UNIVERSITY OF IBADAN LIBRARY INSECTS: FRIENDS OR FOES? An inaugural lecture delivered at the University of Ibadan on Thursday, 04 June, 2015 By FRANCIS KOLAWOLE EWETE Professorof Entomology, Faculty of Agriculture and Forestry, University of Ibadan, Ibadan, Nigeria. UNIVERSITY OF IBADAN UNIVERSITY OF IBADAN LIBRARY , ' Ibadan University Press Publishing House University of Ibadan Ibadan, Nigeria. © University of Ibadan, 2015 Ibadan, Nigeria First Published 2015 All Rights Reserved ISBN: 978 - 978 - 8456 - 81- 0 'Printed by: Ibadan University Printery UNIVERSITY OF IBADAN LIBRARY The Vice-Chancellor, Deputy Vice-Chancellor (Administra- tion), Deputy Vice-Chancellor (Academic), The Registrar and other Principal Officers, Provost of the College of Medicine, Dean of the Faculty of Faculty of Agriculture and Forestry, Deans of other Faculties and Postgraduate School, Dean of Students, Distinguished Ladies and Gentlemen. Introduction I sincerely thank the University of Ibadan for giving me this opportunity to deliver an inaugural lecture on behalf of the Faculty of, Agriculture and Forestry at the twilight of disengaging from the University. I must thank the Dean of " my Faculty for nominating me to deliver this lecture on behalf of the Faculty as well as my Head of Department for ensuring that the lecture is delivered. Were it not for the extension of retirement age to 70 years, I would have missed this golden opportunity to stand before this audience, since I would have retired from the University in 2014. It has taken me 15 years after my elevation to the grade of Professor before this opportunity arose. ' This is the fourth in the series of inaugural lectures for the -2014/2015 academic session and also the 41st from the Faculty of Agriculture and Forestry since its inception in 1949. It is the 10th inaugural lecture from the Department of Crop Protection arid Environmental Biology, the 8th and 9th lectures having been delivered by Professors Bamidele Fawole and Babatunde Ikotun, respectively and the three of us were 1972 graduates of this Faculty. Mr. Vice-Chancellor, I must hasten to say that this is not by design. May I also re-iterate that this is the 4th inaugural lecture from the Entomology Unit of the Department, the first having been delivered by our erudite scholar and",teacher, Professor T. Ajibola Taylor of blessed memory, entitled "Insects and Our Environment", in 1974. This was followed by that of another teacher of ours, Professor Anthony Youdeowei, also titled, "Insects and Nigerians: The Struggle for Existence", in 1977, and the third by Professor J.A. Odebiyi titled, "Understanding and Exploitation of Balance of 1 UNIVERSITY OF IBADAN LIBRARY Nature in Insect Pest Management", in'2004. My interest was aroused in Entomology by Late Professor T. Ajibola Taylor shortly after my admission into the one year Postgraduate" Diploma Programme in Crop Protection, newly introduced in the then Department of Agricultural Biology during the 1973/74 session. There was a problem of obtaining good quality seeds for planting purpose in an okra breeding project initiated by the Late Professor H.R. Chheda in the Department of Agronomy. Here was I, just admitted into the diploma course and assigned the responsibility of tackling the problem under the joint supervision of Professors Taylor and Chheda. It is a well known fact that prior to being harvested for seed production, dry okra fruits are heavily infested by the seed bugs, Dysdercus (superstitiosus) voelkeri and Oxyca- renus gossypinus (fig. 1). Fig. 1: A colony of seed bugs Dysdercus (superstitiosus) voelkeri and Oxycarenus gossypinus. Please permit me to say here that the Late Professor Chheda and I had a good working relationship, having supervised my final year project in 1972 in the Department of Agronomy 2 UNIVERSITY OF IBADAN LIBRARY where I obtained my B.Sc. Honours (Agriculture) in Crop Science. After completing the final year project with him, he called me into his office one day and said: "Evete" (The Indians would normally pronounce the letter OW' as 'V' and vice versa), you just include my name as one of your referees whenever you are applying to anywhere". Before the com- pletion of the Postgraduate Diploma course, the preliminary results that I presented fascinated both of them. One day I was summoned to Professor Taylor's office and both of them asked me if I was ready to proceed to the M.Sc. (Agric.) degree if offered a scholarship. Of course I replied positively. Within one week, my letters of admission and award of Rockefeller Foundation Scholarship were ready. The rest is history and I now stand before you as an entomologist. What is Entomology? This is the branch of science that deals with the study of insects whether in the pure or applied form. What are Insects? Virtually everyone in this audience lives with one form of insects be it flies, mosquitoes, cockroaches, ants, termites, butterflies, et cetera, at home, in the office, or restaurant. Therefore we can all recognize a living creature known as insect. Technically however, insects are invertebrates (animals without backbone) in the Phylum Arthropoda which is the largest member in the Animal Kingdom. The classification of the Phylum and the Class Insecta to which all insects belong had been well enunciated by my three worthy predecessors and teachers in their inaugural lectures. This therefore spares me the time and energy to dissipate on this aspect. But basically for any organism to be grouped into the Class Insecta, the body must be divided into three distinct parts - head, thorax, and abdomen. The head bears a pair of compound eyes for sight, as well as filaments (antennae) for feeling the environment. The thorax is three-segmented, namely: prothorax, mesothorax, and metathorax with a pair of legs attached to each thoracic segment. Consequently, insects 3 UNIVERSITY OF IBADAN LIBRARY possess six (6) legs and hence are referred to as HEXAPODA. Mr. Vice-Chancellor sir, I know no other animal that possesses six legs. The closest relatives, ticks and mites, possess eight (8) legs. Other animals possess two (2) legs, as in birds, and four (4) legs in toads, lizards, pets, primates and ungulates. Insects are the most biologically successful