Stored-Product Insect Resource by Scientific Societies

Contents Chapter 1: Species List . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Table 1.1. Insects and their natural enemies associated with stored products . . . . . . . . . . . . Table 1.2. Full addresses for website keys and website pictures . . . . . . . . . . . . . . . . . . . . Table 1.3. Archaeological records of stored-product insects . . . . . . . . . . . . . . . . . . . . . . Table 1.4. Geographical origins of stored-product insect species . . . . . . . . . . . . . . . . . . . Table 1.5. Ranking of stored-product insect species by number of commodities with which they have been reported . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Table 1.6. Additional insect species reported to be associated with stored products or facilities that have not been included in Table 1.1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Chapter 2: Species by Family . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Table 2.1. Stored-product insect species in each family, listed by family (number of species) (order) and species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Table 2.2. Ranking of families by number of species in each. . . . . . . . . . . . . . . . . . . . . Table 2.3. Ranking of genera with more than one species by number of species in each . . . .

. . . . .

. 1 . 4 200 201 204

. . . . . . . . . . . . . . 205 . . . . . . . . . . . . . . 206

. . . . . . . . . . . . . . . 211 . . . . . . . . . . . . . . . 211 . . . . . . . . . . . . . . . 228 . . . . . . . . . . . . . . . 229

Chapter 3: Species by Key . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233 Table 3.1. Stored-product insect species in Table 1.1 included in each key . . . . . . . . . . . . . . . . . . . . . . . . . . . 233 Chapter 4: Species by Commodity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Table 4.1. Stored-product insect species and their natural enemies reported in literature as associated with a commodity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Table 4.2. Ranking of commodities by number of stored-product insect species associated with each commodity . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Table 4.3. Studies on development, fecundity and population growth of stored-product insects on various commodities . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Table 4.4. Ranking of species by number of commodities in Table 4.3 tested for suitability . . . Table 4.5. Summary of reported numbers of stored-product insect species associations with various types of facilities or habitats . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .

. . . . . . . . . . . . . . 279 . . . . . . . . . . . . . . 279 . . . . . . . . . . . . . . 346 . . . . . . . . . . . . . . 347 . . . . . . . . . . . . . . 363 . . . . . . . . . . . . . . 364

Chapter 5: Species by Rank . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 367 Table 5.1. Ranking for each stored-product insect species . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 367 Chapter 6: Species by Common Name . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Table 6.1. Scientific names of stored-product insects and their natural enemies listed by their common names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 377 Chapter 7: Commodities by Scientific or Common Name . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 Table 7.1. Commodities in Table 1.1 listed by scientific name. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 383 Table 7.2. Commodities in Table 1.1 listed by common name . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 390 Chapter 8: References Cited . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 395 Chapter 9: Preparing Insects for Identification . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 481 Chapter 10: Books and Book Chapters . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 485 Chapter 11: Alphabetical List of Species Names . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 489 Table 11.1. Alphabetical list of current species names in Table 1.1 giving genus name for each. . . . . . . . . . . . . . . 489 Table 11.2. Alphabetical list of synonym species names in Table 1.1 giving the genus for each . . . . . . . . . . . . . . . 505

vii

Preface The initial purpose of our book was to compile a list of the insect species reported to be associated with stored products (any commodity of plant or animal origin other than fresh fruit or vegetable and lumber that is stored for later use) during storage, transportation, processing and marketing. In the process of compiling the list from more than 2,200 publications, we summarized information that will help anyone interested in stored-product entomology identify insects and find relevant information on these species. We hope the biological information from the literature cited in this book will be the basis for developing insect pest management programs. This book supplements the textbook Fundamentals of Stored-Product Entomology by Hagstrum and Subramanyam (2006). Many of the stored-product insects have one or more synonyms. Without knowledge of the 508 synonyms discovered while compiling the list in Table 1.1, some information on many species may be overlooked. The keys available for identifying stored-product insects are catalogued so that the best key can be selected. The more than 9,200 published records of insect association with various commodities and more than 1,800 published records of their association with facilities are summarized for the first time providing insight into the range of commodities and facilities infested by a particular insect species. Much of the information on insect biology may not have been utilized fully because quickly finding the information needed to solve a problem has been difficult. The biggest problem is finding a literature citation. Once the reader has a literature citation, many sources are available to provide a copy of the paper, via libraries, interlibrary loans and commercial online services such as Bioone (http://www.bioone.org/perlserv/?request=searchsimple), Ingentaconnect (http://www.ingentaconnect.com/), JSTOR (http://www.jstor.org/stable/3822224), Science Direct (http://www.sciencedirect.com/), Springerlink (http://www. springerlink.com/journals/) and many others. Chapter 1 is an alphabetical list by genus of the 1,663 insect species that have been reported in the literature as associated with stored products during storage, transportation, processing and marketing. Feeding by some insect pest species can actually reduce the amount of commodity that is available for sale or consumption. Other insect species are scavengers or mold feeders that can contaminate stored products and when found frequently enough or in sufficient numbers can be economically important in making a commodity unsuitable for sale or consumption. Chapter 1 provides extensive information on the commodities or facilities infested, the geographical distributions of each insect species, the ranking of their importance as pest or beneficial insects, common names, synonyms, available keys and

pictures for these insects, natural enemies of pests, hosts/ prey of beneficial insects and sources of biological information to characterize the pest or natural enemy status of an insect species. For a more common stored-product insect species, up to 243 commodities and 39 species of natural enemies are given. Literature citations are provided for up to 21 keys per species and 57 biological studies per species. Literature citations or web addresses with pictures are provided for many insect species. However, many insect species cannot be adequately identified to species using only pictures and selecting the best key is therefore important for correct insect identification. The range of commodities infested by each insect species and records of their geographical distribution are provided to help the reader narrow the number of stored-product insect species that might be expected. The book can also tell the reader where to find information on the biology of a stored-product insect species that will be useful in developing a pest management program. Chapter 1 is the longest chapter and Chapters 2 through 7 reorganize the information given in Chapter 1. Chapter 2 provides an alphabetical list of the families showing the numbers of species of stored-product insects in each of 120 families. There are 259 genera that have more than one species, 22 genera that have 10 or more species and two that have more than 25 species. This is important because the species in the same genus often are most similar in appearance making them most difficult to distinguish using only pictures. Chapter 3 provides an alphabetical list of the storedproduct insect species separated by each key. This will help the reader find the best key for identifying an insect. With ever increasing international commerce, we are more likely to have problems with the less common stored-product insect species. In order to prevent the introduction of these less common insect species into a region or country, effective quarantine and pest management programs may require that we can identify all insect species associated with stored products. Chapter 4 is an alphabetical list of commodities showing the insect species reported to be associated with each commodity. This chapter provides an overview of the number of insect species associated with each commodity. For example, 218 insect species can infest stored wheat, 256 species can infest stored maize, and 505 species can infest at least one of the stored cereal grains. Chapter 5 is a ranked list of stored-product insect pests and their natural enemies showing their relative economic importance. In summarizing the rankings from three published studies, it was clear that the ranking was very dependent upon the commodity infested. We extend this v

