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Pectinophora gossypiella (Saunders) -- Lepidoptera, Gelechiidae




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       When P. gossypiella invaded the lower Colorado Desert of Arizona and California in 1978, within 10 years the following damages resulted:  (1) total cotton area dropped from 79,942 ha. to 37,130 ha., (2) total value of cotton dropped from $245,812,807 to $71,803,418, (3) insect control costs increased from $375.59/ha. to $639.98/ha., and (4) whitefly populations increased exponentially with pyrethroid insecticide use (Johnson et al. 1982) producing severe honeydew, yield loss and disease symptoms in cotton and surrounding vegetables (tomatoes, cucurbits, sugar beets and lettuce).


       Pectinophora is a numerically small genus but based on its distribution and species richness, the center of endemicity appears to lie along the coastal rim of Australia extending from southeast Queensland to northern Western Australia.  From Australia pink bollworm probably taken by humans to Indonesia and then further distributed to parts of Africa, India, Mexico and the United States. 




       Pectinophora may have originated in Australia or in neighboring islands to the northwest of the continent on a malvaceous plant other than domesticated cotton.  Generally ignored in all discussions of pink bollworm movement is the positional relationship of Indonesia.  This region seems geographically pivotal and important in understanding pink bollworm movement.  Indonesia has not been an important cotton producing region and therefore it has been largely ignored in an analysis of pink bollworm problems.  Nevertheless, movement of pink bollworm into the Indonesian Archipelago from Northern Australia seems obvious.  The earliest record of pink bollworm on Java dates from 1903 (Kalshover 1981).  More recently, pink bollworm was observed on Bali, Lombok, Flores and Timor; this would suggest that it is widespread in Indonesia (G. Gordh, unpub. data).


       The shift of pink bollworm from Australia to Indonesia seems probable in that:  (1) the movement could have been affected by trade, probably by indigenous peoples (the Bugis or Makassar of Sulawesi) before Dutch colonization or (2) pink bollworm naturally invaded the islands of Indonesia at an earlier time in geological history (Miocene-Pleistocene).  Once on the islands of Indonesia, pink bollworm was sustained on some of the numerous species of malvaceous plants in that region.  Two host plants commonly used by pink bollworm, Hibiscus tiliaceus and Thespesia populnea, are widespread in Indonesia.


       In this hypothetical scenario, subsequent trade between India and the Indonesian Archipelago, probably during the Hindu expansion, moved pink bollworm into India.  In India, pink bollworm remained at insignificant levels on endemic alternative host plants, such as Abutilon indicum, Hibiscus spp., Thespesia populnea, etc.  During this period of residence in India, pink bollworm may have been transported to Africa by traders.


       When American cotton was imported into India, pink bollworm quickly shifted hosts.  Movement to Egypt probably occurred from India via transport of seed.  Movement elsewhere within Africa probably occurred from the spread of pink bollworm in Tanzania northward into Kenya and thence along the coast into Somalia.  Movement to Brazil and Mexico came through transport of infested seed originating in Egypt.  Movement to Malaysia, China and Hawaii came through transport of infested seed originating in India.  Pink bollworm has been described from India in 1843 and subsequently recovered in Hawaii (1901), East Africa (1904), Malaysia and Burma (1906), Egypt (1907), Australia (1911), Mexico (1911), Brazil (1913), Texas (1917), China (1918) and California (1965).


Spread in Egypt and the United States


       The history of pink bollworm in Egypt is tied to the production of cotton.  During historical times cotton was woven into fabric before it was cultivated in Egypt (Ballou 1919).  Cotton production probably began in the 13th or 14th century.  Production was increased and the industry stimulated about 1820.  Then cotton in Upper Egypt was produced as a perennial crop; cotton in Lower Egypt was produced as an annual crop.  The first cotton seeds used for commercial plantings in Egypt were obtained from ornamental gardens near Cairo.  This so-called Jumel cotton was cultivated for many years as a three-year perennial.  Subsequently Sea Island cotton was imported from Georgia and Florida in the United States, and Peruvian cotton was imported from brazil.  During the latter half of the 19th century Ashmuni and Mit Affifi were the common varieties produced.


