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Datasheet

Nomadacris septemfasciata
(red locust)

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Datasheet

Nomadacris septemfasciata (red locust)

Summary

  • Last modified
  • 20 November 2019
  • Datasheet Type(s)
  • Invasive Species
  • Pest
  • Preferred Scientific Name
  • Nomadacris septemfasciata
  • Preferred Common Name
  • red locust
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Arthropoda
  •       Subphylum: Uniramia
  •         Class: Insecta

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Pictures

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PictureTitleCaptionCopyright
Adult Red Locust (Nomadacris septemfasciata) on wild sorghum in the Wembere Plains in Central Tanzania in February 2003.
TitleAdult
CaptionAdult Red Locust (Nomadacris septemfasciata) on wild sorghum in the Wembere Plains in Central Tanzania in February 2003.
CopyrightReleased into the Public Domain by Christiaan Kooyman.
Adult Red Locust (Nomadacris septemfasciata) on wild sorghum in the Wembere Plains in Central Tanzania in February 2003.
AdultAdult Red Locust (Nomadacris septemfasciata) on wild sorghum in the Wembere Plains in Central Tanzania in February 2003.Released into the Public Domain by Christiaan Kooyman.
Nomadacris septemfasciata (the red locust); close-up of anterior region.
TitleAdult
CaptionNomadacris septemfasciata (the red locust); close-up of anterior region.
Copyright©Alex Franc - CC BY-SA 3.0
Nomadacris septemfasciata (the red locust); close-up of anterior region.
AdultNomadacris septemfasciata (the red locust); close-up of anterior region.©Alex Franc - CC BY-SA 3.0

Identity

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Preferred Scientific Name

  • Nomadacris septemfasciata Audinet-Serville, 1838

Preferred Common Name

  • red locust

Other Scientific Names

  • Acridium coangustatum Lucas, 1862
  • Acridium fasciferum Finot, 1907
  • Acridium purpuriferum Finot, 1907
  • Acridium sanctae-mariae Finot, 1907
  • Acridium septemfasciatum Audinet-Serville, 1938
  • Acridium subsellatum Finot, 1907
  • Cyrtacanthacris coangustata Kirby, 1910
  • Cyrtacanthacris fascifera Walker, 1870
  • Cyrtacanthacris purpurifera Walker, 1870
  • Cyrtacanthacris sanctae-mariae Kirby, 1910
  • Cyrtacanthacris septemfasciata Kirby, 1902
  • Cyrtacanthacris subsellata Walker, 1870
  • Nomadacris fascifera Orian, 1957
  • Nomadacris septemfasciata insularis Chopard, 1936
  • Patanga septemfasciata Jago, 1981
  • Schistocerca septemfasciata

International Common Names

  • English: locust, red; locust, red-winged; red-winged locust
  • Spanish: langosta roja
  • French: criquet nomade

Local Common Names

  • Angola: gafanhoto vermelho
  • Germany: Rote Wanderheuschrecke; Rotflügelige Wanderheuschrecke; Schrecke, Nomadacris-; Wanderheuschrecke, Rote; Wanderheuschrecke, Rotflügelige
  • Madagascar: valala mena elatra
  • Mozambique: gafanhoto vermelho
  • South Africa: rooi sprinkhaan
  • Tanzania: nzige mwekundu

EPPO code

  • NOMAFA (Nomadacris fascifera)
  • NOMASE (Nomadacris septemfasciata)

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Arthropoda
  •             Subphylum: Uniramia
  •                 Class: Insecta
  •                     Order: Orthoptera
  •                         Family: Acrididae
  •                             Genus: Nomadacris
  •                                 Species: Nomadacris septemfasciata

Notes on Taxonomy and Nomenclature

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The species was first described under the genus Acridium by Audinet-Serville in 1838. It was placed in the genus Cyrtacanthacris by Kirby in 1902. A number of closely related species was described in the genera Acridium and Cyrtacanthacris, but these were synonymized with septemfasciata by Uvarov (1923a), who also erected a separate genus for the species: Nomadacris. Dirsh (1979) argued that Nomadacris should be synonymized with the genus Cyrtacanthacris, but this was contested by Jago (1981), who proposed to make Nomadacris a subgenus of Patanga. Though the name Patanga septemfasciata is occasionally encountered, most authors continue to use Nomadacris septemfasciata.

Description

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Eggs

Eggs are laid in pods containing 20-100 eggs (gregarious females) or 20-195 eggs (solitarious females). The egg masses are straight and 2-4 cm long by 8-10 mm wide. They are covered on top by a froth plug of about 3 cm long, but they are not surrounded by a wall of froth mixed with sand grains like in other grasshopper species. The eggs are 5-7 mm long and arranged radially. Their colour is fawn (brownish) and they are covered with a strong chorion exhibiting a hexagonal fine-structure with small tubercles at the angles (Popov et al., 1990).

Nymphs

At hatching, the nymphs are still loosely covered with the embryonic cuticle (Smee, 1936). It is shed as soon as the nymphs emerge from the soil and heaps of these exuviae can be found at recently hatched egg fields. There are six to eight nymphal instars, solitarious locusts having more instars than gregarious ones. Purely solitarious nymphs are mostly green with a thin black line under the eyes. Gregarious ones are brown, black and yellow (Faure, 1932). They typically have a yellow pronotum with a median black stripe and a black spot on the sides, and a black spot on the distal half of the hind femora. Intermediate forms ('transiens') have varying amounts of black depending on the degree of gregarisation.

Adults

N. septemfasciata is a large locust: females are 55-85 mm, males 50-70 mm. Solitarious adults are larger on average than gregarious ones. The former are largely (reddish) brown and grey. Their tegmina carry oblique spots or fasciae (usually seven) and their wings are clear and pale red to purplish at the base. Gregarious adults are more reddish in general colour especially the younger ones.