vi / Preface

analysis using information from Chapter 1 on number of keys, number of commodities infested, number of natural enemies, number of hosts/prey attacked and number of biological studies for each stored-product insect species. The book provides an international overview of the relative importance of stored-product insect species. Chapter 6 is an alphabetical list of the common names of stored-product insects giving the scientific name for each species. This list is useful in finding the scientific name for a species that is used in Chapter 1 when only the common name is known. Chapter 7 is an alphabetical list of the scientific names of the commodities giving the common names for each. This list is useful in finding the common name for a commodity that is used in Chapter 1 when only the scientific name is known. Scientific names are also listed by common names. Chapter 8 has the references cited in Chapters 1 through 7 of the book. Chapter 9 discusses the methods of collect-

ing and preserving insects for identification and Chapter 10 provides an overview of additional books and review articles that are available on stored-product insects. Chapter 11 is an alphabetical list of species providing an index to the generic names for each species. This can be important because insect species are listed alphabetically by their genus name in previous chapters. This book summarizes in one place an extensive database on a surprisingly large number of insect species associated with stored products during storage, transportation, processing and marketing. We sincerely hope that the pest management practitioners use the Stored-Product Insect Resource to improve the quality and reduce the cost of their pest management programs. David W. Hagstrum Bhadriraju Subramanyam

April 6, 2009

Chapter 1: Species List Insects associated with stored products and their natural enemies are in the orders Coleoptera (Col.), Diptera (Dip.), Hemiptera (Hem.), Homoptera (Hom.), Hymenoptera (Hym.), Lepidoptera (Lep.) and Psocoptera (Pso.) (Table 1.1). Hinton (1945) reported that there were 600 species of Coleoptera associated with stored products. Halstead (1986) revised this to 500 species and Delobel and Tran (1993) included 416 species of Coleoptera. In the book edited by Prakash (2003), the authors reported 700 insect species associated with stored products during storage, transportation, processing and marketing. Hagstrum and Subramanyam (2006) included only 101 of the most common storedproduct insect species in their textbook so these publications and the current book provided a much broader view of the insect species associated with stored products. Insect scavengers and mold feeders may be found contaminating stored products. These scavengers and mold feeders and insect species feeding directly on stored products or commodities that are found frequently or in sufficient numbers are considered pests. Pests of fresh fruit or vegetable and lumber have been included in Table 1.1 only when they also have been reported as problems in dried fruit, root crop, finished wood items or commodities other than fresh fruit or vegetable and lumber. Mathew (1987) lists 68 insect species that infest timber stored in India. Records in the literature in which insects were only identified to genus have not been included in Table 1.1. Although Acarina (mites) are also included in many of the publications used to compile Table 1.1, species of Acarina have not been included. By 2009, more than 1,660 insect species have been reported to be associated with stored products during storage, processing, transportation and marketing. The number of insect species in Table 1.1 is larger than the number included in previous estimates. This is mainly a result of including more commodities and larger geographical areas, but some new stored-product insect species have been recently described. A total of 160 insect species associated with stored products have been described since 1950 including 67 species of natural enemies. Studies show that the number of insect species known to be associated with stored products generally has been expanded each time a commodity or a geographical area has been examined more closely. General categories for the types of stored-product insects include: (1) insects that are primarily found with stored commodities and the facilities or habitats with which these commodities are associated (S), (2) insects that infest commodities in the field prior to harvest and may continue to reproduce during storage (F), (3) insects that are primar-

ily field pests and generally do not reproduce during storage but can occur in sufficient numbers to require pest management during commodity storage (C) and (4) parasitoids or predators that are natural enemies of stored-product insects (P). The majority of these natural enemies belong to the orders Hemiptera and Hymenoptera. However, some species of Coleoptera and Diptera in the families Carabidae, Cecidomyiidae, Cleridae, Colydiidae, Eumolpidae, Histeridae, Passandridae, Proctotrupidae, Scenopinidae, Staphylinidae, Stratiomyidae and Tenebrionidae are also predators. Insect species were only marked as parasitoids or predators in Table 1.1 when they have been observed and reported in the literature to be natural enemies. With further research, many more of these species may be found to be predators. Some studies such as those by Donohoe (1939), Hunter et al. (1973), Redlinger and Davis (1982), and Salmond (1956) do not indicate that an insect species infests a particular commodity but only indicate the type of facilities or habitat (T) in which a species has been found. The â&#x20AC;&#x153;Type, Rankâ&#x20AC;? heading in Table 1.1 includes the category for the type of insect species, and the rank (R) and the new rank (NR) from Chapter 5. Some of the species categorized in this book as S may actually be in the categories C or F, but this was not specified in the publications consulted in compiling the list in Table 1.1. Also, some of the species categorized as T may actually be shown to be in the categories S, C or F as more records of these species become available. The list in Table 1.1 has been compiled from large surveys, but also from smaller surveys, from many reports for individual species and from research on insect populations during storage, transportation, processing and marketing of commodities. A book edited by Prakash et al. (2003) provided information on the stored-product insect species found in different geographical regions of the world. Books and papers by Aitken (1975, 1984), Howe and Freeman (1955) and Hurlock (1964) included the stored-product insects found in commodities imported into England. Papers by Haines (1974, 1981) included the stored-product insects mainly from tropical countries submitted for identification. Extensive surveys of stored-product insects have been published by Attia and Kamel (1966) for Egypt, Bousquet (1990, 1991) for Canada, Cornes (1973) for Nigeria, Davies (1960) for Uganda, Delobel (1992) for cassava in Central Africa, Forsyth (1966) for Ghana, Haines and Pranata (1982) for Java, Hayhurst (1942) for England, Loschiavo and Okumura (1979) for Hawaii, Obr (1978) for psocids in Czech Republic and Sauphanor et al. (1987) for yams in Ivory Coast. Several surveys have been done in China (Chao et al. 1980, Chen 1985, Chen and Huang 1

