. , J. E. Overby , and B. J. Watkins . 1968 . Low temperature effect on cigarette beetle infestation in tobacco hogsheads . J. Econ. Entomol . 61 : 992 – 996 . Google Scholar Crossref Search ADS WorldCat Childs D. P. , J. E. Overby , B. J. Watkins , and D. Niffenegger . 1970 . Low temperature effect upon third- and fourth-Instar cigarette beetle larvae . J. Econ. Entomol . 63 : 1860 – 1864 . Google Scholar Crossref Search ADS WorldCat Chuman T. , M. Kohno , K. Kato , and M. Noguchi . 1979 . 4,6-Dimethyl-7-hydroxy-nonan-3-one a sex pheromone of the cigarette beetle (Lasioderma serricorne F.) . Tetrahedron Letter . 20 : 2361 – 2364 . Google Scholar Crossref Search ADS WorldCat Chuman T. , M. Kohno , K. Koto , M. Noguchi , H. Nomi , and K. Mori . 1981 . Stereoselective synthesis of erythro-serricornin, (4R,6R,7S)-, and (4S,6R,7S)-4,6-dimethyl-7-hydroxynonan-3-one, stereoisomers of the sex pheromone of cigarette beetle . J. Agric. Biol. Chem . 45 : 2019 – 2023 . WorldCat Chuman T. , K. Mochizuki , M. Mori , M. Kohno , M. Ono , and I. Onish . 1982 . Pheromone activity of (±)-serricornins for male cigarette beetle (Lasioderma serricorne F.) . Agric. Biol. Chem . 46 : 593 – 595 . WorldCat Chuman T., K. Mochizuki M. Mori M. Kohno K. Kato , and M. Noguchi . 1985 . Lasioderma chemistry sex pheromone of cigarette beetle (Lasioderma serricorne F.) . J. Chem. Ecol . 11 : 417 – 434 . Google Scholar Crossref Search ADS PubMed WorldCat Coelho-Bortolo T., C. A. Mangolin , and A. S. Lapenta . 2016 . Genetic variability in the natural populations of Lasioderma serricorne (F.) (Coleoptera: Anobiidae), detected by RAPD markers and by esterase isozymes . Bull. Entomol. Res . 106 : 47 – 53 . Google Scholar Crossref Search ADS PubMed WorldCat Coffelt J. A . 1975 . Multiple mating by Lasioderma serricorne (F.) - effects on fertility and fecundity, pp. 549 – 553 . In E. U. Brady , J. H. Brower , P. E. Hunter , E. G. Jay , P. T. M. Lum , H. O. Lund , M. A. Mullen , and R. Davis (eds.), Proceedings of the 1st International Working Conference on Stored-Product Entomology , 7–11 October 1974 , Savannah, GA . University of Georgia, Athens, GA and USDA/ARS , Washington, DC. Coffelt J. A. , and W. E. Burkholder . 1972 . Reproductive biology of the cigarette beetle, Lasioderma serricorne 1. Quantitative laboratory bioassay of the female sex pheromone from females of different ages . Ann. Entomol. Soc. Am . 65 : 447 – 450 . Google Scholar Crossref Search ADS WorldCat Coffelt J. A. , and W. E. Burkholder . 1973 . Reproductive biology of the cigarette beetle, Lasioderma serricorne 2. Ovarian development and female sexual maturity . Ann. Entomol. Soc. Am . 66 : 368 – 372 . Google Scholar Crossref Search ADS WorldCat Coffelt J. A. , and K. W. Vick . 1973 . A black mutation of Lasioderma serricorne (F.) (Coleoptera: Anobiidae) . J. Stored Prod. Res . 9 : 65 – 70 . Google Scholar Crossref Search ADS WorldCat Collins P. J., G. J. Daglish M. Bengston T. M. Lambkin , and H. Pavic . 2002 . Genetics of resistance to phosphine in Rhyzopertha dominica (Coleoptera: Bostrichidae) . J. Econ. Entomol . 95 : 862 – 869 . Google Scholar Crossref Search ADS PubMed WorldCat CORESTA . 2009 . Guide No. 2: Phosphine fumigation parameters for the control of cigarette beetle and tobacco moth . https://www.coresta.org/sites/default/files/technical_documents/main/Guide-No02-Fumigation_Oct13.pdf. Accessed 6 September 2018. CORESTA . 2013 . Guide No. 12. Controlled atmosphere parameters for the control of cigarette beetle and tobacco moth . https://www.coresta.org/sites/default/files/technical_documents/main/Guide-No12_Controlled-Atmosphere_May13.pdf. Accessed 8 September 2018. Cornwell P. B. , and J. O. Bull 1960 . Insect control by gamma irradiation: an appraisal of the potentialities and problems involved . J. Sci. Food Agric . 11 : 754 – 768 . Google Scholar Crossref Search ADS WorldCat Cotton R. T. , and N. E. Good . 1937 . Annotated list of the insects and mites associated with stored grain and cereal products, and of their arthropod parasites and predators . Miscellaneous Publication No. 258, USDA , Beltsville, MD . Google Preview WorldCat COPAC Cox P. D . 2004 . Potential for using semiochemicals to protect stored products from insect infestation . J. Stored Prod. Res . 40 : 1 – 25 . Google Scholar Crossref Search ADS WorldCat Crumb S. E. , and F. S. Chamberlin . 1934 . The effect of cool temperatures on some stages of the cigarette beetle . Fla. Entomol . 18 : 11 – 14 . Google Scholar Crossref Search ADS WorldCat Cuyler R. D. , and J. N. Tenhet . 1963 . Distance the cigarette beetle is attracted to black-light . SPIB Internal report, USDA , Beltsville, MD . Google Preview WorldCat COPAC Davis R. , and P. K. Harein . 1985 . Fumigation of structures, pp. 161 – 170 . In F. Baur (ed.), Insect management for food storage and processing , 2nd printing, American Association of Cereal Chemists , St. Paul, MN . Google Preview WorldCat COPAC Dick J . 1937 . Oviposition in certain coleoptera . Ann. Appl. Biol . 24 : 762 – 796 . Google Scholar Crossref Search ADS WorldCat den Doop J. E . 1919 . Aanteekeningen over de Lasioderma bestrijding. Tweede Serie No. III, Mededeelingen van het Deli Proefstation te Medan , Medan, Naaml. venn. "De Deli courant,” Sumatra, Indonesia . Google Preview WorldCat COPAC Douglas A. E . 2017 . The B vitamin nutrition of insects: the contributions of diet, microbiome and horizontally acquired genes . Curr. Opin. Insect Sci . 23 : 63 – 69 . WorldCat Dowd P. F . 1989 . In situ production of hydrolytic detoxifying enzymes by symbiotic yeasts in the cigarette beetle (Coleoptera: Anobiidae) . J. Econ. Entomol . 82 : 396 – 400 . Google Scholar Crossref Search ADS WorldCat Eberhardt H-J . 1997 . Alternative forms of storage protection: biological insecticides for the control of the cigarette beetle. (Lasioderma serricorne) and the tobacco moth (Ephestia elutella) . Beitr. Tabakforsch. Int. /Contrib. Tob. Res . 17 : 31 – 47 . WorldCat Edde P. A . 2012 . A review of the biology and control of Rhyzopertha dominica (F.) the lesser grain borer . J. Stored Prod. Res . 48 : 1 – 18 . Google Scholar Crossref Search ADS WorldCat Edde P. A . 2018 . Principal insects affecting tobacco plants in the field . Beitr Tabakforsch Int. /Contrib. Tob. Res . 