Distribution

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The main breeding areas of N. septemfasciata, from where the majority of invasions and plagues originate, are the Mweru wa Ntipa Valley in north-western Zambia, the Kafue flats in southern Zambia, the Rukwa Valley in south-western Tanzania, the Malagarasi and Wembere plains in central Tanzania, Lake Chilwa in southern Malawi, the Busi-Gorongosa flood plains in Mozambique, and to a lesser extent southern Madagascar. Other important breeding areas are the central Niger delta in Mali and the southern shores of Lake Chad. Swarms do develop from time to time in the latter areas, but plagues have never resulted. Finally, resident breeding populations occur on Mauritius, Réunion and the Cape Verde islands. During the last major plague (1930-1944), which originated in the Mweru wa Ntipa and Rukwa Valleys, swarms invaded areas from the South African Cape Province to Khartoum and from Angola to Kenya and Somalia. Since then, invasions have occasionally occurred, for example, in the early 1970s and most recently in 1996/97, but these never developed into long-lasting plagues.

In addition to the records listed, the species has been captured in Chad and Niger (C. Kooyman, CAB International, African Regional Centre, Nairobi, Kenya, personal communication, 1991/1992).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Last updated: 17 Feb 2021
Continent/Country/Region Distribution Last Reported Origin First Reported Invasive Reference Notes

Africa

AngolaPresent, Few occurrences
BotswanaPresent, Few occurrences
BurundiPresent, Few occurrences
Cabo VerdePresent
CameroonPresent, LocalizedNative
ChadPresent, LocalizedNative
ComorosPresent
Congo, Democratic Republic of thePresent, Few occurrences
EswatiniPresent, Few occurrences
EthiopiaPresent, Few occurrences
GabonPresent, Few occurrences
GuineaPresent, Few occurrences
KenyaPresent, Few occurrences
LesothoPresent, Few occurrences
MadagascarPresent, LocalizedNative
MalawiPresent, Widespread
MaliPresent, LocalizedNative
MauritiusPresent
MozambiquePresent, WidespreadInvasive
NamibiaPresent, Few occurrences
NigeriaPresent, LocalizedNative
RéunionPresent
RwandaPresent, Few occurrences
SomaliaPresent, Few occurrences
South AfricaPresent, Few occurrences
SudanPresent, LocalizedNative
TanzaniaPresent, Widespread
UgandaPresent, LocalizedNative
ZambiaPresent, Widespread
ZimbabwePresent, LocalizedNative

Habitat

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The solitary phase has a preference for wet grasslands, especially those that are subject to flooding. The vegetation in these areas consists essentially of a mosaic of tall and short grasses and sedges without trees. Populations build up over several seasons and gregarisation sets in at sufficiently high densities aided by the fact that adults congregate in patches of green vegetation when the dry season progresses. When gregarisation is complete, swarms leave the outbreak areas. Breeding can then take place in any humid grasslands and even maize fields.

Hosts/Species Affected

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In the solitary phase, N. septemfasciata feeds mainly on grasses, especially species with relatively soft and juicy leaves like Echinochloa pyramidalis, Cynodon dactylon and several species of Cyperus (Chapman, 1957). Gregarisation changes its behaviour and it becomes more polyphagous. However, it continues to show a preference for grassy species, so cereal crops and sugarcane are especially vulnerable to attack. Pasture can also be affected, but the importance of this damage is not clear. The locusts defoliate the plants leaving the midribs in species with hard leaves like sugarcane. Many other plants are attacked during plagues, especially by adult swarms. Prominent among these are citrus and other fruit trees, palm trees, tobacco, cotton, cassava and vegetables (COPR, 1982). Some damage may be done to trees during roosting when branches break off due to the weight of the locusts.

Host Plants and Other Plants Affected

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Plant nameFamilyContextReferences
CitrusRutaceaeOther
    Coffea (coffee)RubiaceaeOther
      Eleusine coracana (finger millet)PoaceaeMain
        Gossypium herbaceum (short staple cotton)MalvaceaeOther
          Manihot esculenta (cassava)EuphorbiaceaeOther
            Nicotiana tabacum (tobacco)SolanaceaeOther
              Oryza sativa (rice)PoaceaeMain
                Poaceae (grasses)PoaceaeWild host
                  Saccharum officinarum (sugarcane)PoaceaeMain
                    Sorghum bicolor (sorghum)PoaceaeMain
                      Zea mays (maize)PoaceaeMain

                        Growth Stages

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                        Flowering stage, Fruiting stage, Seedling stage, Vegetative growing stage

                        Symptoms

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                        Symptoms are not very specific and they depend on the type of plant/tree attacked and the degree of hunger of the pest. The leaves are usually the first plant parts to be attacked and these can be chewed almost completely or if they are rather hard, the major veins, especially the mid ribs, are left (Lea, 1938). In cereals, varying proportions of the ripening grains are chewed back. Seed pods and fruits may also be attacked. When hungry, the locusts may chew stems and bark.

                        List of Symptoms/Signs

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                        SignLife StagesType
                        Fruit / external feeding
                        Fruit / frass visible
                        Inflorescence / external feeding
                        Inflorescence / frass visible
                        Leaves / external feeding
                        Leaves / frass visible
                        Seeds / external feeding
                        Seeds / frass visible
                        Stems / external feeding
                        Stems / visible frass
                        Whole plant / external feeding
                        Whole plant / frass visible

                        Biology and Ecology

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                        The salient feature of all locusts is that they can occur in two phases: a solitary phase, in which population densities are low and the behaviour is just like ordinary grasshoppers, and a gregarious phase, in which high population densities trigger a change in behaviour and in morphological and physiological characteristics. Solitarious locusts are cryptically coloured and tend to avoid each other, except when mating. When they are forced together by circumstances, the first changes occur in their physiology. Within days, they start to produce certain pheromones that in turn affect their behaviour. They now actively group together and form hopper bands or swarms. Over the next few generations, when the conditions are right, morphological changes take place. In the red locust, these changes are most pronounced in the hoppers, which become more strikingly coloured. Gregarious adults are smaller and more reddish brown. Gregarious locusts also become less restricted in their choice of food.