2 / Chapter 1

1985, Guilin et al. 1999, Yan et al. 2006) and in Greece (Buchelos 1981, 1985a,b; Buchelos and Athanassiou 1993a, 1998). The introduction and establishment of stored-product insects have been summarized for several countries including Australia and neighboring pacific countries (Archibald and Chalmers 1983, Archibald and Maddison 1988), Belgium (Casteels et al. 1995), Bulgaria (Tomov et al. 2007), Canada (Majka 2007, 2008; Majka and Cline 2006; Majka and Smetana 2007), Czech Republic (Sefrova and Lastuvke 2005, Stejskal and Kucerova 1993), Germany (Wohlgemuth and Reichmuth 1983), Hungary (Merkl 2006), India (Bhalla et al. 2004; Gupta et al. 2002, 2004, 2005; Kapur and Verma 2004; Wadhi 1963), Italy (Aldini 2004, Contessi 1994, Ratti and Rampini 1977), Japan (Kiritani 1959, Mito and Uesugi 2004), Netherlands (Schulten and Roorda 1984), Poland (Ruta et al. 2004) and United States (Olsen 1981, Zimmerman 1990). The insects infesting bean (Hansson et al. 2004), dried fish (Osuji 1974b, 1980), dried mushroom (Madenjian et al. 1993), kola nut (Daramola 1976a), lentil (Ho and Zainy 1985), maize (Ayertey and Ibitoye 1987, Compton et al. 1998), sorghum (Ayertey and Ibitoye 1987), and yam and yam flour (Osuji 1980, Sika et al. 2004) for sale in African, Asian, Mexican and North American markets have been surveyed. The lists of hosts/prey for some natural enemies are quite long. Many Hymenoptera are both predators and parasitoids, killing many stored-product insects on which they feed but do not lay eggs. Natural enemies do better on some hosts than on others. Laelius pedatus is more than twice as effective against Trogoderma variabile as against Trogoderma glabrum (Klein and Beckage 1990). Conversely, the ability of parasitoids and predators to attack many additional species of stored-product insects may not have been fully observed and reported. Pteromalus cerealellae was once thought to be a parasitoid of only Sitotroga cerealella and is now known to parasitize 13 host species (Brower 1991). The ability of this parasitoid to reduce host population varied from 0% for Acanthoscelides obtectus to 97% for Callosobruchus maculatus. Some insect pest species may be incorrectly identified as hosts for natural enemies because stored-product insects are often used as factitious (artifical/surrogate) host for natural enemies of crop and forest insect pests. The long lists of natural enemies that attack many species of stored-product insects suggest that there are many unexplored opportunities for biological control. Foreign exploration for new and more effective species and strains of natural enemies may have greater than expected potential in developing biological control programs. Also, some of the natural enemies of crop and storage pests may be bred to be more effective against storage pests. The majority (57.0%) of the insect species associated with stored products were named between 1800 and 1899. Fewer stored-product insect species were named earlier than 1800 (15.1%) or later than 1899 (27.9%). There have been many changes over the years in the names used for stored-product insects. For example, the family Bruchidae

has had the names Mylabridae after the genus name Mylabris (Geoffroy 1762), Lariidae after the genus name Laria (Scopoli 1763) and Bruchidae after the genus name Bruchus (Linnaeus 1767) (Arora et al. 1969). These three genus names are synonyms (syn.). Linnaeus described the genus Bruchus under the family Curculionidae in 1758 and Lacordaire removed the family Bruchidae from the Curculionidae in 1845. Mylabridae and Lariidae family names are no longer used because genotypes were not designated for the genera Mylabris and Laria. Recently, the bruchids have been considered a subfamily, Bruchinae, in the family Chrysomelidae (Lingafelter and Pakaluk 1997). Borowski and Wegrzynowicz (2007) included Lyctidae as a subfamily of the Bostrichidae, and Lobl and Smetana (2007) include Anobiidae as a subfamily of the Ptinidae. The classification of stored-product insects is continually changing and the reader may need to find more recent taxonomic revisions for the current name. The 508 synonyms encountered in the non-taxonomic literature that was used to complile Table 1.1 are included. Knowing that two or more names have been used to refer to the same insect species can be important in correctly reporting on a species, locating information on their biology or accurately estimating the number of insect species associated with a commodity. Literature citations are included in Table 1.1 for the biological studies encountered while compiling the list of insects associated with stored products. Taxonomic literature has been consulted to identify synonyms and to describe the geographical distribution of a species. Table 1.1 summarizes the geographical distribution information found in the publications cited in Chapter 8. However, additional observations for most species may be needed to describe their full geographical distribution and the included records may best be considered only examples. Records for the geographical distributions of insect species associated with stored products are arranged roughly by continents into 7 regions, namely, Africa (Af), Asia (As), Central America (C), Europe (E), North America (N), Oceania (O) and South America (S). Islands (Is.) are generally included with the closest continent. Asian region includes the Middle East and many of the islands in the Indian Ocean. Central America includes islands in the Carribean. North America includes Canada, Mexico and United States of America (USA). Distributions of insect species among states within the USA and provinces within Canada are reported much more frequently than distributions within other countries and these are included in parentheses. When insect species have been reported to be introduced, the continent or country name is proceeded by the letter â&#x20AC;&#x153;iâ&#x20AC;?. When reported, the year that species is first discovered is given in parentheses after the continent, country, province or state name. When importation data were used to compile Table 1.1, only the exporting country is included unless there is evidence that the species has become established in the importing country. Identifying new synonyms can expand the geographic distribution of a species and discovering misidentifications in an insect collection can reduce the geographical distri-