       Pink bollworm was first noted in spinning mills near Alexandria in 1906-07 (Pearson 1958).  Willcocks (1916) first detected pink bollworm in field plots during November-December 1910.  The moth may have been present in Egypt as early as 1903.  Accounts of pink bollworm in Egypt around 1879 have not been confirmed.  That is the year in which the cotton worm, Prodenia litura, was discovered in Egypt.  Records of pink bollworm in Egypt around 1879 may refer to cotton worm.  Alternative host plants for pink bollworm in Egypt include Bamia, Hibiscus esculentus, teel, Hibiscus cannabinus, and hollyhock, Althea rosea (Willcocks 1916).  The source of the infestation has been traced to badly ginned cotton received from India (Willcocks 1916).  It was noted that Caravonica cotton seed had been introduced into Egypt from Queensland, but no pink bollworms were detected.  By 1912 pink bollworm was the most common pest of cotton in Lower Egypt.


       Pink bollworm was first intercepted in the United States at Hearne, Texas on September 10, 1917 by Mr. Ivan Schiller (Hunter 1918).  Infestations in Texas developed from Mexican cotton seed taken to Texas for oil extraction.  Subsequent infestations may have resulted from bales of ginned cotton swept from the wharves of Galveston during hurricanes of August 1915 (Hunter 1918).  Presumably the cotton bales washed ashore near cultivated cotton.


       Pink bollworm was first detected on wild cotton in southern Florida during 1932, but origin of the infestation cannot be documented.  The pest was detected on cultivated cotton in northern Florida and southern georgia at about the same time, and persisted until eradicated during 1936.  Pink bollworm persists at very low levels on wild cotton in the Florida Keys (G. Gordh pers. commun.).


       Pink bollworm was first reported in California during 1965 (Legner & Medved 1979).  Problems with this insect in the southwestern United States since 1965 have centered in New Mexico, Arizona and the California lower desert.  More substantial acreage of cotton growing in the Central Valley of California have experienced sporadic infestations which are controlled by releases of sterile pink bollworm males.




       During the past 80 years, 160 species in 43 genera of parasitic Hymenoptera have been collected with or reared from pink bollworm infested cotton (Gordh 1989).  The genera Apanteles, Bracon, Brachymeria, Chelonus and Elasmus among the parasitic Hymenoptera contribute numerous species of potentially useful pink bollworm parasitoids.  Efforts to permanently establish 14 imported parasitic Hymenoptera on the pink bollworm in the lower Colorado Desert of California and Arizona during 1969-78 we thwarted by widespread insecticide application, even though field reproduction of eight species was recorded.  Inundative releases of parasitoids produced varying levels of pink bollworm reduction, the best performance being attained with egg-larval parasitoids, Chelonus spp. (Braconidae).  A Chelonus sp. nr. curvimaculatus (Cameron) obtained from the presumed native range of pink bollworm in northwestern Australia was most effective, giving an adjusted 69.9% infested boll reduction by August 24 at the equivalent release rate of 2,667 females/ha. (Legner & Medved 1979).  Extensive collections in northwestern Australia in 1991-92 revealed that Apanteles oenone and a Dirhinnus sp. were very active on P. gossypiella  (J. Altmann, unpub. data).  Nutritional studies have been conducted by Legner & Thompson (1977)


       Insecticide applications eventually eliminated attempts to biologically control pink bollworm in the United States.  Insecticides represent the primary control measure which has been successful in limiting damage of pink bollworm in commercial cotton.  However, during more than 40 years of application, insecticides have not solved the problem anywhere in the world:  each growing season finds pink bollworm present and developing resistance to toxic compounds. 




       All of the common predators in cotton fields are capable of feeding on pink bollworm eggs and first instar larvae (Jackson 1980).  Biological control programs were initiated at the University of California during the late 1960's.  Field work during this period focused on survey of predators occurring naturally in the lower desert.  Laboratory studies were summarized by Orphanides et al. (1971);  field studies were summarized by Irwin et al. (1974).  These studies show that several groups of arthropods attack pink bollworm naturally, including mites, predaceous Dermaptera, Hemiptera, Coleoptera and Neuroptera.  The egg stage is most vulnerable to attack by predators because it is relatively exposed when compared to larvae and pupae.  Most predators lack morphological modification of the legs and mouthparts necessary to penetrate bolls to feed on pink bollworm larvae.  The dermapteran Labidura riparia (Pallas) attacks all immature stages of pink bollworm including the pupa (Orphanides et al. 1971).  However, the predator is not a dominant element in a predatory complex.  Hemiptera are abundant in cotton in southeastern California, and at least five species in five genera of Hemiptera have been recovered from pink bollworm.  Hemiptera seem to express the broadest range of attack, feeding on eggs, larvae, cocooned larvae and probably pupae.  Irwin et al. (1974) found that Nabis, Geocoris and Orius all demonstrated effectiveness in field studies.  Coleoptera are well represented with four species in four genera attacking pink bollworm.  Most beetles focus on eggs and early instar larvae as prey.  Chrysoperla carnea (Stephens) is the only neuropteran reported attacking pink bollworm in California, and seems to prefer eggs and early instar larvae (Orphanides et al. 1971).  There is little data about the consumption of pink bollworm by predators under natural field conditions.