                        Eggs are laid in bare patches of soil between clumps of grass. Gregarious females tend to lay in groups, preferring relatively loose soil. The density of egg pods can be very high: up to 50 per square foot (Smee, 1936). Up to three egg pods can be laid during a female's lifetime. The incubation period is roughly 30 days with a minimum of 18 days (Mozambique, 1937) and a maximum of 54 days (Botha, 1969) mainly depending on the temperature. There are 6-8 hopper (nymphal) instars, solitarious locusts having more instars than gregarious ones. The development periods of the second to fifth instars are much shorter than the other instars. Development from egg to adult usually takes about 2 months, but it can be as short as 37 days in warm areas (Frappa, 1935) and as long as 78 days in cool areas (Lea, 1938). Adults remain immature during the dry season and mature at the onset of the rains. Even in areas without a marked dry season, like south-western Uganda, they mature only after about 6 months (Morstatt, 1912).

                        In southern Africa, egg laying takes place from November to December (Frappa, 1935; Jack, 1936; Têtefort and Wintrebert, 1965). The first hoppers can appear in December. Many different instars are usually present at the same time because of repeated egg laying. Adults appear between February and May. In the central Niger delta and the Lake Chad basin, adults mature at the beginning of the rainy season between April and June, and egg laying can continue until mid August (Golding, 1934; Saraiva, 1939; Descamps, 1953). Hoppers may be around from July to October. Where the rains continue throughout the year, breeding is continuous, but the species is still univoltine (Morstatt, 1912).

                        After hatching, the hoppers immediately look for cover. The first two instars do not move around much. They are generally to be found in lower grasses, like Cynodon dactylon and Cyperus spp. (Smee, 1936). Grouping usually starts in the second or third instar. Later instars and adults prefer roosting in tall grasses, like Echinochloa and Sporobolus. They feed mainly on grasses with soft leaves and high water content (Cynodon, Echinochloa, Cyperus). Drying of the vegetation forces late hoppers and adults in increasingly smaller patches of green grass, which encourages gregarisation (Burnett, 1951; Anderson, 1964). Though a good rainy season provides sufficient food for an increasing population, several good seasons in a row do not necessarily cause populations to build up. This is most likely because a more closed vegetation cover provides less opportunity for egg laying. Another reason may be that wetter conditions favour the development of fungal diseases.

                        Swarm formation is favoured by dry conditions after a few good breeding seasons and is further enhanced by the burning of grass in the dry season (Symmons, 1959). A return to wetter conditions in the outbreak areas reverses the trend to gregarisation. Plagues can last for many years if swarms continue to find good opportunities for laying and hopper survival is good. At the same time, conditions should continue to encourage grouping, which keeps the locusts in the gregarious phase. It seems that an essential condition is the existence of certain 'retention areas', where swarms can survive the dry season without the need for risky migrations (Symmons, 1964). During the last plague, the most important retention areas were from southern Malawi to north-eastern Rhodesia and west of Lake Victoria and into Rwanda and the Democratic Republic of Congo. Indications are that it is adverse weather conditions that put an end to plagues, rather than control measures, though the latter may decrease their impact.

                        Natural enemies

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                        Natural enemyTypeLife stagesSpecificityReferencesBiological control inBiological control on
                        Entomophaga grylli Pathogen Adults; Arthropods|Nymphs
                        Epicauta brevipennis Predator Eggs
                        Epicauta ruficollis Predator Eggs
                        Epicauta velata Predator Eggs
                        Metacemyia calloti Parasite Adults; Arthropods|Nymphs
                        Metarhizium anisopliae var. acridum Pathogen Adults; Arthropods|Nymphs
                        Mylabris Predator Eggs
                        Mylabris dicincta Predator Eggs
                        Mylabris pertinax Predator Eggs
                        Podapolipus elongatus Parasite Adults; Arthropods|Nymphs
                        Scelio howardi Parasite Eggs
                        Scelio sudanensis Parasite Eggs
                        Scelio zolotarevskyi Parasite Eggs
                        Stomorhina lunata Predator Eggs

                        Notes on Natural Enemies

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                        Locusts and grasshoppers are attacked by a range of specialist and generalist pathogens, parasites and predators (Greathead, 1963; Greathead et al., 1994). These can conveniently be divided into organisms attacking the eggs and those attacking nymphs and adults.

                        There is only one group of parasitoids of grasshopper eggs: the Scelionidae (Hymenoptera). Three species are known to parasitize the eggs of N. septemfasciata. Eggs further fall prey to predators. Specialised predators are Stomorhina lunata and species of Anastoechus. It is likely that Systoechus spp. attack Nomadacris eggs as well, but no records are available. Among beetle larvae, those of some species of Mylabris (Meloidae) apparently specialise on grasshopper eggs. Most other predators seem to be generalists or occasional predators.

                        Two specialised parasitoids are known from N. septemfasciata: the dipterans S. lunata (Sarcophagidae) and Metacemyia calloti (Tachinidae). Predators of locusts are all considered opportunists, with the possible exception of some sphecid wasps, especially the genus Prionyx. No records are available on Prionyx spp. taking Nomadacris as prey. Species of herons and storks, especially the Abdim's stork, Ciconia abdimii, appear to concentrate on locusts and grasshoppers when these are abundant. Most other predators become interested only when hopper bands and swarms are present.

                        Pathogens of locusts include viruses, bacteria and fungi. Records of viruses and bacteria from N. septemfasciata are not available. However, the fungi Entomophthora grylli, Beauveria bassiana and Metarhizium anisopliae are known to infect red locust. Finally, unidentified nematodes (probably Mermis spp.) are often found parasitizing both hoppers and adults.

                        The importance of natural enemies in regulating red locust populations is still being debated. It is however clear that they can not always prevent outbreaks and plagues. Few detailed studies have been conducted. Stortenbecker (1967) claims that very high hopper mortalities (60-90%) were caused in the Rukwa Valley by robber-flies (Asilidae) and dragonflies.