Species List / 3

bution of a species. Also, the geographical distributions of many insect species are still being expanded through international trade. Central heating in residences and food-processing plants has allowed more of the tropical insect species to become established in temperate climates (Halstead 1975b, Howe and Lefkovitch 1957, Katz 1972). Changes in harvesting methods, in pest management practices and in transportation have resulted in some species becoming more important and others becoming less important (Conway 1987, Halstead 1975b, Howe and Lefkovitch 1957, Katz 1975, Yoshida 1984). Populations of insect species like Trogoderma granarium have become temporarily established in some countries but have been eradicated through effective pest management programs or local populations have naturally become extinct (Armitage 1956, 1958; Banks 1977; Hagstrum and Subramanyam 2006; Shephard 1957; Sonda 1968; Viljoen 1990; Whelan 1955). Literature and websites at which pictures of storedproduct insects can be found are cited in Table 1.1. Table 1.2 includes the full addresses for these websites (Web1Web13 in Table 1.1). In many cases, pictures may be more quickly accessed on the internet when the reader does not have access to a university library with entomology books and journals. In a few cases, pictures are available for adults (A) and immatures (E (egg), L (larva), P (pupa), N (nymph)) or male (M) and female (F) adults. Only the websites with pictures of many stored-product insect species are cited here and websites with pictures of other storedproduct insect species can be found by searching the internet. Also, the insect pictures in the books by Bousquet (1990) and Kingsolver (2004) are available at http://home.cc.umanitoba.ca/~fieldspg/fields/beetles.pdf and http://www.ars.usda.gov/is/np/Bruchidae/BruchidaeIntro.htm, respectively. Because many stored-product insects are similar in appearance, keys should be consulted for identifying insects rather than relying on pictures alone. References are cited in Table 1.1 for the keys that are available in the literature to identify many of the insect species listed. Table 3.1 lists the insect species covered by each key. This table can help the reader select the best key. However, each of the keys cited includes only a few of the species in Table 1.1. This leaves the user uncertain as to whether they have one of the insect species that is not in that key. At some point, consulting the taxonomic descriptions of several insect species or a taxonomist may be necessary. Chapter 9 provides the information needed to prepare insects for identification by yourself or a taxonomist. Adult insects can be identified first to family by using a key in a general entomology textbook. Many of the references cited for species have keys to many of the genera in a family. To simplify the keys, often keys are developed for the genera within a subfamily or family and then a separate key is available for the species within each genus. In addition to the keys cited in Table 1.1, keys to genera also are available for the families Anobiidae (White 1971a), Anthicidae (larva, Kitayama 1982), Anthribidae (Valentine 1998), Braconidae (Baltazar 1962, Marsh et al. 1987), Bruchidae (Borowiec 1987a), Ciidae (Lawrence 1971),

Cleridae (Barr 1962, Solervicens 2001), Dermestidae (Beal 2003, Hava 2004, Rees 1943), Erotylidae (larva, Rymer Roberts 1958), Eurytomidae (Burks 1971), Ichneumonidae (Townes 1969a,b,c; Townes 1971; Townes and Townes 1966, 1973; Townes et al. 1961, 1965), Laemophloeidae (Thomas 1988b), Mycetophagidae (Parsons 1975), Passandridae (Thomas 1988b), Pteromalidae (Boucek and Rasplus 1991), Pyrrhocoridae (Robertson 2004), Silvanidae (Thomas 1988b), Staphylinidae (Moore and Legner 1974) and Tenebrionidae (Doyen 1989). Yoshimoto (1984) and Grissell and Schauff (1997) have keys to the subfamilies and families, and Gibson et al. (1997) to the families and genera of the Chalcidoidea Hymenoptera which include Chalcididae, Encyrtidae, Eulophidae, Eurytomidae, Leucospidae, Mymaridae, Perilampidae, Pteromalidae, Torymidae and Trichogrammatidae. Crowson (1956) and Halstead (1986) have keys to the Coleoptera families of stored-product insects, New (1977) and New and Lienhard (2007) have keys to Asian families and genera of Psocoptera, Oldroyd (1954) has a key to Diptera families in England, Dolling (1991) has keys to the English families and genera of Hemipteran adults and nymphs, Priesner and Alfieri (1953) has a key to Egyptian families of Hemiptera, Richards (1956) has a key to Hymenoptera families in England and Slater and Baranowski (1978) have keys to the families and genera of Hemiptera. Some insect identification keys are available online (Table 1.2). Keys to insect orders by Choate (Key1), by Queensland Museum (Key2), adapted from Oldroyd (1958) (Key3) and adapted from Borror and White (1970) (Key4) are available online. Keys by Choate are available online for the families of Coleoptera (Key5) and for the families of Hemiptera (Key6). A key by Grissell and Schauff (1997) is available for the families and subfamilies of the Chalcidoidea Hymenoptera (Key7). A key to well over 100 species of Coleoptera and Lepidoptera associated with stored products by Weidner and Rack (1984) is available in French (Key8) and in English (Key9). These websites also include keys to the families of the most common Hymenopteran natural enemies. Keys that include stored-product insect species are available online for the pyralid larvae by Solis (1999) (Key10), for tenebrionid adults of Florida and eastern United States by Dunford, Thomas and Choate (Key11) and for laemophloeid, passandrid and silvanid adults by Thomas (1993) (Key12Key15). Also, keys to 110 species of Coleoptera in the book by Bousquet (1990) and 17 species of Bruchidae in the book by Kingsolver (2004) are available at http:// home.cc.umanitoba.ca/~fieldspg/fields/beetles.pdf and http:// www.ars.usda.gov/is/np/Bruchidae/BruchidaeIntro.htm, respectively. Lawrence and co-workers describe characteristics for quick identification of Coleoptera families at http:// www.inbio.ac.cr/ES/papers/coleoptera/MELYRIIN.html. Judy Johnson’s “Stored product insects: identification and biology” at http://www.cardinalproproducts.com/Misc/Stored ProductInsecthandoutcolor.pdf provides a very simplified guide to the identification of the most common storedproduct insect species. Additional keys are available in journal articles that are available online (see preface for online access to journal papers).

4 / Chapter 1

Table 1.1. Insects and their natural enemies associated with stored productsa Acanthocnemis nigricans (Hope 1845) Type, Rank: T Taxonomy: Col., Melyridae, also family Acantocnemidae Distribution (4): iAf: Algeria; iAs; iE: Cyprus, France, Italy; O: Australia Commodities (2): wheat, wooden walking stick Facilities: grain elevators Acantholyctus cornifrons (Lesne 1898) Type, Rank: S Taxonomy: Col., Lyctidae Pictures: Borowski and Wegrzynowicz 2007, Web1 Distribution (2): Af: Algeria, Eritrea, Ethiopia, Morocco, Mozambique, Namibia, Senegal, Somalia, Tunisia; As: Iran, Israel, Jordan, Saudi Arabia Commodities (2): cotton seed, wood walking stick Natural enemies (1): Cephalonomia mateui Acanthoscelides argillaceus (Sharp 1885) Type, Rank: C, 2, NR=1 Common names: lima bean weevil Taxonomy: Col., Bruchidae, syn. Acanthoscelides obreptus Bridwell 1942 Keys (2): Johnson 1983, 1990 Pictures: Johnson 1990 Distribution (3): C: Guadeloupe, Guatemala, Haiti, Honduras, Panama, Trinidad; N: Mexico; S: Brazil, Chile, Colombia, Peru Commodities (5): cowpea, hyacinth bean, kidney bean, lima bean, pigeonpea Natural enemies (4): Chryseida bennetti, Eupelmus cushmani, Stenocorse bruchivora, Torymus atheatus Acanthoscelides chilensis (Schilsky1905) Type, Rank: C, 2, NR=1 Taxonomy: Col., Bruchidae Keys (1): Johnson 1983 Pictures: Johnson 1983 Distribution (1): S: Argentina, Chile, Uruguay Commodities (1): bean Acanthoscelides clandestinus (Motschulsky 1874) Type, Rank: C Taxonomy: Col., Bruchidae Distribution (1): S Commodities (1): cowpea Acanthoscelides collusus (see Acanthoscelides pallidipennis) Acanthoscelides dominicanus (see Mimosestes mimosae) Acanthoscelides griseolus (Fall 1910) Type, Rank: C Taxonomy: Col., Bruchidae, syn. Bruchus griseolus Fall 1910 a

See text for explanation of headings and abbreviations. Addresses for web sites Web1-Web13 are given in Table 1.2.