28 : 117 – 165 . WorldCat Edde P. A. , and T. W. Phillips . 2006 . Potential host affinities for the lesser grain borer, Rhyzopertha dominica: behavioral responses to host odors and pheromones and reproductive ability on non-grain hosts . Entomol. Exp. Appl . 119 : 253 – 263 . Google Scholar Crossref Search ADS WorldCat Eliopoulos P. A. , C. G. Athanasiou and C. H. Buchelos . 2002 . Occurrence of Hymenoptera parasitoids of stored product pests in Greece . IOBC WRPS Bull . 25 : 127 – 139 . EPA (U.S. Environmental Protection Agency) . 2004 . Indian meal moth Granulosis Virus PC Code 108896 , Biopesticides Registration Action Document , Washington, D.C. Google Preview WorldCat COPAC Fabricius J. C . 1792 . Entomologia systematica emendata et aucta. Secundum classes, ordines, genera, species adjectis synonimis, locis, observationibus, descriptionibus . Entomol. Systematica . 1 : 1 – 330 . [Article in Latin]. WorldCat Fall H. C . 1905 . Revision of the Ptinidae of boreal America . T. Am. Entomol. Soc . 31 : 97 – 296 . WorldCat Fardisi M. , and L. J. Mason . 2013 . Influence of temperature, gender, age, and mating status on cigarette beetle (Lasioderma serricorne (F.)) (Coleoptera: Anobiidae) flight initiation . J. Stored Prod. Res . 52 : 93 – 99 . Google Scholar Crossref Search ADS WorldCat Farnham A. S. , W. J. W. Flora , S. S. Ingram , and D. L. Faustini . 2007 . No evidence of substantial nicotine metabolism by Lasioderma serricorne (Fabricius) (Coleoptera: Anobiidae) reared on tobacco . J. Stored Prod. Res . 43 : 171 – 176 . Google Scholar Crossref Search ADS WorldCat Faustini D. L. , A. V. Barak , W. E. Burkholder , and J. Leos-Martinez . 1990 . Combination –type trapping for monitoring stored-product insects: a review . J. Kansas Entomol. Soc . 63 : 539 – 547 . WorldCat Fields P. G . 2003 . Laboratory selection for resistance to diatomaceous earth, pp. 776 – 778 . In P. F. Credland , D. M. Armitage , C. H. Bell , P. M. Cogan , and E. Highley (eds.), Proceedings of the 8th International Working Conference on Stored Product Protection , 22–26 July 2002 , York, UK . CAB International , Wallingford, UK . Fletcher L. W . 1977a . A procedure for collecting large numbers of eggs of Lasioderma serricorne (F.) . J. Stored Prod. Res . 13 : 87 – 88 . Google Scholar Crossref Search ADS WorldCat Fletcher L. W . 1977b . Influence of food odors on oviposition by the cigarette beetle on nonfood materials . J. Econ. Entomol . 64 : 770 – 771 . Google Scholar Crossref Search ADS WorldCat Fletcher L. W. , and J. S. Long . 1976 . Cigarette beetle oviposition and egg hatch at selected temperatures . Tob. Sci . 20 : 172 – 173 . WorldCat Fraenkel G. , and M. Blewett . 1943 . The basic food requirements of several insects . J. Exp. Biol . 20 : 28 – 34 . WorldCat Fullaway D. T . 1914 . Tobacco insect in Hawaii . Bulletin No. 34, Hawaii Agricultural Experiment Station , Honolulu, HI . Google Preview WorldCat COPAC Gautam S. G. , G. P. Opit , D. Margosan , J. S. Tebbets , and S. Walse . 2014 . Egg morphology of key stored-product insect pests of the United States . Ann. Entomol. Soc. Am . 107 : 1 – 10 . Google Scholar Crossref Search ADS WorldCat Gay L. , P. E. Eady , R. Vasudev , D. Hosken , and T. Tregenza . 2009 . Costly sexual harassment in a beetle . Physiol. Entomol . 34 : 86 – 92 . Google Scholar Crossref Search ADS WorldCat Getchell A. I. , and Bh. Subramanyam . 2007 . Evaluation of a dry spinosad formulation on two extruded pet foods for controlling four stored-product insects . Biopestic. Int . 3 : 108 – 116 . WorldCat Giga D. P. , and R. H. Smith . 1985 . Oviposition markers in Callosobruchus maculatus F. and Callosobruchus rhodesianus Pic. (Coleoptera, Bruchidae): Asymmetry of interspecific responses . Agr. Ecosyst. Environ . 12 : 229 – 233 . Google Scholar Crossref Search ADS WorldCat Gredilha R. , A. R. Carvalho , A. F. Lima and R. P. Mello . 2006 . Parasitismo de Anisopteromalus calandrae Howard, 1881 (Hymenoptera: Pteromalidae) sobre formas imaturas de Lasioderma serricorne Fabricius, 1792 (Coleoptera: Anobiidae) na cidade do Rio De Janeiro, RJ . Arq. Inst. Biol., São Paulo . 73 : 489 – 491 . WorldCat Hagstrum D. W. , T. Z. Klejdysz , Bh. Subramanyam , and J. Nawrot . 2013 . Atlas of stored-product insects and mites . AACC International , St. Paul, MN . Google Preview WorldCat COPAC Hallman G. J . 2013 . Control of stored product pests by ionizing radiation . J. Stored Prod. Res . 52 : 36 – 41 . Google Scholar Crossref Search ADS WorldCat Halstead D. G . 1963 . External sex differences in stored-products Coleoptera . Bull. Entomol. Res . 54 : 119 – 134 . Google Scholar Crossref Search ADS WorldCat Hansen J. D. , J. A. Johnson , and D. A. Winter . 2011 . History and use of heat in pest control: a review . Int. J. Pest Manage . 57 : 267 – 289 . Google Scholar Crossref Search ADS WorldCat Harein P. K. , and A. F. Press . 1968 . Mortality of stored-peanut insects exposed to mixtures of atmospheric gases at various temperatures . J. Stored Prod. Res . 4 : 77 – 78 . Google Scholar Crossref Search ADS WorldCat Hartnack H . 1939 . 202 Common household pests of North America , Hartnack Publishing Co. , Chicago, IL . Google Preview WorldCat COPAC Hertlein M. B. , G. D. Thompson B. Subramanyam , and C. G. Athanassiou . 2011 . Spinosad: a new natural product for stored grain protection . J. Stored Prod. Res . 47 : 131 – 146 . Google Scholar Crossref Search ADS WorldCat Highland H. A . 1983 . Exit holes by cigarette beetles and lesser grain borers through paper bags treated with permethrin or synergized pyrethrins, J. Georgia Entomol. Soc . 18 : 65 – 71 . WorldCat Hori M . 2004a . Repellency of shiso oil components against the cigarette beetle, Lasioderma serricorne (Fabricius) (Coleoptera: Anobiidae) . Appl. Entomol. Zool . 39 : 357 – 362 . Google Scholar Crossref Search ADS WorldCat Hori M . 2004b . Repellency of hinokitiol against the cigarette beetle, Lasioderma serricorne (Fabricius) (Coleoptera: Anobiidae) . Appl. Entomol. Zool . 39 : 521 – 526 . Google Scholar Crossref Search ADS WorldCat Hori M. , M. Miwa , and H. Iizawa . 2011 . Host suitability of various stored food products for the cigarette beetle, Lasioderma serricorne (Coleoptera: Anobiidae) . Appl. Entomol. Zool . 