       The alfalfa/cotton ecosystems are basically composed of eight genera of numerically important primary consumers, six genera of predators and associated parasitoids and hyperparasitoids in New Mexico (Gordon et al. 1986).  A number of parasitoid species and most predaceous arthropods are non host specific (Huffaker & Rabb 1984).  Ehler & van den Bosch (1974) suggested that most common predators in cotton fields are opportunists that switch between primary consumer and other predaceous prey species depending on availability.  Although animal populations may be associated for a number of reasons, predation is one of them, and the measurement of interspecific association may be used to reduce the total possible testable combinations (Smith 1980).  Ellington (1988) analyzed 17 predator/primary consumer and predator/predator genera from 862 cotton samples for co-occurrence from 1981-84 and 1987.  There were 273 significant co-occurrences above three (3-14).  A switching parasitoid or predator can stabilize an otherwise unstable host/parasitoid interaction (Murdock 1969, Royama 1971).  A primary consumer like pink bollworm entering an ecosystem with a high density, species rich arthropod complex, may experience a damped density response.


Arthropod Complexity in Cotton


       The cotton ecosystem is very complex.  In Arkansas about 600 species of predators representing 45 families of insects, 9 families of spiders and 4 families of mites may be associated with cotton (Whitcomb & Bell 1964).  In the San Joaquin Valley, California, 300-350 arthropod species may be found in cotton (van den Bosch & Hagen 1966).  This complex arthropod group may be composed of three trophic levels:  primary consumers, parasitoids and hyperparasitoids and predators.  Some 15-30 genera may be present in sufficiently high numbers to warrant evaluation.  Included in this group are four superfamilies of minute, parasitic Hymenoptera which are responsible for the biological control of many key phytophagous species and are often unfortunately the first arthropods to be removed by insecticide application because they have lower genetic variability and exhibit lower detoxification capacities (Croft 1990). 


Potential For Biological Control


       The gelechiid moth genus Pectinophora contains three species:  P. gossypiella (Saunders), P. scutigera (Holdaway), and P. endema (Common).  All three species occur in Australia:  P. gossypiella occurs in the Northern Territory and Western Australia; P. scutigera and P. endema occur in Queensland.


       Life history data is provided by Fullaway (1909) and Busck (1917) for pink bollworm in Hawaii.  Similar information is provided by Gough (1916, 1920) and Ballou (1918) for pink bollworm in Egypt and Hunter (1918) and Loftin et al. (1921) for pink bollworm in Mexico.  Russo (1940) provided an extensive description of anatomy for all stages of development, based on studies in Somalia.  Holdaway (1926) discussed the bionomics and biology of spotted pink bollworm in relation to pink bollworm.  He concluded that the spotted species was native to Queensland and that its primary host plants were Hibiscus tiliaceus and Thespesia populnea; commercial cotton was a secondary host.  Common (1958) reviewed the species of Pectinophora associated with cotton in Australia, described P. endema from eastern Australia, and provided a key to the species.


       All species of Pectinophora are potential pests because they feed upon the buds, flowers and seed capsules of malvaceous plants.  The list of hostplants for pink bollworm is extensive and has been summarized by Li (1936) and Noble (1969).  Pectinophora gossypiella and P. scutigera are pests of cotton; P. endema attacks hibiscus in Australia but not cotton.  Pectinophora gossypiella became a commercial problem because its larval stage frequently enters diapause while in seed capsules, which enabled the pest to become widespread.  In contrast the spotted bollworm does not enter larval diapause within seed capsules and therefore has not become widespread (Sloan 1946).  In Queensland P. scutigera is not an especially important pest of cotton, but can cause damage to late maturing bolls and reach more serious levels of infestation when cotton is grown near other host plants.


       During the past 80 years, 160 species in 43 genera of parasitic Hymenoptera have been collected with or reared from pink bollworm infesting cotton (Gordh 1989), most of which are probably casual.  The Hymenoptera are divided into at least 13 superfamilies which contain about 80 families and more than 100,000 identified species.  Within this numerically large and biologically diverse group, species which attack pink bollworm are restricted to the superfamilies Ichneumonoidea, Chalcidoidea and Chrysidoidea. 