Keys (3): Johnson 1983, 1990; Kingsolver 2004 Pictures: Johnson 1983, 1990; Kingsolver 2004 Distribution (3): As: Japan; C: Costa Rica; N: Mexico, USA (AZ, CA, TX) Commodities (1): sesbania seed Acanthoscelides macrophthalmus (Schaeffer 1907) Type, Rank: C, NR=1 Taxonomy: Col., Bruchidae Keys (3): Johnson 1983, Kingsolver 2004, Pfaffenberger and Johnson 1976 (1st instar larva) Pictures: Johnson 1983, Kingsolver 2004, Web2 Distribution (3): As: iJapan (2000), C: El Salvador, Guatemala, Nicaragua; N: Mexico, USA (TX) Commodities (1): jumbie bean Natural enemies (5): Eupelmus amicus, Heterospilus prosopidis, Horismenus missouriensis, Lariophagus texanus, Stenocorsa bruchivora Acanthoscelides morosus (see Sennius morosus) Acanthoscelides obreptus (see Acanthoscelides argillaceus) Acanthoscelides obtectus (Say 1831) Type, Rank: F, 2, 4, NR=5 Common names: bean weevil Taxonomy: Col., Bruchidae Keys (14): Anton et al. 1997; Bousquet 1990; Carvalho 1979; Haines 1991; Herford 1935; Johnson 1983, 1990; Kingsolver 1991e, 2004; McFarlane and Wearing 1967 (pupa); Pfaffenberger and Johnson 1976 (1st instar larva); Pratt and Scott 1962; Shomar 1963; Weidner and Rack 1984 Pictures: Borowiec 1987a; Bousquet 1990; Carvalho 1979; Delobel and Tran 1993; Gorham 1991; Haines 1991; Hill 2002 (A,L,P); Johnson 1983, 1990; Kingsolver 2004; Linsley and Michelbacher 1942 (A,E,L,P); Patton 1931; Shomar 1963; VanRyckeghem 2004; Web1; Web5; Wightman and Southgate 1982 (E) Distribution (7): Cosmopolitan. Af: Burundi, Egypt, Ethiopia, Ghana, Kenya, Lesotho, Libya, Malawi, Nigeria, Rwanda, Swaziland, Tanzania, Uganda, Zambia; As: India, Iran, Israel, iJapan, Lebanon, Syria, Turkey; C: Bahamas, Guatemala, Honduras, Nicaragua, Puerto Rico; E: Albania, Armenia, Austria, iBulgaria (1955), Germany, Greece, Ireland, Netherlands, Poland, Portugal, Romania, Russia, Spain, Switzerland, Yugoslavia; N: Canada (BC, MB, ON, PQ), Mexico, USA (AZ, CA (1769), HI, RI (1860), NY (1891)); O: Australia, New Caledonia, New Zealand, Papua New Guinea, Tonga; S: Argentina, Chile, Peru, Venezuela. The bean weevil originated in South America and has been found with red lima beans in ancient Indian graves in Peru dating from 1-500 A.D. The bean weevils were introduced in California by Mission Padres with the founding of the first Mission at San Diego. Bean weevil was not reported in France before the late 19th century, a good 300 years after bean crops were introduced. From southeastern France, bean weevil colonized a large part of Europe and was later

Species List / 5

spread to Africa following the introduction of Phaseolus vulgaris as crop. Commodities (18): bambara groundnut, bean, broad bean, chickpea, cowpea, kidney bean, lentil, lima bean, lupine seed, pea, pigeonpea, pulse, sorghum, soybean, tepary bean, vetch, wattle seed, white bean. Black gram, chickling vetch, hyacinth bean, maize, moth bean, mung bean, rice bean, scarlet runner bean, sesbania and yardlong bean are also suitable host plants. Facilities: tobacco curing barns, tobacco fields Natural enemies (18): Anisopteromalus calandrae, Bracon vestiticida, Chryseida bennetti, Dinarmus basalis, Eupelmus cushmani, Eupelmus cyaniceps, Eupelmus vesicularis, Eurytoma bruchophaga, Heterospilus prosopidis, Horismenus depressus, Pteromalus cerealellae, Stenocorse bruchivora, Theocolax elegans, Torymus atheatus, Triaspis thoracica, Trichogramma evanescens, Uscana semifumipennis, Uscana senex Biology (30): Alvarez et al. 2005; Arbogast 1991b; Biemont and Bonet 1981; Bonet et al. 1987; Caswell 1962; Essig 1929; Hodek et al. 1981; Horler 1970; Howe and Currie 1964; Jarry 1981; Labeyrie 1981; Larson 1932; Leroi 1981; Leroi and Jarry 1981; Menten and Menten 1984; Menten et al. 1981; Menusan 1934, 1935, 1936; Perez and Bonet 1984; Perttunen 1972; Perttunen and Haryrinen 1969, 1970a,b; Pfaffenberger 1985; Schoonhoven 1977; Shade et al. 1987; Stamopoulos 1989; Thiery and Jarry 1985; Weiss 1944 Acanthoscelides obvelatus Bridwell 1942 Type, Rank: C, 2, NR=1 Taxonomy: Col., Bruchidae Keys (2): Johnson 1983, 1990 Pictures: Johnson 1983, 1990 Distribution (3): C: Guatemala, Puerto Rico; N: Mexico; S: Colombia Commodities (2): bean, kidney bean. Scarlet runner also is a suitable host. Natural enemies (4): Chryseida bennetti, Eupelmus cushmani, Stenocorse bruchivora, Torymus atheatus Biology (1): Alvarez et al. 2005 Acanthoscelides pallidipennis (Motschulsky 1874) Type, Rank: C Taxonomy: Col., Bruchidae, syn. Acanthoscelides collusus Fall 1910 Pictures: Rogers and Garrison 1975, Tuda et al. 2001 Distribution (3): iAs: China, Japan, Korea; iE: England, Hungary; N: USA Commodities (1): false indigo seed (planted for erosion control) Natural enemies (1): Horismenus productus Biology (1): Tuda et al. 2001 Acanthoscelides puellus (Sharp 1885) Type, Rank: C, 2, NR=2 Taxonomy: Col., Bruchidae Keys (2): Johnson 1983, 1990 Pictures: Johnson 1983, 1990