                        Impact

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                        There are few good quantitative data on economic damage. The best attempts are from South Africa, though these are from the 1930s. Damage to pasture, maize and sugarcane was estimated at UK£20,000 during the invasions of 1933-35. An added complication is that many published estimates combine the damage caused by N. septemfasciata with that by other species, notably the migratory locust, Locusta migratoria migratorioides. There are of course many anecdotes of serious damage, but the economic implications on a national level are not always clear. Even so, the red locust is considered a serious pest and considerable resources are geared to its control. Expenditure by the International Red Locust Control Organization for Central and Southern Africa (IRLCO-CSA) amounted to about UK£240,000 in 1978 (COPR, 1982) and during the outbreaks of 1986/87 about US$600,000 was spent on insecticides alone (Musuna, 1988).

                        Detection and Inspection

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                        Monitoring the red locust populations in their known outbreak areas will give early warning of incipient outbreaks. Before these areas become inundated, hopper concentrations can be found by driving or walking through the grass bearing in mind that the first two instars are usually to be found on patches of Cynodon and Cyperus, while later instars prefer taller grasses like Echinochloa. When the terrain becomes impassable, light helicopters can be used to flush adults. Early in the morning, even hopper bands can be observed basking in the tops of the grasses. Helicopters or fixed-wing aircraft can also be used to detect escaping swarms.

                        Similarities to Other Species/Conditions

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                        There are a few other large locust or grasshopper species that can be confused with N. septemfasciata. Confusion arises most often with species of Ornithacris, especially O. turbida. This species has reddish wing bases as well, but the shade tends more to orange. O. cyanea and O. imperialis magnifica are also often confused with N. septemfasciata, but their wings are more violet and the second species is even larger and its pronotum has a sharp arcuate median keel. Ornithacris spp. can easily be recognized by an oblique cream stripe along the lower margins of the pronotum, while in Nomadacris, this stripe is horizontal. Male Ornithacris have short, pointed cerci, while the cerci of male Nomadacris are elongated with a truncated tip.

                        Two other sympatric species can easily be confused with N. septemfasciata: Cyrtacanthacris tatarica and C. aeruginosa. Especially the first of these species is superficially similar because it also has oblique fasciae on its tegmina. However, both species have colourless wings or they are tinted yellow at the base. The male cerci are slender and pointed.

                        Other large locusts that often have reddish to purplish wing bases, are the tree locusts (Anacridium spp.). They can be distinguished by their black antennae, black fascia on the wings (very faint in A. melanorhodon melanorhodon) and the male subgenital plate having three lobes. The desert locust, Schistocerca gregaria, is not usually found in the same areas as the red locust. It has no fasciae on its tegmina and its wings have no red. The two remaining sympatric locust species, the migratory locust Locusta migratoria and the brown locust Locustana pardalina, are smaller than the red locust. They belong to the subfamily Oedipodinae, which lacks the prosternal peg of the Cyrtacanthacridinae.

                        Prevention and Control

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                        Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

                        Chemical Control

                        Fenitrothion has became the favoured product for swarm control because of its quick action. When the importance of the outbreak areas was recognized, it was decided in 1941 to establish the International Red Locust Control Service. Its successor, the International Red Locust Control Organization for Central and Southern Africa (IRLCO-CSA), is now responsible for the co-ordination of control efforts.

                        When previously used chemicals were banned in the 1980s, no good insecticide was available for hopper control. Organophosphates and the more recent pyrethroids are not as persistent  have therefore to be applied repeatedly as blanket sprays. The latest insecticide on the locust control market, fipronil, is probably not acceptable for the control of red locust in its outbreak areas because of its high toxicity to fish and aquatic invertebrates. Insect growth regulators, like diflubenzuron and teflubenzuron, might have some potential but have not yet been used against red locust.

                        Ecological Control

                        Several forms of ecological control have been tried. It was hoped that the introduction of cattle ranging would reduce the tall grasses favoured by red locust. However, it turned out to be difficult to make this commercially viable. Another idea was to control the burning of grass by the local inhabitants at the end of the dry season to reduce the number of bare patches where the locusts prefer to lay eggs. This proved costly and almost impossible to enforce. Tree planting trials failed, because most trees did not survive in this environment of highly variable water levels. Finally, an idea was investigated to control the water level of Mweru wa Ntipa, but it was considered to be too expensive (COPR, 1982).

                        Biological Control

                        Though many natural enemies have been recognized for a long time, none was ever considered effective enough to be used for control purposes. One attempt at biological control was the mass production and distribution of Beauveria bassiana in South Africa. The product failed to work and a subsequent investigation into the cause revealed that it contained mainly a saprophytic fungus instead of B. bassiana. Confidence in the product had disappeared and no new attempt was made. On Mauritius and Réunion, red locust populations have apparently been brought under control by the introduction of the Indian mynah bird, Acridotheres tristis, though the bird has become mainly frugivorous on Réunion (Bordage, 1913).

                        Recently, a new biological product has been developed by the LUBILOSA programme of CABI, IITA, GTZ and CILSS (Lomer et al., 1997a, b). Based on the fungus Metarhizium anisopliae var. acridum (previously identified as M. flavoviride), it was initially targeted for use against the desert locust, Schistocerca gregaria. During its development, it became clear that the fungus could infect most members of the superfamily Acridoidea (short-horned grasshoppers). In fact, it turned out to be almost totally specific to this group. It appears to be safe for humans and other vertebrates, and though it can infect other species of insects when these are under stress, no infections have been noted at recommended doses under (semi-)field conditions. Tests in the laboratory and on a small scale in the field have confirmed its infectivity to N. septemfasciata (Price et al., 1998). The product looks very promising, since the wet habitat of the red locust favours the development of fungi. Its slow mode of action (2-3 weeks) is no problem when applied against hoppers in the outbreak areas, because there, they do not attack crops. Large-scale trials are being planned.

                        References

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                        Anderson NL, 1964. Observations on some grasshoppers of the Rukwa Valley, Tanganyika. Proceedings of the Zoological Society of London, 143:395-403.

                        Anderson TJ, 1911. Annual Report of the entomologist for the year 1910-1911. Report of the Department of Agriculture of British East Africa, 161-171.