46 : 463 – 469 . Google Scholar Crossref Search ADS WorldCat Howe R. W . 1957 . A Laboratory study of the cigarette beetle, Lasioderma serricorne (F.) (Col., Anobiidae) with a critical review of the literature on its biology . Bull. Entomol. Res . 48 : 9 – 56 . Google Scholar Crossref Search ADS WorldCat Howe R. W . 1965 . A summary of estimates of optimal and minimal conditions for population increase of some stored products insects . J. Stored Prod. Res . 1 : 177 – 184 . Google Scholar Crossref Search ADS WorldCat Howlader A. J. , and P. M. Ambadkar . 1995 . Oviposition deterring influence of female body wash in tobacco beetle, Lasioderma serricorne (F.) (Coleoptera: Anobiidae) . J. Stored Prod. Res . 31 : 91 – 95 . Google Scholar Crossref Search ADS WorldCat Hsiao T. H . 1986 . Specificity of certain chrysomelids for Solanaceae, pp. 345 – 377 . In W. G. D’Arcy (ed.), Solanaceae: biology and systematics , Columbia University Press , New York, NY . Google Preview WorldCat COPAC Hunter D. K. , and D. F. Hoffmann . 1972 . Cross infection of a granulosis virus of Cadra cautella, with observations on its ultrastructure in infected cells of Plodia interpunctella . J. Invertebr. Pathol . 20 : 4 – 10 . Google Scholar Crossref Search ADS WorldCat Imai T . 2010 . Suppression of the population of Lasioderma serricorne in stored tobacco by relocation of warehouses to cooler areas, pp. 668 – 670 . In M. O. Carvalho , P. G. Fields , C. S. Adler , F. H. Arthur , C. G. Athanassiou , J. F. Campbell , F. Fleurat-Lessard , P. W. Flinn , R. J. Hodges , A. A. Isikber , S. Navarro , R. T. Noyes , J. Riudavets , K. K. Sinha , G. R. Thorpe , B. H. Timlick , P. Trematerra and N. D. G. White (eds.), Proceedings of the 10th International Working Conference on Stored Product Protection , 27 June to 2 July 2010 , Estoril, Portugal . Julius Kühn-Institut , Berlin, Germany . Imai T. , and H. Harada . 2006 . Low-temperature as an alternative to fumigation to disinfest stored tobacco of the cigarette beetle, Lasioderma serricorne (F.) (Coleoptera: Anobiidae) . Appl. Entomol. Zool . 41 : 87 – 91 . Google Scholar Crossref Search ADS WorldCat Imai T., H. Kodama T. Chuman , and M. Kohno . 1990 . Female-produced oviposition deterrents of the cigarette beetle, Lasioderma serricorne (F.) (Coleoptera: Anobiidae) . J. Chem. Ecol . 16 : 1237 – 1247 . Google Scholar Crossref Search ADS PubMed WorldCat Imai T. , M. Onozawa , T. Takekawa , and T. Shakudo . 2006 . Lethal and sterile effects of X-ray irradiation on cigarette beetle, Lasioderma serricorne (F.) (Coleoptera: Anobiidae), Beitr Tabakforsch Int. /Contrib. Tob. Res . 22 : 1 – 5 . WorldCat Jackson R. H., and M. E. Long . 1965 . Characterization of a cell-free bacterial extract larvicidal to lasioderma serricorne (cigarette beetle) . Biochim. Biophys. Acta 100 : 418 – 425 . Jagadeesan R. , P. J. Collins , G. J. Daglish , P. R. Ebert , and D. I. Schlipalius . 2012 . Phosphine resistance in the rust red flour beetle, Tribolium castaneum (Coleoptera: Tenebrionidae): inheritance, gene interactions and fitness costs . PLoS One 7 : e31582 . Jagadeesan R., P. J. Collins M. K. Nayak D. I. Schlipalius , and P. R. Ebert . 2016 . Genetic characterization of field-evolved resistance to phosphine in the rusty grain beetle, Cryptolestes ferrugineus (Laemophloeidae: Coleoptera) . Pestic. Biochem. Physiol . 127 : 67 – 75 . Google Scholar Crossref Search ADS PubMed WorldCat Jay E. G . 1971 . Suggested conditions and procedures for using carbon dioxide to control insects in grain storage facilities . Agricultural Research Services, Miscellaneous Publication No. 51-46, USDA , Beltsville, MD . Google Preview WorldCat COPAC Jay E. G. , R. T. Arbogast , and G. C. Pearman Jr. 1971 . Relative humidity: its importance in the control of stored-product insects with modified atmospheric gas concentrations . J. Stored Prod. Res . 6 : 325 – 329 . Google Scholar Crossref Search ADS WorldCat Jones C. R . 1913 . The cigarette beetle (Lasioderma serricorne Fabr.) in the Philippine islands . J. Sci . 8 : 1 – 51 . Google Scholar Crossref Search ADS WorldCat Jurzitza G . 1969 . The vitamin requirements of normal and aposymbiotic Lasioderma serricorne F. (Coleoptera, Anobiidae) and the role of symbiotic fungi as vitamin sources for their hosts . Oecologia . 3 : 70 – 83 . [Article in German]. Google Scholar Crossref Search ADS PubMed WorldCat Kaelin P., P. Morel , and F. Gadani . 1994 . Isolation of Bacillus thuringiensis from stored tobacco and Lasioderma serricorne (F.) . Appl. Environ. Microbiol . 60 : 19 – 25 . Google Scholar PubMed WorldCat Kaelin P. , L. Zaugg , A. M. Albertini , and F. Gadani . 1999 . Activity of Bacillus thuringiensis isolates on Lasioderma serricorne (F.) (Coleoptera: Anobiidae) . J. Stored Prod. Res. 35 : 145 – 158 . COPAC Katsuki M. , K. Arikawa , M. Wakakuwa , Y. Omae , K. Okada , R. Sasaki , K. Shinoda , and T. Miyatake . 2013 . Which wavelength does the cigarette beetle, Lasioderma serricorne (Coleoptera: Anobiidae), prefer? Electrophysiological and behavioral studies using light-emitting diodes (LEDs) . Appl. Entomol. Zool . 48 : 547 – 551 . Google Scholar Crossref Search ADS WorldCat Kawakami Y. , and H. Takahashi . 2007 . Sanitary insect pests as a carrier of Aspergillus ochraceus . Mycotoxins . 57 : 47 – 56 . Google Scholar Crossref Search ADS WorldCat Kawakami Y. , I. Shimizu , and H. Takahashi . 2002 . Some fungi isolated from the cigarette beetles, Lasioderma serricorne Fabricius (Coleoptera: Anobiidae) in Japan . Med. Entomol. Zool . 3 : 249 – 256 . Google Scholar Crossref Search ADS WorldCat Kawakami Y. , I. Shimizu , H. Takahashi , and K. Okano . 2016 . Some fungi isolated from the cigarette beetles, Lasioderma serricorne in Japan − Part3 . Med. Entomol. Zool . 55 : 40 . Google Scholar Crossref Search ADS WorldCat Kearns C. W . 1934 . A hymenopterous parasite (Cephalonomia gallicola Ashm.) new to the cigarette beetle (Lasioderma serricorne Fab.) . J. Econ. Ent . 27 : 801 – 806 . Google Scholar Crossref Search ADS WorldCat Kirk H., S. Dorn , and D. Mazzi . 2013 . Molecular genetics and genomics generate new insights into invertebrate pest invasions . Evol. Appl . 