Distribution (2): C: El Salvador, Honduras, Panama; N: Mexico Commodities (1): bean Natural enemies (6): Eupelmus amicus, Heterospilus bruchi, Heterospilus prosopidis, Horismenus missouriensis, Stenocorse bruchivora, Zatropis incertus Acanthoscelides surrubresus (see Meibomeus surrubresus) Acanthoscelides zeteki Kingsolver 1969 Type, Rank: C, 2, NR=1 Taxonomy: Col., Bruchidae Keys (2): Johnson 1983, 1990 Pictures: Johnson 1983, 1990 Distribution (3): C: Bahamas, Guatemala, Panama, Puerto Rico, Trinidad, West Indies; N: Mexico; S: Curacao, Venezuela Commodities (2): cowpea, pigeonpea Natural enemies (3): Eupelmus amicus, Horismenus missouriensis, Stenocorse bruchivora Acedes fuscipunctella (see Niditinea fuscella) Achaea catocaloides GuenĂŠe Type, Rank: S Taxonomy: Lep., Noctuidae Distribution (1): Af Commodities (1): palm kernel Achaea lienardi (Boisduval 1833) Type, Rank: S Taxonomy: Lep., Noctuidae Distribution (1): Af Commodities (1): cocoa bean Achroia grisella (Fabricius 1794) Type, Rank: S, NR=2 Common names: honey moth, lesser bee moth, lesser wax moth Taxonomy: Lep., Pyralidae Keys (7): Arbogast et al. 1980 (egg), Carvalho 1979, Corbet and Tams 1943a, Ferguson 1991, Hinton 1943b (larva), Weisman 1991 (larva), Zimmerman 1978a (larva) Pictures: Arbogast et al. 1980 (E); Carvalho 1979; Corbet and Tams 1943a, 1943b (M,F); Goater 1986; Gorham 1991; Web1; Web2; Web4; Web9; Web12; Zimmerman 1958 Distribution (6): Probably of European origin. Af: Egypt, Libya, Nigeria, South Africa; As: Hong Kong, India; C: Jamaica; E: England, Germany, Guernsey, Ireland, Isle of Scilly, Jersey, Scotland, Wales; N: USA (iHI); O: New Zealand Commodities (10): apple (dried), beeswax, blanket, currant, dried fruit, fig (dried), honeycomb, insect (dried), peach (dried), tobacco seed Natural enemies (4): Antrocephalus galleriae, Apanteles galleriae, Dibrachys boarmiae, Venturia canescens (lab) Biology (4): Carter 1984, Greenfield and Coffelt 1983, Shylesha and Rajagopal 1995, Uckan et al. 2007

6 / Chapter 1

Achryson surinamensis Linnaeus 1767 Type, Rank: S Taxonomy: Col., Cerambycidae Pictures: Web4 Distribution (2): N: USA (southern); S: Argentina, Peru Commodities (1): lonchocarpus root. Does not breed during storage.

Distribution (4): Af: Ivory Coast, Madagascar, Mauritius, Reunion Is., Tanzania; As: India, Japan, Java; C: Grenada, St. Vincent, Tobago, Trinidad; O: Fiji Commodities (4): ginger root, potato, turmeric root, yam Hosts/Prey (1): Aspidiotus hartii

Acritus analis LeConte 1853 Type, Rank: S Taxonomy: Col., Histeridae Distribution (2): C: Cuba, Guatemala, Puerto Rico; N: USA (CA) Commodities (1): barley

Adistemia watsoni (Wollaston 1871) Type, Rank: S, NR=1 Taxonomy: Col., Lathridiidae, syn. Lathridius watsoni Wollaston Keys (6): Hinton 1941a, 1945, 1945 (larva), 1945 (pupa); Kingsolver and Andrews 1991; Mound 1989 Pictures: Gorham 1991, Halstead 1986, Hinton 1945 Distribution (5): Af: Algeria, Canary Is., Egypt, Madeira, Morocco; As: Israel, Japan; E: Belgium, Czech Republic, Denmark, England, Finland, France, Germany, Ireland, Italy, Netherlands, Norway, Portugal, Spain, Sweden, Switzerland; iN: Canada (ON, PQ), S Commodities (3): drug, hay, tamarind product Facilities: granaries, herbariums Biology (2): Hinton 1945, Kerr and McLean 1956

Acrobasis caryae Grote 1881 Type, Rank: C Common names: pecan nut case-bearer Taxonomy: Lep., Pyralidae Keys (2): Corbet and Tams 1943a, Hinton 1943b (larva) Pictures: Corbet and Tams 1943a Distribution (1): N: Canada (ON), USA (AR, IL, MI, SC) Commodities (1): pecan Natural enemies (4): Eupelmus amicus, Goniozus columbianus, Macrocentrus instabilis, Trichogramma minutum Biology (1): Gecan et al. 1971 Acrobasis nuxvorella Neunzig 1970 Type, Rank: C Common names: pecan nut case-bearer Taxonomy: Lep., Pyralidae Keys (1): Weisman 1991 (larva) Pictures: Web4 Distribution (1): N: USA (AL, FL, GA, IL, LA, MO, MS, NC, NM, OK, SC, TX) Commodities (1): pecan Natural enemies (3): Eupelmus amicus, Goniozus columbianus, Macrocentrus instabilis Acrobasis vaccinii Riley 1884 Type, Rank: C Common names: cranberry fruitworm Taxonomy: Lep., Pyralidae Keys (1): Weisman 1991 (larva) Pictures: Web4, Web12 Distribution (1): N: Canada (NS), USA (FL, TX, WI) Natural enemies (3): Bracon mellitor, Trichogramma minutum, Trichogramma pretiosum Biology (1): Murray et al. 1996

Adistemia rileyi (see Aridius rileyi)

Adontomerus amygdali Boucek 1958 Type, Rank: P Taxonomy: Hym., Torymidae Distribution (1): As: Jordan Hosts/Prey (1): Eurytoma amygdali Biology (1): Doganlar et al. 2006 Aethus indicus (Westwood 1837) Type, Rank: S Taxonomy: Hem., Cydnidae Distribution (3): Af: Sierra Leone; As; O: Australia Commodities (1): cocoa bean Africobatus harpaloides (GuĂŠrin-MĂŠneville 1847) Type, Rank: S Taxonomy: Col., Carabidae Distribution (1): Af: Malawi, Nigeria Commodities (1): tobacco Agathis cylasovorus (see Bassus cylasovorus)