                        Audinet-Serville JG, 1838. Histoire naturelle des Insectes. In: Roret, Collection des Suites a BUFFON, OrthoptFres, Paris.

                        Barros Rodrigues Queiroz J de, 1934. Angola: locust movements. International Bulletin of Plant Protection, 8:M25-M29.

                        Barros Rodrigues Queiroz J de, 1935. Angola: movements of locusts (Nomadacris septemfasciata). International Bulletin of Plant Protection, 8:M147-M150.

                        Bordage E, 1913. Notes biologiques recueillies à l'île de la Réunion. Bulletin Scientifique France-Belgique, 47:395-396.

                        Botha HD, 1969. Locusts and their control in South Africa. Part II. The red locust. Farming South Africa, 45:52-53, 55.

                        Bredo HJ, 1936. Sommaire des observations faites au Congo Belge et projet des futures recherches sur les acridiens migrateurs. Bulletin Agriculture du Congo Belge, 27:298-302.

                        Bruner L, 1910. Acridoidea from Madagascar, Comoro Islands and eastern Africa. In: Voeltzkow A, ed. Reise in Ostafrika in den Jahren 1903-1905, Band 2. Stuttgart, Germany: E. Schweitzerbart'sche Verlagsbuchhandlung, NSgele & Dr. Sproesser, 623-644.

                        Burnett GF, 1951. Field observations on the behaviour of the Red Locust (Nomadacris septemfasciata Serville) in the solitary phase. Anti-Locust Bulletin No. 8.

                        Chapman RF, 1957. Observations on the feeding of adults of the Red Locust (Nomadacris septemfasciata (Serville)). British Journal of Animal Behaviour, 5:60-75.

                        Chopard L, 1936. Mission de M. A. Chevalier aux îles du Cap Vert (1934). II. OrthoptFres. Revue frantaise d'Entomologie, 3:88-96.

                        Chopard L, 1958. La rTserve naturelle intTgrale du Mont Nimba. III. Acridiens. MTmoires de l'Institut frantais de l'Afrique Noire, 53:127-153.

                        Chopard L, 1958. Les Orthoptéroïdes des Comores. Mémoires de l'Institut scientifique de Madagascar (E), 10:3-40.

                        Chopard L, 1958. Orthopteroidea. Résultats de l'expédition zoologique du Professeur Dr. Hakan Lindberg aux îles du Cap Vert durant l'hiver 1953-1954. No. 16. Commentat. Biol., Helsingfors, 17:1-17.

                        Chorley JK, 1939 Southern Rhodesia: locust invasion, 1932-1938. International Bulletin of Plant Protection, 13:M52.

                        COPR, 1982. The Locust and Grasshopper Agriculture Manual. London, UK: Centre for Overseas Pest Research.

                        Davey JT, Descamps M, Demange R, 1959. Notes on the Acrididae of the French Sudan with special reference to the Central Niger Delta. Part I. Bulletin de l'Institut frantais d'Afrique noire (sTrie A), 21:60-112.

                        Davey JT, Duhart AJ, Koné I, 1964. Notes on an incipient outbreak of the Red Locust (Nomadacris septemfasciata Serv.) in the central Niger delta (1959). Locusta, No. 9.

                        Descamps M, 1953. Observations relatives au criquet migrateur africain et a quelques autres espFces d'Acrididae du Nord-Cameroun. Agronomie Tropicale, Nogent 8:567-613.

                        Descamps M, 1965. Acridoïdes du Mali (DeuxiFme contribution). Régions de San et Sikasso (Zone Soudanaise). 1re partie. Bulletin de l'Institut frantais d'Afrique Noire, (série A) 27:922-962.

                        Dirsh VM, 1962. Acridoidea (Orthoptera) collected by Dr. F. Keiser in Madagascar. Verhandlungen der Naturforschungsgesellschaft von Basel, 73:270-275.

                        Dirsh VM, 1962. The Acridoidea (Orthoptera) of Madagascar. I. Acrididae (except Acridinae). Bulletin of the British Museum (Natural History) Entomology, 12:275-350.

                        Dirsh VM, 1979. The species and synonymy of the genus Cyrtacanthacris (Orth.: Acrididae). Eos, Madrid, 35:25-50.

                        Du Plessis C, 1949. Recent advances in the control of locusts in South Africa. Journal of the Entomological Society of South Africa, 12:3-12.

                        Faure JC, 1932. The phases of locusts in South Africa. Bulletin of Entomological Research, 23:293-405.

                        Faure JC, 1935. Life-history of the Red Locust (Nomadacris septemfasciata). Bulletin of the Department of Agriculture and Forestry of the Union of South Africa No. 144.

                        Finot A, 1907. Sur le genre Acridium. Annales de la Société Entomologique de France, 76:247-354.

                        Frappa C, 1935. Etude sur la sauterelle migratrice Nomadacris septemfasciata Serv. et sa présence à Madagascar de 1926 à 1935. Bulletin Économique de Madagascar, 3:203-221.

                        Golding FD, 1934. On the ecology of Acrididae near Lake Chad. Bulletin of Entomological Research, 25:263-303.

                        Golding FD, 1948. The Acrididae (Orthoptera) of Nigeria. Transactions of the Royal Entomological Society of London, 99:517-587.

                        Greathead DJ, 1963. A review of the insect enemies of Acridoidea (Orthoptera). Transactions of the Royal Entomological Society, 114:437-517.

                        Greathead DJ, Kooyman C, Launois-Luong MH, Popov GB, 1994. Les ennemis naturels des criquets du Sahel. Niamey, Niger: CILSS/DFPV.

                        Griffini A, 1897. Intorno ad alcuni Ortotteri raccolti dal Rev. L. Jalla a Kazungula (Alto Zambesi). Bolletin del Museo Zoologico e Anatomico comparativo de Torino, 12:1-12.

                        Hargreaves H, 1936. Report of the Government Entomologist for 1935. Report of the Department of Agriculture of Uganda 1935-1936:8-11.