6 : 842 – 856 . Google Scholar Crossref Search ADS PubMed WorldCat Kirst H. A. , K. H. Michel , I. S. Mynderse , E. H. Chao , R. C. Yao , W. M. Nakatsukasa , L. D. Boeck , J. Occlowitz , J. W. Paschel , J. B. Deeter , et al. 1992 . Discovery, isolation and structure elucidation of a family of structurally unique fermentation-derived tetracyclic macrolides, pp. 214 – 225 . In D. R. Baker , I. G. Fenyes , L. L. Steffens (eds.), Synthesis and chemistry of Agrochemicals III . America Chemical Society , Washington, DC . Google Preview WorldCat COPAC Knipling E. E . 1955 . Possibilities of insect control or eradication through the use of sexually sterile males . J. Econ. Entomol . 48 : 459 – 462 . Google Scholar Crossref Search ADS WorldCat Kohno M. , T. Chuman , K. Kato , and M. Noguchi . 1983 . The olfactory response of the cigarette beetle, Lasioderma serricorne Fabricius, to various host foods and cured tobacco extracts . Appl. Entomol. Zool . 18 : 401 – 406 . Google Scholar Crossref Search ADS WorldCat Kongratarpon T. , P. Tauthong , M. Sutantawong , and A. Wongpiyasatid . 2003 . Effects of gamma radiation on the cigarette beetle, Lasioderma serricorne (F.) . 9th Nuclear Science and Technology Conference , 19–21 June 2003 , Bangkok, Thailand . International Atomic Energy Agency (IAEA), Vienna, Austria. Korunić Z . 2010 . Effectiveness of enhanced diatomaceous earth on stored cocoa beans grains against five stored grain insect species, pp. 347 – 353 . In Zbornik Radova 22. Znanstveno Stručno Edukativni Seminar DDD i ZUPP 2010: Djelatnost dezinfekcije, dezinsekcije, deratizacije i zaštite uskladištenih , 24–26 March 2010 , Pula, Croatia . Korunić d.o.o. Zagreb, Zagreb, Croatia. Korunić Z . 2013 . Diatomaceous earths – natural insecticides . Pestic. Phytomed. (Belgrade) . 28 : 77 – 95 . Google Scholar Crossref Search ADS WorldCat Kucerová Z. , and V. Stejskal . 2010 . External egg morphology of two stored-product anobiids, Stegobium paniceum and Lasioderma serricorne (Coleoptera: Anobiidae) . J. Stored Prod. Res . 46 : 202 – 205 . Google Scholar Crossref Search ADS WorldCat Kurup A. , and D. P. Parkhe . 1962 . Some observations on the biology of the cigarette beetle (Lasioderma serricorne F.) . Indian J. Entomol . 23 : 274 – 278 . WorldCat Lacey L. A., D. Grzywacz D. I. Shapiro-Ilan R. Frutos M. Brownbridge , and M. S. Goettel . 2015 . Insect pathogens as biological control agents: back to the future . J. Invertebr. Pathol . 132 : 1 – 41 . Google Scholar Crossref Search ADS PubMed WorldCat Lawrence J. F . 1991 . Anobiidae (Bostrichoidea), pp. 441 – 444 . In F. W. Stehr (ed.), Immature insects . Kendall/Hunt Publishing Co. , Dubuque, IA . Google Preview WorldCat COPAC Lawrence J. F. , and A. F. Newton . 1995 . Families and subfamilies of Coleoptera (with selected genera, notes and references, and data on family-group names), pp. 779 – 1006 . In J. Pakaluk and S. A. Slipinski (eds.), Biology, phylogeny, and classification of Coleoptera: Papers Celebrating the 80th Birthday of Roy A. Crowson . Muzeum i Instytut Zoologii PAN , Warsaw, Poland . Google Preview WorldCat COPAC LeCato G. L. , and B. R. Flaherty . 1974 . Description of eggs of selected species of stored-product Insects (Coleoptera and Lepidoptera) . J. Kansas Entomol. Soc . 47 : 308 – 317 . WorldCat Lefkovitch L. P. , and J. E. Currie . 1963 . The effects of food shortage upon larvae of Lasioderma serricorne (F.) (Coleoptera, Anobiidae) . Bull. Entomol. Res . 54 : 535 – 547 . Google Scholar Crossref Search ADS WorldCat Lefkovitch L. P. , and J. E. Currie . 1967 . Factors affecting adult survival and fecundity in Lasioderma serricorne (F.) (Coleoptera, Anobiidae) . J. Stored Prod. Res . 3 : 199 – 212 . Google Scholar Crossref Search ADS WorldCat Leptinotarsa Sequencing Consortium . 2018 . A model species for agricultural pest genomics: the genome of the Colorado potato beetle, Leptinotarsa decemlineata (Coleoptera: Chrysomelidae) . Sci. Rep . 8 : 193 . Crossref Search ADS PubMed WorldCat Levinson H. Z. , and A. R. Levinson . 1987 . Pheromone biology of the tobacco beetle (Lasioderma serricorne F., Anobiidae) with notes on the pheromone antagonism between 4S,6S,7S- and 4S,6S,7R-serricornin . J. Appl. Entomol . 103 : 217 – 240 . Google Scholar Crossref Search ADS WorldCat Levinson H. Z. , A. R. Levinson , G. E. Kahn , and K. Schafer . 1983 . Occurrence of a pheromone-producing gland in female tobacco beetles . Experientia . 39 : 1095 – 1097 . Google Scholar Crossref Search ADS WorldCat Lim J. , M. Oh , J. Lee , and S. Lee . 2007 . Cephalonomia gallicola (Hymenoptera: Bethylidae), New to Korea, an ectoparasitoid of the cigarette beetle, Lasioderma serricorne . J. Asia-Pac Entomol . 10 : 335 – 338 . Google Scholar Crossref Search ADS WorldCat Livingstone E. M. , and W. D. Reed . 1936 . Insect fauna of cured tobacco in storage in the United States . J. Econ. Ent . 29 : 1017 – 1022 . Google Scholar Crossref Search ADS WorldCat Lü J. , and S. Liu . 2016 . The behavioral response of Lasioderma serricorne (Coleoptera: Anobiidae) to citronellal, citral, and rutin . SpringerPlus . 5 : 798 . Google Scholar Crossref Search ADS PubMed WorldCat Mahroof R. M . 2013 . Stable isotopes and elements as biological markers to determine food resource use pattern by Lasioderma serricorne (Coleoptera: Anobiidae) . J. Stored Prod. Res . 52 : 100 – 106 . Google Scholar Crossref Search ADS WorldCat Mahroof R. M. , and T. W. Phillips . 2007 . Orientation of the cigarette beetle, Lasioderma serricorne (F.) (Coleoptera: Anobiidae) to plant-derived volatiles . J. Insect Behav . 20 : 99 – 115 . Google Scholar Crossref Search ADS WorldCat Mahroof R. M. , and T. W. Phillips . 2008 . Life history parameters of Lasioderma serricorne (F.) as influenced by food sources . J. Stored Prod. Res . 44 : 219 – 226 . Google Scholar Crossref Search ADS WorldCat Mahroof R. M. , and T. W. Phillips . 2014 . Mating disruption of Lasioderma serricorne (Coleoptera: Anobiidae) in stored product habitats using the synthetic pheromone serricornin . J. Appl. Entomol . 138 : 378 – 386 . Google Scholar Crossref Search ADS WorldCat Mamta B., and M. V. Rajam . 2017 . RNAi technology: a new platform for crop pest control . Physiol. Mol. Biol. Plants . 23 : 487 – 501 . Google Scholar Crossref Search ADS PubMed WorldCat Manoto E. C. , E. B. Lapis , and A. Parungao . 