Adelencyrtus femoralis (see Adelencyrtus moderatus)

Agathis gibbosa (Say 1936) Type, Rank: P Taxonomy: Hym., Braconidae Distribution (2): C: Bermuda; N: Canada (AB, ON, PQ), USA (CA, CO, GA, IL, IN, KS, LA, MA, MD, MI, MO, NJ, NM, NY, OR, TX, VA, WA) Hosts/Prey (1): Phthorimaea operculella Biology (1): Odebiyi and Oatman 1972

Adelencyrtus moderatus Howard 1897 Type, Rank: P Taxonomy: Hym., Encyrtidae, syn. Adelencyrtus femoralis Compere and Annecke 1961

Agathis unicolor (Schrottky 1902) Type, Rank: P Taxonomy: Hym., Braconidae, syn. Agathis unicolorata Shenefelt 1970

Species List / 7

Distribution (3): As: Borneo, Sarwawak; iN; S: Argentina Hosts/Prey (1): Phthorimaea operculella Biology (2): Chundurwar 1978, Odebiyi and Oatman 1977 Agathis unicolorata (see Agathis unicolor) Aglenus brunneus (Gyllenhal 1813) Type, Rank: P Taxonomy: Col., Colydiidae Keys (3): Carvalho 1979, Halstead 1986, Hinton 1945 Pictures: Carvalho 1979, Delobel and Tran 1993, Halstead 1986, Hinton 1945 Distribution (5): Af: Algeria, Canary Is., Egypt, Morocco; As: Turkestan; E: Azores, Caucasus, England, Germany, Greenland, Iceland, Scotland; N: USA (CA, MO); S: Chile Commodities (4): animal feed, bone, grain, grain residue Facilities: cellars, commercial mushroom beds, cowsheds, granaries, haystacks, manure heaps, mills, primitive tanneries, stables Hosts/Prey (1): Xylodromus concinnus Biology (2): Kenward 1975, Woodroffe 1967 Aglossa caprealis (H端bner 1800-1809) Type, Rank: S, NR=2 Common names: murky meal caterpillar, murky meal moth, small stable tabby moth, small tabby moth Taxonomy: Lep., Pyralidae Keys (6): Carvalho 1979, Corbet and Tams 1943a, Ferguson 1991, Hinton 1943b (larva), Solis 1999 (larva), Weisman 1991 (larva) Pictures: Arbogast and Byrd 1981 (E); Carvalho 1979; Corbet and Tams 1943a,b (M,F); Goater 1986; Web2; Web7; Web9; Web12 Distribution (6): Cosmopolitan. Af: Algeria, Nigeria; As: China, Myanmar; E: England, Italy; N: USA (OH, southern); O: Australia, New Zealand; S Commodities (13): bean, cereal, cocoa bean residue, cork in wine bottle, cracker, linen, maize, mushroom (dried), pabulum, rice, tobacco (nearly rotten), wheat, wheat flour Facilities: feedmills, flour mills, grain elevators, peanut shelling plants, peanut warehouses Biology (3): Carter 1984, Trematerra 1987a, Watt 1965a Aglossa dimidiata (Haworth 1809) Type, Rank: S, 2, NR=1 Common names: black rice worm, tea tabby Taxonomy: Lep., Pyralidae Keys (1): Corbet and Tams 1943a Pictures: Corbet and Tams 1943a, 1943b (M,F), Goater 1986 Distribution (2): As: China, India, iJapan, Korea, Taiwan; E: England Commodities (7): Chinese traditional medicine, herbal medicine, herbarium specimen, insect (dried), rice, tea (dried), wheat Facilities: tea warehouses Natural enemies (1): Peregrinator biannulipes Biology (1): Trematerra 1987a

Aglossa ocellalis Lederer 1863 Type, Rank: S, 3, NR=1 Taxonomy: Lep., Pyralidae, includes three subspecies, Aglossa ocellalis indistincta Corbet and Tams (Baluchistan, northwest India), Aglossa ocellalis pinguinalis Linnaeus (Cameroon, Europe, North America) and Aglossa ocellalis asiatica Erscoff (Egypt to Iran, Turkestan) Pictures: Goater 1986 Distribution (4): Af: Cameroon, Egypt, Ivory Coast, Nigeria, Sierra Leone; As: Baluchistan, India, Iran, Pakistan, Turkestan; E: England; N Commodities (7): ginger, hoof, horn, maize, oilseed, palm kernel, yam Aglossa pinguinalis (Linnaeus 1758) Type, Rank: S Common names: grease moth, Italian moth of fat and lard, large tabby moth Taxonomy: Lep., Pyralidae, three subspecies: Aglossa pinguinalis asiatica Erscoff (Cyprus, Egypt, Iran, Jordan, Palestine, Turkestan, Yemen); Aglossa pinguinalis indistincta Corbet and Tams (Baluchistan, northwest India); Aglossa pinguinalis pinguinalis Linnaeus (Cameroon, Europe, North America) Keys (2): Corbet and Tams 1943a, Hinton 1943b (larva) Pictures: Corbet and Tams 1943a (M,F), 1943b (M,F); Goater 1986; Web5; Web9; Web12 Distribution (4): Eurasian origin. Af: Cameroon, Egypt, Libya, Morocco; As: Afganistan, China, India, Iran, Iraq, Pakistan, Turkestan, Yemen; E: England, Greece, Italy, Russia, Sweden; O: Australia, Tasmania Commodities (9): barley, book binding, butter, cereal, cereal residue, dung (dried), lard, seed (dried), vegetable material (dried). Natural enemies (1): Stomatoceras pomonellae Biology (2): Carter 1984, Trematerra 1987a Agonoderus lecontei (see Stenolophus lecontei) Agonopterix applana (see Agonopterix heracliana) Agonopterix heracliana (Linnaeus 1758) Type, Rank: T Taxonomy: Lep., Oecophoridae, syn. Agonopterix applana Fabricius 1777, Depressaria applana (Fabricius 1777) Distribution (1): E: England, Scotland Facilities: grain warehouses Agonum mulleri (Herbst 1784) Type, Rank: T Taxonomy: Col., Carabidae Keys (2): Lindroth 1974, Luff and Turner 2007 Pictures: Luff and Turner 2007 Distribution (3): As: Siberia, Turkey; E: Albania, Austria, Azores, Belgium, Bosnia Herzegovina, Bulgaria, Croatia, Czech Republic, Denmark, England, Estonia, Finland, France, Germany, Hungary, Ireland, Italy, Latvia, Liechtenstein, Lithuania, Malta, Moldavia, Netherlands, Norway, Poland, Portugal, Russia, Scotland, Serbia and Montenegro,