                        Harris WV, 1933. The Red Locust. Pamphlet of the Department of Agriculture of Tanganyika, No. 10. Dar-es-Salaam, Tanzania: Department of Agriculture, Tanganyika.

                        Hemming CF, 1964. Red Locusts in Mauritius (Nomadacris septemfasciata Serville). Technical Circular of the Mauritius Sugar Industry Research Institute, 22:1-24.

                        Jack RW, 1931. Locusts in Southern Rhodesia. Rhodesia Agricultural Journal, 28:81-91.

                        Jack RW, 1936. Southern Rhodesia: locust invasion, 1932-1935. International Bulletin of Plant Protection, 10:M28-M29.

                        Jago ND, 1981. The genus Nomadacris Uvarov, 1923 and its recent incorrect synonymy under Cyrtacanthacris Walker, 1870 (Acrididae: Cyrtacanthacridinp), with new nomenclatural changes in the Patanga-Nomadacris-Austracris complex. Plant Protection Bulletin, India, 33(3/4):39-43

                        Kirby WF, 1902. Report on a collection of African Locustidae formed by Mr. W. L. Distant, chiefly from the Transvaal. Transactions of the Entomological Society of London, 1902:57-114.

                        Kirby WF, 1910. A Synonymic Catalogue of Orthoptera. Vol 3. Orthoptera Saltatoria. Part 2 (Locustidae vel Acrididae). London, UK: British Museum (Natural History).

                        Lea A, 1935. The Red Locust in Natal. South African Sugar Journal, 19:41-107.

                        Lea A, 1938. Investigations on the Red Locust Nomadacris septemfasciata (Serv.) in Portuguese East Africa and Nyasaland in 1935. Scientific Bulletin of the Department of Agriculture of the Union of South Africa No. 176.

                        Lean OB, 1931. Notes on the breeding of Nomadacris septemfasciata (Orth., Acrid.) on the shores of Lake Chad. Bulletin of Entomological Research, 22:571-572.

                        Lewin CJ, 1933. The locust invasion. Report of the Department of Agriculture of North Rhodesia 1932:15-18.

                        Lewin CJ, 1935. Locusts. Report of the Department of Agriculture of North Rhodesia 1934:14-16.

                        Lomer CJ, Prior C, Kooyman C, 1997. Development of Metarhizium spp. for the control of grasshoppers and locusts. Memoirs of the Entomological Society of Canada, 171:265-286.

                        Lomer CJ, with LUBILOSA project staff and collaborators, 1997. Metarhizium flavoviride: recent results in the control of locusts and grasshoppers. In: Krall S, Peveling R, Ba Diallo D, eds. New Strategies in Locust Control. Basel, Switzerland: BirkhSuser Verlag, 159-169.

                        Lucas H, 1862. OrthoptFres. In: Maillard L, ed. Notes sur l'Ile de la Réunion, Annexe I., 22-25.

                        Miller NCE, 1925. A list of Acrididae (Orthoptera) collected in the Tukuyu (New Langenburg) District, Tanganyika Territory. Annals and Magazine of Natural History, 15:618-634.

                        Miller NCE, 1929. Acrididae collected in Tanganyika Territory. Transactions of the Entomological Society of London, 77:61-97.

                        Miller NCE, 1936. A collection of Acrididae made in Southern Rhodesia (Orthoptera). Proceedings of the Royal Entomological Society of London, B 5:153-161.

                        Morant V, 1947. Migrations and breeding of the Red Locust (Nomadacris septemfasciata Serville) in Africa, 1927-1945. Anti-Locust Memoir No. 2, 66 pp.

                        Morstatt H, 1912. Die SchSdlinge und Krankheiten des Kaffeebaumes in Ostafrika. Pflanzer, 8:1-87.

                        Morstatt H, 1913. Liste schSdlichen Insekten Deutsch-Ostafrikas. Pflanzer, 9:288-296.

                        Mozambique, 1937. Contribution à l'étude biologique du Nomadacris septemfasciata Serv. 4th International Locust Conference, Cairo 1936 App. 35:1-10.

                        Musuna ACZ, 1988. Cereal crop losses caused by locusts in eastern, central and southern Africa region. Insect Science and its Application, 9(6):701-707

                        Okhoba MM, Zitsanza ES, Katheru JN, 2012. Dimba plains- a new red locust (Nomadacris septemfasciata) outbreak area in central Mozambique. Agricultural Science Research Journals, 2(10):534-540.

                        Orian AJE, 1957. Saltatoria, Phasmidae and Dictyoptera of Mauritius. Annals and Magazine of Natural History, 12th Series, 10:513-520.

                        Pinhey ECG, 1965. Check list of the short-horned grasshoppers of Syringa Farm, Turk Mine, Southern Rhodesia. Arnoldia, 2:1-20.

                        Popov GB, Launois-Luong MH, Weel JJ van der, 1990. Les oothFques des criquets du Sahel. Niamey, Niger: CILSS/DFPV.

                        Poulton EB, 1926. Protective resemblance borne by certain African insects to the blackened areas caused by grass fires. Verhandlungen des IIIen internationales entomologisches Kongress Zürich 1925, 2:12-94.

                        Rehn JAG, 1944. South African bird locust records and notes (Orthoptera, Acrididae, Cyrtacanthacridinae: group Cyrtacanthacres). Notulae Naturae, No. 137, 11 pp.

                        Robertson IAD, Chapman RF, 1962. Notes on the biology of some grasshoppers from the Rukwa Valley, S. W. Tanganyika (Orth. Acrididae). Eos, Madrid, 38:51-114.

                        Roblot M, 1951. Le criquet nomade (Nomadacris septemfasciata Serv.) au Soudan frantais. Agronomie Tropicale, Nogent 6:565-605.

                        Saraiva AC, 1939. A preliminary list of the insect pests of crops and fruit trees in Portuguese East Africa. Journal of the Entomological Society of South Africa, 2:101-114.

                        Schulthess A, 1898. OrthoptFres du pays des Somalis recueillis par L. Robecchi-Brichetti en 1891 et par le Prince E. Ruspoli en 1892-93. Annali del Museo di Storia naturale de Genova, 19:161-216.