       Although several genera of ichneumonids have been reported attacking pink bollworm, no group prevails.  In contrast, the genera Apanteles, Bracon and Chelonus among the Braconidae contribute numerous species of parasitoids.  Gordon Gordh (1992) reported that only two genera of Trichogrammatidae have been taken from pink bollworm eggs, but it is doubtful that these collections were actually from that host.  Trichogramma is cosmopolitan in distribution and presently contains more than 12 species.  Biological information developed on the genus indicates that host specificity is not common.  Gordh stated that the genus is substantially larger than presently recognized and probably contains many species which potentially attack pink bollworm eggs.  The problem with this is that true pink bollworm, P. gossypiella sequesters its eggs under the calyx and other plant structures in a manner that is not generally available to parasitoids in the Trichogrammatidae.


       The genus Trichogrammatoidea has generally been ignored in biological control work.  This genus contains about 30 species and is predominantly Indo-Australian in distribution.  A few records exist for the new world, but these probably represent expansions of the natural geographical distribution by species transported in commerce.  Most recently two species have been recovered from Pectinophora scutigera in Queensland, Australia, which deposits its eggs in an exposed position on host plants.  Gordon Gordh reported that they also readily attack P. gossypiella (G. Gordh, unpub. data), but this information is undoubtedly wholly from data secured in confinement cages in the laboratory, where indeed trichogrammatids are easily mass cultured (J. Altmann & E. F. Legner, personal observations).


History of Biological Control Effort


       Biological control efforts were undertaken in Egypt during 1928-35, with the introduction of Bracon mellitor (Say) from Hawaii, Bracon kirkpatricki (Wilkinson) from Kenya and the Sudan, and Bracon lefroyi (Dudgeon & Geough) from India (Kamal 1951).  No practical results were reported although the latter species became established.


       In the United States importations to Texas during 1932-35 were made of Bracon brevicornis (Wesmael) from Europe, and B. mellitor from Hawaii.  Chelonus blackburni (Cameron) from Hawaii and Exeristes roborator F. from Egypt.  Additional strains of these species as well as B. nigrorufum (Cushman) and Chelonus pectinophorae (Cushman), were made from Korea during 1937-44.  Although large numbers were released, they were not established.  Failure to do so was attributed to concurrent heavy insecticide usage.  Final attempts to establish parasitoids from India were made in 1953-55.  Bracon brevicornis, B. gelechiae (Ashmead), Chelonus heliope (Gupta), and Apanteles angaleti (Muesebeck) importations also failed, reportedly due to intensive pesticide treatment and low winter temperatures (McGough & Noble 1955, 1957; Noble 1969, Noble & Hunt 1937).


       Bracon kirkpatricki and C. blackburni were introduced in southern Arizona during 1917-74, with only minor impact on P. gossypiella being recorded (Bryan et al. 1973, 1976; Fye & Jackson 1973).  It is significant that most of these parasitoids were known from other hosts than P. gossypiella, and that they were colonized under unfavorable circumstances.  Therefore, when biological control efforts began in California in 1969, explorations were extended to acquire not only previously tested species, but specific ones as well (Legner & Medved 1979).


       In the biological control effort in California, parasitic Hymenoptera that were successfully reared from P. gossypiella were obtained during explorations in India, eastern Africa, southern Europe, Australia and Hawaii.  The parasitoids were cultured from 5-20 founders and colonized in cotton fields infested with P. gossypiella in southeastern California and western Arizona.  The potato tuberworm, Phthorimaea operculella (Zeller) was used for mass propagation, with no known deleterious effects on the biology of the parasitoids (Legner & Thompson 1977).  Specific experiments were designed to measure the ability of a particular parasitoid to permanently establish in cotton fields, and its regulative impact on the pink bollworm population (Legner & Medved 1979).  Eight out of 14 parasitic species introduced in the area reproduced in the field, but no species was ever recovered in the summer following the year of release, even though field reproduction in the release year was often significant and overwintering possibilities existed (Fye & Jackson 1973, Legner 1979).


       During the year in which releases were made, parasitoid recovery was greatest in autumn months following peaks in host abundance, which was probably due to a combination of parasitoid reproduction and more favorable environmental conditions for development in the cooler autumn period (Legner & Medved 1979).  Significant regulative impact against P. gossypiella by several parasitic species was detected (Legner & Medved 1979), but as no carry-over to the second year was possible, compounded control with time was not observable.  It was concluded that in the absence of widepread insecticide applications, annual early season parasitoid releases over large areas of cotton might result in a drop of pink bollworm density if continued for several years, with decreasing numbers of parasitoids required for control in succeeding years as pink bollworm densities drop through parasitization.