1976 . Effects of gamma radiation on cigarette beetle I. Radiosensitivity of different stages of Lasioderma serricorne, Report No. PAEC (D) 76018. Philippine Atomic Energy Commission , Diliman, Quezon City Philippine . Google Preview WorldCat COPAC Marzke F. O. , J. A. Coffelt , and D. L. Silhacek . 1977 . Impairment of reproduction of the cigarette beetle, Lasioderma serricorne (Coleoptera: Anobiidae) with the insect growth regulator, methoprene . Entomol. Exp. Appl . 22 : 294 – 300 . Google Scholar Crossref Search ADS WorldCat Maxwell-Lefroy H . 1906 . India insect pests . The Superintendent of Government Printing , Calcutta, India . Google Preview WorldCat COPAC Mbata G. N., and T. W. Phillips . 2001 . Effects of temperature and exposure time on mortality of stored-product insects exposed to low pressure . J. Econ. Entomol . 94 : 1302 – 1307 . Google Scholar Crossref Search ADS PubMed WorldCat McGaughey W. H . 1975 . A granulosis virus for Indian meal moth control in stored wheat and corn . J. Econ. Entomol . 68 : 346 – 348 . Google Scholar Crossref Search ADS WorldCat McGaughey W. H . 1994 . Problems of insect resistance to Bacillus thuringiensis . Agr. Ecosyst. Environ . 49 : 95 – 102 . Google Scholar Crossref Search ADS WorldCat McGaughey W. H. , and R. W. Beeman . 1988 . Resistance to Bacillus thuringiensis in colonies of Indian meal moth and almond moth (Lepidoptera: Pyralidae) . J. Econ. Entomol . 81 : 28 – 33 . Google Scholar Crossref Search ADS WorldCat Meng L. , L. Xiao-Juan , L. Jian-Hua , and H. Ming-Fei . 2018 . The effect of acclimation on heat tolerance of Lasioderma serricorne (Fabricius) (Coleoptera: Anobiidae) . J. Therm. Biol . 71 : 153 – 157 . Google Scholar Crossref Search ADS PubMed WorldCat Miller W. R. , J. M. Munita , and C. A. Arias . 2014 . Mechanisms of antibiotic resistance in enterococci . Expert Rev. Anti Infect. Ther . 12 : 1221 – 1236 . Google Scholar Crossref Search ADS PubMed WorldCat Milne M . 1961 . The mechanism of growth retardation by nicotine in the cigarette beetle, Lasioderma serricorne . S. Afr. J. Agri. Sci . 4 : 277 – 278 . WorldCat Mitcham E., T. Martin , and S. Zhou . 2006 . The mode of action of insecticidal controlled atmospheres . Bull. Entomol. Res . 96 : 213 – 222 . Google Scholar Crossref Search ADS PubMed WorldCat Miyatake T. , T. Yokoi , T. Fuchikawa , N. Korehisa , T. Kamura , K. Nanba , S. Ryouji , N. Kamioka , M. Hironaka , M. Osada , et al. 2016 . Monitoring and detecting the cigarette beetle (Coleoptera: Anobiidae) using ultraviolet (LED) direct and reflected lights and/or pheromone traps in a laboratory and a storehouse . J. Econ. Entomol . 109 : 2551 – 2560 . Google Scholar Crossref Search ADS PubMed WorldCat Mochizuki K. , T. Chuman , M. Mori , M. Kohno , and K. Kato . 1984 . Activity of stereoisomers of serricornin, sex pheromone of the cigarette beetle (Lasioderma serricorne F.) . Agric. Biol. Chem . 48 : 2833 – 2834 . WorldCat Mokhtar A. S. , G. S. Sridhar , R. Mahmud , J. Jeffery , Y. L. Lau , J-J. Wilson , and N. M. Abdul-Aziz . 2016 . First case report of canthariasis in an infant caused by the larvae of Lasioderma serricorne (Coleoptera: Anobiidae) . J. Med. Entomol . 53 : 1234 – 1237 . Google Scholar Crossref Search ADS WorldCat Nagasawa A., Y. Kamada Y. Kosaka N. Arakida , and M. Hori . 2014 . Catechol–an oviposition stimulant for cigarette beetle in roasted coffee beans . J. Chem. Ecol . 40 : 452 – 457 . Google Scholar Crossref Search ADS PubMed WorldCat Nakagawa A. , Y. Kawakami , K. Hashimoto , Y. Hatakeyama , A. Uchida , H. Takahashi , and H. Iwano . 2008 . Adhesion rate and patterns of fungi spores on the body surface of the cigarette beetle, Lasioderma serricorne . Med. Entomol. Zool . 59 : 85 – 89 . Google Scholar Crossref Search ADS WorldCat Nasir H., and H. Noda . 2003 . Yeast-like symbiotes as a sterol source in anobiid beetles (Coleoptera, Anobiidae): possible metabolic pathways from fungal sterols to 7-dehydrocholesterol . Arch. Insect Biochem. Physiol . 52 : 175 – 182 . Google Scholar Crossref Search ADS PubMed WorldCat Navon A . 2000 . Bacillus thuringiensis insecticides in crop protection – reality and prospects . Crop Protect . 19 : 669 – 676 . Google Scholar Crossref Search ADS WorldCat Niiho C . 1984 . Ecological study of the tobacco beetle, Lasioderma serricorne (F.) II. Growth of tobacco beetles fed on bread crumbs . Jpn. J. Appl. Entomol. Zool . 28 : 209 – 216 . Google Scholar Crossref Search ADS WorldCat Noda H., and K. Kodama . 1996 . Phylogenetic position of yeastlike endosymbionts of anobiid beetles . Appl. Environ. Microbiol . 62 : 162 – 167 . Google Scholar PubMed WorldCat Okada K. , A. Watanabe , M. Mori , K. Shimazak , T. Chuman , F. Mochizuki , and T. Shibuya . 1992 . Olfactory responses to the sex pheromone component and its behavioural inhibitor in the male cigarette beetle, Lasioderma serricorne . J. Insect Physiol . 38 : 705 – 709 . Google Scholar Crossref Search ADS WorldCat Okada K. , Y. Suzaki , R. Sasaki , and M. Katsuki . 2017 . Fitness costs of polyandry to female cigarette beetle Lasioderma serricorne . Behav. Ecol. Sociobiol . 71 : 86 . Google Scholar Crossref Search ADS WorldCat Onyearu A. K. , and W. M. Graham . 1967 . Life history studies of two body colour mutants of Tribolium castaneum (Herbst) (Coleoptera, Tenebrionidae) . J. Stored Prod. Res . 3 : 65 – 70 . Google Scholar Crossref Search ADS WorldCat Palli S. R . 2014 . RNA interference in Colorado potato beetle: steps toward development of dsRNA as a commercial insecticide . Curr. Opin. Insect Sci . 6 : 1 – 8 . Google Scholar Crossref Search ADS PubMed WorldCat Pant N. C., and G. Fraenkel . 1950 . The function of the symbiotic yeasts of two insect species, Lasioderma serricorne F. and Stegobium (sitodrepa) paniceum L . Science . 112 : 498 – 500 . Google Scholar Crossref Search ADS PubMed WorldCat Papadopoulou S. Ch . 2004 . Experimental verification of Lasioderma serricorne (F.) pupa development threshold previously defined using the thermal constant equation K=D(T–t) . J. Pest. Sci . 77 : 137 – 138 . WorldCat Papadopoulou S. Ch . 2006a . Observations on the mating behavior of Lasioderma serricorne (F.) adults and experiments on their nutritional requirements in dried tobacco . Coleopt. Bull . 