                        Schulthess A, 1899. La faune entomologique du Delagoa. II. OrthoptFres. Bulletin de la Société vaudoise des Sciences Naturelles, 35:191-215.

                        Sjöstedt Y, 1913. Neue Orthopteren aus Ost- und Westafrika nebst einigen anderen zugehörigen Formen. Arkiv för Zoologi, 8:1-26.

                        Smee C, 1934. Report of the Entomologist. Report of the Department of Agriculture of Nyasaland 1933:46-53.

                        Smee C, 1936. Notes on the Red Locust (Nomadacris septemfasciata) in Nyasaland 1933-34. Bulletin of Entomological Research, 27:15-35.

                        Smit B, 1936. The Red Locust invasion of the eastern Cape Province during 1935. Farming South Africa 1936.

                        Stortenbecker CW, 1967. Observations on the population dynamics of the Red Locust, Nomadacris septemfasciata (Serville), in its outbreak areas. Pudoc Agricultural Research Reports, No. 694, 118 pp.

                        Stsl C, 1873. Stockholm. Recensio Orthopterorum, 1:154 pp.

                        Symmons PM, 1959. The effect of climate and weather on the numbers of the Red Locust, Nomadacris septemfasciata (Serv.), in the Rukwa outbreak area. Bulletin of Entomological Research, 50:507-521.

                        Symmons PM, 1964. The dynamics of the most recent plague of the red locust, Nomadacris septemfasciata (Serville), with special reference to the importance of climate and weather. PhD Thesis. Bristol, UK: University of Bristol.

                        TOtefort JP, Wintrebert D, 1965. Notes de mission au sujet de Nomadacris septemfasciata (Serville) et de Locusta migratoria L. à Maurice et à la Réunion. Agronomie Tropicale, 20:649-656.

                        Uvarov BP, 1923. A revision of the Old World Cyrtacanthacrini. I. Annals and Magazine of Natural History, 9th Series 11:130-144. II. Ibid. 11:473-490. III. Ibid. 12:345-366.

                        Uvarov BP, 1923. Notes on locusts of economic importance with some new data on the periodicity of locust invasions. Bulletin of Entomological Research, 14:31-39.

                        Uvarov BP, 1953. Grasshoppers (Orthoptera, Acrididae) of Angola and Northern Rhodesia, collected by Dr. Malcolm Burr in 1927 - 1928. Publicatoes Culturaes da Companhia de Diamantes de Angola, 21:9-217.

                        Walker F, 1870. Catalogue of the Specimens of Dermaptera Saltatoria in the Collection of the British Museum. Part III. London, UK: British Museum (Natural History), 485-594.

                        Walker F, 1871. Catalogue of the specimens of Dermaptera Saltatoria in the collection of the British Museum, Supplement, Part V. London, UK: British Museum (Natural History), 49-89.

                        Williams JR, 1964. Cane pests. Report of the Mauritius Sugar Industry Research Institute, 1963:91-93.

                        Distribution References

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                        Anderson T J, 1911. Report of the Department of Agriculture of British East Africa. 161-171.

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                        Bam A J, Conlong D E, Addison P, 2014. Grasshopper (Orthoptera) outbreaks on sugarcane in the Empangeni region of Zululand, KwaZulu-Natal, South Africa. In: 86th Annual Congress of the South African Sugar Technologists' Association (SASTA 2013), Durban, South Africa, 6-8 August 2013 [86th Annual Congress of the South African Sugar Technologists' Association (SASTA 2013), Durban, South Africa, 6-8 August 2013.], Mount Edgecombe, South Africa: South African Sugar Technologists' Association. 301-305. http://www.sasta2013.co.za

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                        Bordage E, 1913. [English title not available]. (Notes biologiques recueillies à l'île de la Réunion.). Bulletin Scientifique France-Belgique. 395-396.

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                        Descamps M, 1953. [English title not available]. (Observations relatives au criquet migrateur africain et a quelques autres especes d'Acrididae du Nord-Cameroun.). Agronomie Tropicale (Nogent). 567-613.

                        Descamps M, 1965. (Acridoïdes du Mali (DeuxiFme contribution). Régions de San et Sikasso (Zone Soudanaise). 1re partie). In: Bulletin de l'Institut frantais d'Afrique Noire, (série A), 27 922-962.

                        Dirsh VM, 1962. Acridoidea (Orthoptera) collected by Dr. F. Keiser in Madagascar. In: Verhandlungen der Naturforschungsgesellschaft von Basel, 73 270-275.

                        Dirsh VM, 1962a. The Acridoidea (Orthoptera) of Madagascar. I. Acrididae (except Acridinae). In: Bulletin of the British Museum (Natural History) Entomology, 12 275-350.

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                        Finot A, 1907. On the genus Acridium. (Sur le genre Acridium.). Annales de la Societe Entomologique de France. 247-354.

                        Frappa C, 1935. [English title not available]. (Etude sur la sauterelle migratrice Nomadacris septemfasciata Serv. et sa présence à Madagascar de 1926 à 1935.). Bulletin Économique de Madagascar. 203-221.

                        GOLDING F D, 1934. On the Ecology of Acrididae near Lake Chad. Bulletin of Entomological Research. 25 (pt. 2), 263-303 pp. DOI:10.1017/S0007485300012670

                        Golding F D, 1948. The Acrididae (Orthoptera) of Nigeria. Transactions of the Royal Entomological Society of London. 517-587.

                        Griffini A, 1897. [English title not available]. (Intorno ad alcuni Ortotteri raccolti dal Rev. L. Jalla a Kazungula (Alto Zambesi).). Bolletin del Museo Zoologico e Anatomico comparativo de Torino. 1-12.

                        Hargreaves H, 1936. Report of the Department of Agriculture of Uganda 1935-1936. 8-11.

                        Harris W V, 1933. The Red Locust. In: Pamphlet of the Department of Agriculture of Tanganyika, Dar-es-Salaam, Tanzania: Department of Agriculture.