8 / Chapter 1

Slovakia, Slovenia, Spain, Sweden, Switzerland, Ukraine; iN: Canada (BC, NB, NF, iNS (1897), PE, ON, PQ) Facilities: mills, warehouses Ahasverus advena (Waltl 1834) Type, Rank: F, 1 or 2 (cassava root flour, copra, maize, peanut, sesame seed), 3 (cocoa bean, ginger, palm kernel, rice, sheanut, sorghum, tobacco, yam), NR=4 Common names: foreign grain beetle Taxonomy: Col., Silvanidae Keys (16): Bousquet 1990; Carvalho 1979; Cutler 1971 (larva); Delobel and Tran 1993; Grouvelle 1912; Haines 1991; Hinton and Corbet 1955; Kingsolver 1991a; Klein 1986; Kucerova and Stejskal 2002 (egg); Mound 1989; Pratt and Scott 1962; Rees 2004; Thomas 1993; Weidner and Rack 1984, 1984 (larva) Pictures: Anon. 1986; Bousquet 1990; Carvalho 1979; Delobel and Tran 1993; Gorham 1991; Haines 1991; Halstead 1986, 1993; Hayhurst 1942; Hill 2002; Klein 1986; Kucerova and Stejskal 2002 (E); Mound 1989; Patton 1931; Rees 1996, 2004; VanRyckeghem 2004; Web2; Web5; Web6; Web13 (A,L); Weidner and Rack 1984 (A,L) Distribution (7): American, Ethiopian or Oriental origin. Af: Algeria, Burundi, Cameroon, Canary Is., Congo, Egypt, Ethiopia, Ghana, Lesotho, Liberia, Libya, Madeira, Malawi, Morocco, Mozambique, Nigeria, Rwanda, South Africa, Uganda; As: Bangladesh, China, India, Indonesia, Iran, Israel, iJapan, Malaysia, Pakistan, Philippines, Siberia, Singapore, Sri Lanka, Taiwan, Thailand, Vietnam; C: Cuba, Dominican Republic, Grenada, Honduras, Jamaica, Panama, Puerto Rico, St. Lucia, Trinidad and Tobago; E: Albania, Austria, Belarus, Belgium, Bosnia Herzegovina, Croatia, Czech Republic, Denmark, England, Finland, France, Germany, Greece, Hungary, Italy, Liechtenstein, Lithuania, Poland, Portugal, Romania, Russia, Slovakia, Slovenia, Spain, iSweden, Switzerland; iN: Canada (AB, BC, MB, ON, PQ, SK), Mexico, USA (AL, AR, AZ, CA, FL, GA, HI, IA, IN, KS, KY, MA, MD, MI, NC, NE, NJ, NH, NY, OH, OK, OR, SC, TX, VT, WA, WI); O: Australia, Micronesia, New Caledonia, New Zealand, Papua New Guinea, Samoa, Solomon Is., Tonga; S: Argentina, Brazil, Chile, Colombia, Ecuador, Paraguay, Suriname Commodities (120): algarroba pod, almond hull, animal (dried), animal feed, apple pomace, bamboo leaf (dried), banana (dried), barley, basketware, BBQ sauce, bean, bean curd, biscuit, black pepper, broom, broom corn, bulb, cake (stale), capsicum pod, cashew, cassava root, cassava root flour, cassava root meal, castor seed, cereal, chili bean sauce, chili paste, chili pepper, chili pod (dried), Chinese traditional medicine, chocolate, clothing, cocoa, cocoa bean, cocoa cake, coffee, coffee bean, copra, coriander, cotton seed, dandelion, date, dried fruit, fennel seed, fig, fig (dried), fish (preserved), flax, fungus (dried), ginger, ginger root, grain, grain (decaying), grain product, grass seed, groundnut cake, hemp (marijuana), herb, herb (dried), horsebean, illipenut, leaf (dried), licorice apricot, lily

flower (dried), lima bean, linseed, litchi nut, lonchocarpus root, maize, maize bran, maize cob, malt, meal, melon rind, millet, mixed grain residue, mushroom, mussel (dried), nut, nutmeg, oat, oilcake, oilseed, olive, onion, orange pomace, paddy, palm kernel, peanut, pigeonpea, pine nut, plum, prune, raisin, rice, root (dried), safflower seed, sago flour, sesame seed, sheanut, shrimp (dried), sorghum, soup mix (dried), soybean, soybean residue, spice, sunflower seed, tamarind, tapioca shred, taro, tobacco, tomato (dried), turmeric, turtle shell, vegetable (preserved), wheat, wheat flour, yam, yellow bean sauce. Does not damage clean, dry grain. Facilities: barley mills, cocoa storages, currant raisin storages, empty cargo containers, farm grain bins, farm storages of rice, feedmills, flat grain storages, flour mills, grain elevators, peanut shelling plants, peanut warehouses, pet stores, railroad cars, semolina mills, sultana raisin storages Biology (13): Arbogast 1991b; Campbell 1989; Cox et al. 2007; David and Mills 1975; David et al. 1974; Hagstrum 2000, 2001; Hill 1965, 1978; Jacob 1996; Pierce et al. 1991; Toews et al. 2006; Woodroffe 1962 Ahasverus excisus (Reitter 1876) Type, Rank: S Taxonomy: Col., Silvanidae, syn. Cathartus excisus Reitter 1876 Keys (1): Grouvelle 1912 Distribution (5): As; C: Guatemala, Panama, Trinidad, West Indies; iE: Germany, Italy, Netherlands; N: Mexico; S: Brazil, Suriname, Venezuela Commodities (1): tobacco Ahasverus rectus (LeConte 1854) Type, Rank: T Taxonomy: Col., Silvanidae Keys (2): Grouvelle 1912, Thomas 1993 Distribution (2): C: Cuba, Puerto Rico; N: USA (AL, AZ, FL, GA, MO, NC, TX) Facilities: botanical warehouses, retail stores Akis elongata Brulle 1832 Type, Rank: S Taxonomy: Col., Tenebrionidae Distribution (1): E: Greece Commodities (5): barley, hay, maize, wheat, wheat flour Facilities: flour mills Akis reflexa Fabricius 1775 Type, Rank: S Taxonomy: Col., Tenebrionidae Distribution (1): Af: Egypt Commodities (5): barley, hay, maize, wheat, wheat flour Alaptus globosicornis Girault 1908 Type, Rank: P Taxonomy: Hym., Mymaridae Keys (2): Girault 1908, 1929