60 : 291 – 296 . Google Scholar Crossref Search ADS WorldCat Papadopoulou S. C . 2006b . Tyrophagus putrescentiae (Schrank) (Astigmata: Acaridae) as a new predator of Lasioderma serricorne (F.) (Coleoptera: Anobiidae) in tobacco stores in Greece . J. Stored Prod. Res . 42 : 391 – 394 . Google Scholar Crossref Search ADS WorldCat Papadopoulou S. Ch. , and C. G. Athanassiou . 2004 . Lariophagus distinguendus (F.) (Hyme., Chalcidoidea, Pteromalidae), an ectoparasitoid of Lasioderma serricorne (F.) (Col., Anobiidae), found for the first time in tobacco stores in Greece . J. Pest. Sci . 77 : 183 – 184 . WorldCat Papadopoulou S. Ch. and C. Th. Buchelos . 2002a . Identification of female adult Lasioderma serricorne (F.) by simple external observation of the abdomen . J. Stored Prod. Res . 38 : 315 – 318 . Google Scholar Crossref Search ADS WorldCat Papadopoulou S. Ch. , and C. Th. Buchelos . 2002b . Definition of the flight period of Lasioderma serricorne (F) in stored tobacco . J. Pest. Sci . 75 : 81 – 83 . WorldCat Papadopoulou S. Ch. , and C. Th. Buchelos . 2002c . Comparison of trapping efficacy for Lasioderma serricorne (F.) adults with electric, pheromone, food attractant and control-adhesive traps . J. Stored Prod. Res . 38 : 375 – 383 . Google Scholar Crossref Search ADS WorldCat Park Y. , and R. W. Beeman . 2008 . Postgenomics of Tribolium: targeting the endocrine regulation of diuresis . Entomol. Res . 38 : 93 – 100 . Google Scholar Crossref Search ADS WorldCat Parkin E. A . 1933 . The larvae of some wood-boring Anobiidae (Coleoptera) . Bull. Entomol. Res . 24 : 33 – 68 . Google Scholar Crossref Search ADS WorldCat Philips K. T . 2002 . Anobiidae Fleming 1821, pp. 245 – 260 . In R. H. Arnett Jr. M. C. Thomas P. E. Skelley , and J. H. Frank . (eds.), American beetles volume 2, Polyphaga: Scarabaeoidea through Curculionoidea . CRC Press LLC , Boca Raton, FL . Google Preview WorldCat COPAC Phoonan W. , S. Deowanish , and W. Chavasiri . 2014 . Food attractant from mulberry leaf tea and its main volatile compounds for the biocontrol of Lasioderma serricorne F. (Coleoptera: Anobiidae) . J. Stored Prod. Res . 59 : 299 – 305 . Google Scholar Crossref Search ADS WorldCat Powell T. E. Jr. 1931 . An ecological study of the tobacco beetle, Lasioderma Serricorne Fabr., with special reference to its life history and control . Ecol. Monogr . 1 : 333 – 393 . Google Scholar Crossref Search ADS WorldCat Punzo F . 1975 . Effects of temperature, moisture and thermal acclimation on the biology of Tenebrio molitor (Coleoptera: Tenebrionidae) . Ph.D. dissertation, Iowa State University , Ames, IA . Google Preview WorldCat COPAC Rafter M. A. , G. A. McCulloch , G. J. Daglish , K. Gurdasani , and G. H. Walter . 2017 . Polyandry, genetic diversity and fecundity of emigrating beetles: understanding new foci of infestation and selection . J. Pest Sci . 91 : 287 – 298 . Google Scholar Crossref Search ADS WorldCat Rajendran S. , and K. S. Narasimhan . 1994 . Phosphine resistance in the cigarette beetle Lasioderma serricorne (Coleoptera: Anobiidae) and overcoming control failures during fumigation of stored tobacco . Int. J. Pest Manage . 40 : 207 – 210 . Google Scholar Crossref Search ADS WorldCat Rao C. V. N. , B. N. Rao , and T. R. Babu . 2002 . New record of predation on the eggs of cigarette beetle, Lasioderma serricorne (Fabricius) (Coleoptera: Anobiidae), a stored tobacco pest . J. Bio. Contr . 16 : 169 – 170 . WorldCat Rayner V. I . 1951 . Some aspects of the biology of the tobacco beetle, Lasioderma serricorne (F.) (Coleoptera: Anobiidae) . M.S. thesis, University of Cape Town , Cape Town, South Africa . Google Preview WorldCat COPAC Reed W. D. , and J. P. Vinzant . 1942 . Control of insect attacking stored tobacco and tobacco products . Circular No. 635. USDA , Beltsville, MD . Google Preview WorldCat COPAC Reed W. D. , A. W. Morril Jr. , and E. M. Livingstone . 1935 . Trapping experiments for the control of the cigarette beetle . Circular No. 356. USDA , Beltsville, MD . Google Preview WorldCat COPAC Reichmuth C . 2010 . Fumigants in stored product protection . Julius-Kühn-Archiv . 429 : 56 – 64 . WorldCat Retief E. , and A. Nicholas . 1988 . The cigarette beetle Lasioderma serricorne (F.) (Coleoptera: Anobiidae): a serious herbarium pest . Bothalia . 18 : 97 – 99 . Google Scholar Crossref Search ADS WorldCat Richards O. W . 1952 . A new species of Bethylidae (Hymenoptera) from Palestine . Ann. Mag. Nat. Hist . 5 : 409 – 410 . Google Scholar Crossref Search ADS WorldCat Ridley M . 1988 . Mating frequency and fecundity in insects . Biol. Rev . 63 : 509 – 549 . Google Scholar Crossref Search ADS WorldCat Riudavets J. , I. Salas , and M. J. Pons . 2007 . Damage characteristics produced by insect pests in packaging film . J. Stored Prod. Res . 43 : 564 – 570 . Google Scholar Crossref Search ADS WorldCat Roesli R. , Bh. Subramanyam , F. J. Fairchild and K. C. Behnke . 2003 . Trap catches of stored-product insects before and after heat treatment in a pilot feed mill . J. Stored Prod. Res . 39 : 521 – 540 . Google Scholar Crossref Search ADS WorldCat Rumbos C. I. , and C. G. Athanassiou . 2012 . Insecticidal effect of six entomopathogenic nematode strains against Lasioderma serricorne (F.) (Coleoptera: Anobiidae) and Tribolium confusum Jacquelin Du Val (Coleoptera: Tenebrionidae) . J. Stored Prod. Res . 50 : 21 – 26 . Google Scholar Crossref Search ADS WorldCat Rumbos C. I. and C. G. Athanassiou . 2017a . Use of entomopathogenic fungi for the control of stored-product insects: can fungi protect durable commodities ? J. Pest. Sci . 9 : 839 – 854 . Google Scholar Crossref Search ADS WorldCat Rumbos C. I. , and C. G. Athanassiou . 2017b . The use of entomopathogenic nematodes in the control of stored product insects . J. Pest. Sci . 90 : 39 – 49 . Google Scholar Crossref Search ADS WorldCat Rumbos C. I. , M. Sakka , S. Schaffert , T. Sterz , J. W. Austin , C. Bozoglou , P. Klitsinaris , and C. G. Athanassiou . 2018 . Evaluation of Carifend®, an alpha-cypermethrin-coated polyester net, for the control of Lasioderma serricorne and Ephestia elutella in stored tobacco . J. Pest. Sci . 91 : 751 – 759 . Google Scholar Crossref Search ADS WorldCat Runner G. A . 1916 . Effect of Röntgen rays on the tobacco, or cigarette, beetle and the results of experiments with a new form of roentgen tube . J. Agr. Res . 