                        HEMMING C F, 1964. Red locusts in Mauritius (Nomadacris septemfasciata Serville). In: Technical Circular. Mauritius Sug. Ind. Res. Inst. 1-24 pp.

                        JACK R W, 1931. Locusts in Southern Rhodesia. Rhodesia Agricultural Journal. 28 (1), 81-91 pp.

                        Jack R W, 1936. Southern Rhodesia: locust invasion, 1932-1935. International Bulletin of Plant Protection. M28-M29.

                        Kirby W F, 1902. Report on a collection of African Locustidae formed by Mr. W. L. Distant, chiefly from the Transvaal. Transactions of the Entomological Society of London. 57-114.

                        Lea A, 1935. The Red Locust in Natal. South African Sugar Journal. 41-107.

                        LEAN O B, 1931. Notes on the Bleeding of Nomadacris septemfasciata (Orth., Acrid.) on the Shores of Lake Chad. Bulletin of Entomological Research. 22 (pt. 4), 571-572 pp. DOI:10.1017/S0007485300035380

                        Lewin C J, 1933. Report of the Department of Agriculture of North Rhodesia 1932. 15-18.

                        Lucas H, 1862. Orthopteres. (Orthopteres.). In: Notes sur l'Ile de la Réunion, Annexe I. [ed. by Maillard L]. 22-25.

                        Miller N C E, 1925. A list of Acrididae (Orthoptera) collected in the Tukuyu (New Langenburg) District, Tanganyika Territory. Annals and Magazine of Natural History. 618-634.

                        Miller N C E, 1929. Acrididae collected in Tanganyika Territory. Transactions of the Entomological Society of London. 61-97.

                        Miller N C E, 1936. A collection of Acrididae made in Southern Rhodesia (Orthoptera). Proceedings of the Royal Entomological Society of London, B. 153-161.

                        Morant V, 1947. Migrations and breeding of the Red Locust (Nomadacris septemfasciata Serville) in Africa, 1927-1945. In: Anti-Locust Memoir, 66 pp.

                        Morstatt H, 1913. List of noxious insects (in German E. Africa). (Liste schádlicher Insekten.). Pflanzer. 9 (6), 288-296.

                        Okhoba MM, Zitsanza ES, Katheru JN, 2012. Dimba plains- a new red locust (Nomadacris septemfasciata) outbreak area in central Mozambique. In: Agricultural Science Research Journals, 2 (10) 534-540.

                        Orian AJE, 1957. Saltatoria, Phasmidae and Dictyoptera of Mauritius. In: Annals and Magazine of Natural History, 12th Series, 10 513-520.

                        Pinhey ECG, 1965. Check list of the short-horned grasshoppers of Syringa Farm, Turk Mine, Southern Rhodesia. In: Arnoldia, 2 1-20.

                        Poulton E B, 1926. Protective resemblance borne by certain African insects to the blackened areas caused by grass fires. Verhandlungen des IIIen internationales entomologisches Kongress Zürich 1925. 12-94.

                        Rehn J A G, 1944. Notulae Naturae, 11 pp.

                        ROBERTSON I A D , CHAPMAN E F, 1962. Notes on the biology of some grasshoppers from the Rukwa Valley, S.W. Tanganyika (Orth. Acridi-dae). Eos. 38 (pt. 1), 51-114 pp.

                        Roblot M, 1951. The Nomadic grasshopper (Nomadacris septemfasciata Serv.) in the French Soudan. (Le criquet nomade (Nomadacris septemfasciata Serv.) au Soudan francais.). Agronomie Tropicale, Nogent. 565-605.

                        Schulthess A, 1898. [English title not available]. (Orthopteres du pays des Somalis recueillis par L. Robecchi-Brichetti en 1891 et par le Prince E. Ruspoli en 1892-93.). Annali del Museo di Storia naturale de Genova. 161-216.

                        Schulthess A, 1899. [English title not available]. (La faune entomologique du Delagoa. II. Orthopteres.). Bulletin de la Société vaudoise des Sciences Naturelles. 191-215.

                        Sjöstedt Y, 1913. [English title not available]. (Neue Orthopteren aus Ost- und Westafrika nebst einigen anderen zugehörigen Formen.). Arkiv för Zoologi. 1-26.

                        Smee C, 1934. Report of the Department of Agriculture of Nyasaland 1933. 46-53.

                        SMEE C, 1936. Notes on the Red Locust (Nomadacris septemfasciata, Serv.) in Nyasaland 1933-34. Bulletin of Entomological Research. 27 (pt. 1), 15-35 pp. DOI:10.1017/S0007485300058090

                        Smit B, 1936. The Red Locust invasion of the eastern Cape Province during 1935. In: Farming South Africa, 1936

                        SYMMONS P, 1959. The effect of climate and weather on the numbers of the red locust, Nomadacris septemfasciata (Serv.), in the Rukwa Valley outbreak area. Bulletin of Entomological Research. 50 (pt. 3), 507-521 pp. DOI:10.1017/S0007485300053074

                        UVAROV B P, 1923. Notes on Locusts of Economic Importance, with some new Data on the Periodicity of Locust Invasion. Bulletin of Entomological Research. 14 (pt. 1), 31-39 pp. DOI:10.1017/S0007485300028182

                        Uvarov B P, 1953. Grasshoppers (Orthoptera, Acrididae) of Angola and Northern Rhodesia, collected by Dr. Malcolm Burr in 1927 - 1928. Publicatoes Culturaes da Companhia de Diamantes de Angola. 9-217.

                        Walker F, 1870. Catalogue of the Specimens of Dermaptera Saltatoria in the Collection of the British Museum. Part III. London, UK: British Museum (Natural History). 485-594.

                        Walker F, 1871. Catalogue of the specimens of Dermaptera Saltatoria in the collection of the British Museum, Supplement, Part V. London, UK: British Museum (Natural History). 49-89.

                        Williams JR, 1964. Cane pests. In: Report of the Mauritius Sugar Industry Research Institute, 1963 91-93.

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