       A lack of genetic heterogeneity in imported parasitoids also could have restricted their influence especially as founder cultures never exceeded 20 individuals.  A further search in the presumed endemic range of pink bollworm in northwestern Australia and southern Indonesia (Common 1958, Wilson 1972), might yield additional effective species and strains.  However, a permanent self-perpetuating regulation may be difficult to attain as natural restrictions on the early germination of cotton at the higher latitudes in springtime may cause an asynchrony whereby emerging overwintering parasitoids cannot find hosts upon which to develop successfully (Legner 1979 ). 


       A search for natural enemies in the endemic range of P. gossypiella in northwestern Australia was supported by the University of California in 1981-82 (Sands & Hill 1982), and US/AID in 1981-83.  Two species of Elasmus (Westwood), E. broomensis and E. bellicaput (Girault) were found.  Elasmus broomensis was determined to be a primary parasitoid and considered as a useful agent for biological control (Naumann & Sands 1984).  Additional species found were Apanteles oenone Nixon and Chelonus sp. nr. curvimaculatus.  As part of the survey, 21 species of Lepidoptera including 5 Gelechiidae) were reared from the seed capsules of Malvaceae.  The possibility of hyperparasitic activity by the Elasmus was eliminated experimentally.


Genetic Incorporation Into Cotton of Bacillus thuringiensis Toxin


       The gene for a larvicidal toxin produced by Bacillus thuringiensis was incorporated into the genome of commercial cotton and an array of other plant species that are grown commercially for food and fiber.   The degree of pest control never reached economically acceptable levels, and resistance of Lepidoptera to the toxin as of 2017 has very high, as would be expected from any non-living killing agent.  The levels of control from many previous widespread field applications of Bacillus thuringiensis were never spectacular, so that the poor performance was not surprising. 


       Additional detail on biological / integrated control, biology of hosts  natural enemies and pheromone disruption may be found in the "References" section. 



REFERENCES:          [Additional references may be found at:   MELVYL Library ]


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Abdel-Fattah, M. I., M. M. Hosny & G. El-Saadany.  1980.  The spacing and density of cotton plants as factors affecting populations of the bollworms, Earias insulana Boisd. and Pectinophora gossypiella (Saund.).  Bull. Ent. Soc. Egypt 60:  85-94.


Abdel-Rahim, W. A., S. M. I. Metwally & F. El-Dakrousy.  1980.  Effect of certain physical and chemical characteristics of cotton varieties on susceptibility to infestation by pink and spiny bollworm.  Plant Prot. Dept., Tanta Univ., Kaft-el-Sheikh, Egypt.  p. 727-31.  (RAE, A:  69:  5118).


Abou-Zeid, N. A., M. S. I. El-Dakroury, A. H. El-Heneidy & M. S. T. Abbas  1978.  Biology of Microplitis rufiventris Kok. parasitising Heliothis armigera Hb. in Egypt (Hymenoptera: Braconidae: Lepidoptera: Noctuidae).  Agric. Res. Rev 1978:  31-36.


Abul-Nasr, S., E. D. Ammar & A. I. Merdan.  1978/1979.  The control of the cotton bollworms, Pectinophora gossypiella (Saund.) and Earias insulana (Boisd.).  Bull. Ent. Soc. Egypt, Econ. Ser. 11:  35-9.


Abul-Nasr, S., E. D. Ammar & S. M. Farrag.  1979.  Rates of infestation by Pectinophora gossypiella Saunders and Earias insulana Boisd. on flowering sites of the cotton plant (Lep.).  Deutsche Entomologische Z. 26:  165-72.


Adkisson, P. L., L. H. Wilkes & S. P. Johnson.  1958.  Chemical, cultural and mechanical control of the pink bollworm.  Texas Agric. Expt. Sta. Bull. 920.


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Awate, B. G., L. M. Naik & G. Y. Parlekar.  1977.  Possibility of introducing exotic parasite Trichogramma brasiliensis Ashmead in the integrated control of cotton bollworms.  Cotton Development 7:  21-2.


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Bryan, D. E., R. E. Fye, C. G. Jackson & R. Patana.  1973.  Releases of bracon kirkpatricki (Wilkinson) and Chelonus blackburni Cameron for pink bollworm control in Arizona.  USDA ARS Prod. Res. Rept. 150.  22 p.


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