6 : 383 – 388 . WorldCat Runner G. A . 1919 . The tobacco beetle: an important pest in tobacco product . Bulletin No. 737. USDA , Beltsville, MD . Google Preview WorldCat COPAC Ryan L . 1995 . Post-harvest tobacco infestation control . Kluwer-Academic Publishers , Dordrecht, The Netherlands . Google Preview WorldCat COPAC Saeed M. B. E. E. E. M. , M. D. Laing , R. M. Miller , and B. Bancole . 2017 . Ovicidal, larvicidal and insecticidal activity of strains of Beauveria bassiana (Balsamo) Vuillemin against the cigarette beetle, Lasioderma serricorne Fabricius (Coleoptera: Anobiidae), on rice grain . J. Stored Prod. Res . 74 : 78 – 86 . Google Scholar Crossref Search ADS WorldCat Sağlam Ö., P. A. Edde , and T. W. Phillips . 2015 . Resistance of Lasioderma serricorne (Coleoptera: Anobiidae) to Fumigation with Phosphine . J. Econ. Entomol . 108 : 2489 – 2495 . Google Scholar Crossref Search ADS PubMed WorldCat Saleh A . 2012 . Ecological and biological study of the cigarette beetle, Lasioderma serricorne and survey its natural enemies . M.S. thesis, Damascus University, Damascus, Syria . Google Preview WorldCat COPAC Salem H. M. , M. A. Fouda , A. A. Abas , W. M. Ali , and A. Gabarty . 2014 . Effects of gamma irradiation on the development and reproduction of the greasy cutworm, Agrotis ipsilon (Hufn.) . J. Radiat. Res. Appl. Sci . 7 : 110 – 115 . Google Scholar Crossref Search ADS WorldCat Salgado V. L . 1998 . Studies on the mode of action of spinosad: insect symptoms and physiological correlates . Pestic. Biochem. Phys . 60 : 91 – 102 . Google Scholar Crossref Search ADS WorldCat Schlipalius D. I., Q. Cheng P. E. Reilly P. J. Collins , and P. R. Ebert . 2002 . Genetic linkage analysis of the lesser grain borer Rhyzopertha dominica identifies two loci that confer high-level resistance to the fumigant phosphine . Genetics . 161 : 773 – 782 . Google Scholar PubMed WorldCat Self L. S., F. E. Guthrie , and E. Hodgson . 1964a . Metabolism of nicotine by tobacco-feeding insects . Nature . 204 : 300 – 301 . Google Scholar Crossref Search ADS WorldCat Self L. S. , F. E. Guthrie , and E. Hodgson . 1964b . Adaptation of tobacco hornworms to the ingestion of nicotine . J. Insect Physiol . 10 : 907 – 914 . Google Scholar Crossref Search ADS WorldCat Shah A. A. , and A. A. Khan . 2014 . Use of diatomaceous earth for the management of stored-product pests . Int. J. Pest Manage . 60 : 100 – 113 . Google Scholar Crossref Search ADS WorldCat Shen S. K. , and P. F. Dowd . 1991 . Detoxification spectrum of the cigarette beetle symbiont Symbiotaphrina kochii in culture . Entomol. Exp. Appl . 60 : 51 – 59 . Google Scholar Crossref Search ADS WorldCat Shinoda K. , and K. Fujisaki . 2001 . Effect of adult feeding on longevity and fecundity of the cigarette beetle, Lasioderma serricorne F. (Coleoptera: Anobiidae) . Appl. Entomol. Zool . 36 : 219 – 223 . Google Scholar Crossref Search ADS WorldCat da Silva A. P. O. , H. F. Goulart , A. E. G. Santana , J. R. Martins , A. Riffel , and J. C. Vaz . 2018 . Pest management in stored products: the case of the cigarette beetle, Lasioderma serricorne (Coleoptera: Anobiidae), pp. 61 – 89 . In E. Lichtfouse (ed.), Sustainable agriculture reviews 27 . Springer International Publishing AG , Basel. Switzerland . Google Preview WorldCat COPAC Sivik F. P. , J. N. Tenhet , and C. D. Delamar . 1957 . An ecological study of the cigarette beetle in tobacco storage warehouses . J. Econ. Entomol . 50 : 310 – 316 . Google Scholar Crossref Search ADS WorldCat Stanley D. C. , and H. J. Ball . 1962 . Mode of action and insecticidal value of a diatomaceous earth as a grain protectant . J. Econ. Entomol . 55 : 964 – 970 . Google Scholar Crossref Search ADS WorldCat Steidle J. L. M . 1998 . The biology of Lariophagus distinguendus: a natural enemy of stored product pests and potential candidate for biocontrol . Bull. OILB-SROP 21 : 103 – 109 . WorldCat Stephen J. F . 1832 . Illustration of British entomology or a synopsis of indigenous insects: containing their generic and specific distinction; with an account of their metamorphoses, times of appearance, localities, food, and economy, as far as practicable. Mandibulata Vol. V . Baldwin and Cradock , London, UK . Google Preview WorldCat COPAC Subramanyam B . 2009 . Controlling cigarette beetles . Milling J . 2nd Quarter : 34 – 35 . WorldCat Sun X. , L. F. Wang , Y. Feng , H. Xie , X. Y. Zheng , A. He , M. R. Karim , Z. Y. Lv , and Z. D. Wu . 2016 . A case report: a rare case of infant gastrointestinal canthariasis caused by larvae of Lasioderma serricorne (Fabricius, 1792) (Coleoptera: Anobiidae) . Infect. Dis. Poverty . 5 : 34 . Google Scholar Crossref Search ADS PubMed WorldCat Suneethamma K . 2016 . Biology and morphometrics of cigarette beetle, Lasioderma serricorne (Fabricius) on fennel and coriander and its management using modified atmosphere with CO2 and temperature . M.S. thesis, Sri Venkateswara Agricultural College, Acharya N. G. Ranga Agricultural University , Andhra Pradesh, India . Google Preview WorldCat COPAC Swingle M. C . 1938 . Low temperature as a possible means of controlling the cigarette beetle in stored tobacco . Circular No. 462. USDA , Beltsville, MD . Google Preview WorldCat COPAC Tabashnik B. E., T. Brévault , and Y. Carrière . 2013 . Insect resistance to Bt crops: lessons from the first billion acres . Nat. Biotechnol . 31 : 510 – 521 . Google Scholar Crossref Search ADS PubMed WorldCat Tamang S . 2015 . Biology and management of cigarette beetle, Lasioderma serricorne (F.) on turmeric powder . M.S. thesis, Anand Agricultural University , Gujarat, India . Google Preview WorldCat COPAC Tenhet J. N . 1961 . Controlling the cigarette beetle in the tropics. Agricultural Marketing Service , Market Quality Research Division Publication No. AMS 439. USDA , Beltsville, MD . Google Preview WorldCat COPAC Tenhet J. N. , and C. O. Bare . 1951 . Control of insects in stored and manufactured tobacco . Circular No. 869. USDA , Beltsville, MD . Google Preview WorldCat COPAC Thompson J. V. , and L. W. Fletcher . 1972 . A pathogenic strain of Bacillus cereus isolated from the cigarette Beetle, Lasioderma serricorne . J. Invertebr. Pathol . 20 : 341 – 350 . Google Scholar Crossref Search ADS WorldCat Throne J. E . 2010 . Overview of North American stored product research . Julius-Kühn-Archiv . 425 : 42 – 49 . doi: 10.5073/jka.2010.425.099. WorldCat Tilton E. W. , and J. H. Brower . 1987 . Ionizing radiation for insect control in grain and grain products . Cereal Foods World . 32 : 330 – 335 . WorldCat Tilton E. W., W. E. Burkholder , and R. R. Cogburn . 1966 . Effects of gamma radiation on Rhyzopertha dominica, Sitophilus oryzae, Tribolium confusum, and Lasioderma serricorne . J. Econ. Entomol . 59 : 1363 – 1368 . Google Scholar Crossref Search ADS PubMed WorldCat Tobin E. N. , and L. W. Smith Jr. 1971 . Note on the mating behavior of the cigarette beetle (Lasioderma serricorne (F): Anobiidae . Entomol. News . 82 : 23 – 25 . WorldCat Trematerra P. C. Athanassiou V. Stejskal A. Sciarretta N. Kavallieratos , and N. Palyvos . 2011 . Large scale mating disruption of Ephestia spp. and Plodia interpunctella in Czech Republic, Greece and Italy . J. Appl. Entomol . 135 : 749 – 762 . Google Scholar Crossref Search ADS WorldCat Tribolium Sequencing Consortium . 2008 . The genome of the model beetle and pest Tribolium castaneum . Nature . 52 : 949 – 955 . WorldCat Tsuchiya S., Y. Kasaishi H. Harada T. Ichimatsu H. Saitoh E. Mizuki , and M. Ohba . 2002 . Assessment of the efficacy of Japanese Bacillus thuringiensis isolates against the cigarette beetle, Lasioderma serricorne (Coleoptera: Anobiidae) . J. Invertebr. Pathol . 81 : 122 – 126 . Google Scholar Crossref Search ADS PubMed WorldCat USDA (U.S. Department of Agriculture) . 1971 . Stored tobacco insects: biology and control . Agricultural Handbook No. 233. USDA , Beltsville, MD . Google Preview WorldCat COPAC Vinzant J. P. , and W. D. Reed . 1941 . Type of wire screen required for excluding cigarette beetles and tobacco moths from warehouses . J. Econ. Entomol . 34 : 724 . Google Scholar Crossref Search ADS WorldCat Wakil W. , M. U. Ghazanfar , and M. Yasin . 2014 . Naturally occurring entomopathogenic fungi Infecting stored grain insect species in Punjab, Pakistan . J. Insect Sci . 14 : 182 . Google Scholar Crossref Search ADS PubMed WorldCat White R. E . 1971 . Key to North American genera of Anobiidae, with phylogenetic and synonymic notes (Coleoptera) . Ann. Entomol. Soc. Am . 64 : 179 – 191 . Google Scholar Crossref Search ADS WorldCat Wijayaratne L. K. W . 2011 . Effects of methoprene on Tribolium castaneum (Herbst) (Coleoptera: Tenebrionidae) . Ph.D. dissertation, University of Manitoba , Winnipeg, Manitoba, Canada . Google Preview WorldCat COPAC Worden B. D. , and P. G. Parker . 2001 . Polyandry in grain beetles, Tenebrio molitor, leads to greater reproductive success: material or genetic benefits ?. Behav. Ecol . 12 : 761 – 767 . Google Scholar Crossref Search ADS WorldCat Wright V. F. , E. de las Casas , and P. K. Harein . 1980 . Evaluation of Penicillium mycotoxins for activity in stored product Coleoptera . Environ. Entomol . 9 : 217 – 221 . Google Scholar Crossref Search ADS WorldCat Yamamoto R. T. , and G. Fraenkel . 1960 . The suitability of tobaccos for the growth of the cigarette beetle, Lasioderma serricorne. J. Econ. Entomol . 53 : 381 – 384 . Google Scholar Crossref Search ADS WorldCat Yaman M. , I. Aslan , A. Görmez , and Ö. Ertürk . 2008 . Bacterial flora of Lasioderma serricorne (F.) (Coleoptera: Anobiidae) from several tobacco stores in Turkey . Integrated Protection of Stored Products, IOBC/WPRS Bull . 40 : 49 – 52 . WorldCat Yang C. , Z. Jin , L. Guangtao , Q. Guiqiang , Z. Sixu , and L. Tao . 2008 . Respiration of Tribolium castaneum (Herbst) at different oxygen concentrations, pp. 15 – 20 . In G. Daolin , S. Navarro , Y. Jian , T. Cheng , J. Zuxun , L. Yue , L. Yang , and W. Haipeng , (eds.), Proceedings of the 8th International Conference on Controlled Atmosphere and Fumigation in Stored Products , 21–26 September 2008 , Chengdu, China . Sichuan Publishing House of Science & Technology , China . Yang W-J. , K-K. Xu , X-Y. Zhu , C-X. Chen , X-Y. Cai , Y. Cao , Y-L. Meng , H. Yang and C. Li . 2017 . The complete mitochondrial genome of the cigarette beetle, Lasioderma serricorne (Coleoptera: Anobiidae) . Mitochondrial DNA Part B . 2 : 430 – 431 . Google Scholar Crossref Search ADS WorldCat Yezerski A., G. Cussatt D. Glick , and M. Evancho . 2005 . The effects of the presence of stored product pests on the microfauna of a flour community . J. Appl. Microbiol . 98 : 507 – 515 . Google Scholar Crossref Search ADS PubMed WorldCat Yu C . 2008 . Susceptibility of Lasioderma serricorne (F.) life stages exposed to elevated temperatures. M.S. thesis, Kansas State University , Manhattan, KS . Google Preview WorldCat COPAC Yu C., B. Subramanyam P. W. Flinn , and J. A. Gwirtz . 2011 . Susceptibility of Lasioderma serricorne (Coleoptera: Anobiidae) life stages to elevated temperatures used during structural heat treatments . J. Econ. Entomol . 104 : 317 – 324 . Google Scholar Crossref Search ADS PubMed WorldCat Zettler L. , and D. W. Keever . 1994 . Phosphine resistance in cigarette beetle (Coleoptera: Anobiidae) associated with tobacco storage in the southeastern United States . J. Econ. Entomol . 87 : 546 – 550 . Google Scholar Crossref Search ADS WorldCat Zhou S., R. S. Criddle , and E. J. Mitcham . 2000 . Metabolic response of Platynota stultana pupae to controlled atmospheres and its relation to insect mortality response . J. Insect Physiol . 46 : 1375 – 1385 . Google Scholar Crossref Search ADS PubMed WorldCat © The Author(s) 2019. Published by Oxford University Press on behalf of Entomological Society of America. All rights reserved. For permissions, please e-mail: journals.permissions@oup.com. This article is published and distributed under the terms of the Oxford University Press, Standard Journals Publication Model (https://academic.oup.com/journals/pages/open_access/funder_policies/chorus/standard_publication_model)
TI - Biology, Ecology, and Control of Lasioderma serricorne (F.) (Coleoptera: Anobiidae): A Review
JF - Journal of Economic Entomology
DO - 10.1093/jee/toy428
DA - 2019-05-22
UR - https://www.deepdyve.com/lp/oxford-university-press/biology-ecology-and-control-of-lasioderma-serricorne-f-coleoptera-jKRYT7PWzj
SP - 1011
VL - 112
IS - 3
DP - DeepDyve
ER -