animals in the Animal Kingdom. For instance, their small size accounts for their extremely low food requirement and the colonization of small niches in their thousands. In addition, they are highly adaptive to. extreme living conditions of hotness and coldness ranging from the equator to the arctic and subarctic regions of the earth, being poikilothermic (cold-blooded). Besides, insects are highly fecund (producing many offsprings) and exhibit different types of reproduction, namely: oviparous, -ovoviviparous, viviparous, polyembryony, paedogenesis, parthenogenesis and hermaphroditism. Oviparous reproduction is one in which eggs are laid and hatch into young ones. Majority of insects practice this form of reproduction. In ovoviviparous reproduction, eggs are retained within the body of the insect until they hatch as they are being laid. This occurs in most aphids and some flies. Viviparous reproduction is one in which embryos develop in the body cavity and when fully mature they are brought forth alive as found in man and other mammals. The tsetse fly . (Glossina spp.) that transmits sleeping sickness in cattle and man, is a good example. Polyembryony is one in which many offsprings emerge from a single egg. This is particularly common among endoparasitic insects that develop within the tissue of their host and are hence regarded as natural enemies. Paedogenesis is a form of reproduction in which immature stages (larvae) mature their ovaries very fast and begin egg- laying before attaining the adult stage, as it occurs in some gall midges. Parthenogenesis is a type of reproduction in which young ones develop from unfertilized eggs. Even within this type, there are three different forms based on the sex of the offsprings; it is thelytoky if the offsprings are 4 UNIVERSITY OF IBADAN LIBRARY exclusively female; arrhenotoky if exclusively male; and amphitoky if both male and female offsprings are produced. Hermaphroditism is reproduction in which both spermatozoa and eggs are produced by the same insect and these are fertilized by the spermatozoa and develop into offsprings, while the unfertilized eggs develop into males. Mr. Vice-Chancellor sir, I am neither attempting to bog down the audience with these entomological jargons nor make everyone an entomologist within this one-hour lecture, but I am only exposing the complexity of these organisms known as insects. They are also the only known lower organisms that possess wings to conquer the challenges of aerodynamics by flying to safety in times of danger or for any other exploratory activities to enhance their living. This is particularly true of the Monarch butterfly (Danaus sp) in Canada that flies southwards to Mexico it].late summer where they mate, deposit eggs and all die. The newly emerged adults (tenerals) now fly northwards to Canada in spring. Rote of Insects toMankind Mr. Vice-Chancellor sir, having highlighted what insects are, and that they are the most successful animal group on earth, one can now proceed with the role of insects to mankind. This will help us to discern those insects that are beneficial and/or harmful to man and his agricultural produce. The Beneficial Insects These are insects that contribute positively to the welfare and/or well-being of man in a number of ways through their activities in the ecosystem. The Pollinators These are groups of insects that visit flowers of cultivated crops and other plants to collect nectar and pollen grains for their food and in the manufacture of other useful commodities for man's use. Notable among these are the honey bees as well as butterflies, moths, wasps, flies and beetles. In the process of the visit, they inadvertently transfer pollen grains 5 UNIVERSITY OF IBADAN LIBRARY to the stigma of the flowers and hence effect pollination. This has resulted in fruit and seed production on an unimaginable scale leading to increased agricultural productivity and satisfying the food needs of man. Manufacturers of Useful Commodities A number of useful products are being manufactured by different insect species across various orders. These include. the following: (i) Honey, Propolis and Bees Wax These are products manufactured by the honey bee, Apis mellifera (Hymenoptera: Apidae). Honey is useful as a food source as well as for the medical treatment of woundslburns by preventing bacterial infection. The bees wax from hives is used in the production of candle. Bee-keeping is a big business world-wide and I must confess here that our Department keys well into such a venture through funding by the University Management, to generate funds for this University as pari of the Internally Generated Revenue (IGR). (ii) Wax This is a commodity from the nests of wasps (Hymenoptera: Vespidae) used in the manufacture of candle and polish. (iii) Shellac .This is a commodity obtained from scale insects (Hemiptera: Coccidae) and used in the making of paints as well as gums for sticking motor engine gaskets together. (iv) Cocoon (fig. 2) This is a woven silken material produced by the larvae of lepidopterous insects such as Anaphe venata when about to pupate. The harvested cocoons are unwounded into yams which are used in 6 UNIVERSITY OF IBADAN LIBRARY sericulture for making cloths such as silk cloth, and traditional Yoruba "aso oke (sanyan)". This is another commercial enterprise which individuals and corporate bodies can engage in for revenue generation. An attempt at this venture by the Department ended abruptly owing to abscondment by a Ph.D. student placedon the project. Fig. 2: A woven silken material from the larvae of Anaphe venata. Natural Enemies These are groups of insects that kill other insects which infest our agricultural products. They are the predators and parasites that feed on harmful insects on our crops. Predators are generally bigger than their prey, while parasites are smaller than their hosts. The predatory insects include those in figures 3,4 and 5. 7 UNIVERSITY OF IBADAN LIBRARY Fig. 3: Praying mantis - Mantis religiosa. Fig. 4: Assasin bugs - Pseudophonoctonus-formosus. 8 UNIVERSITY OF IBADAN LIBRARY Fig. 5: Rhinoeoris bieolor. Others are: Ladybird beetle - Cheilomenes lunata, Lacewings (Order: Neuroptera: Chrysopidae), Dragon flies (Order: Odonata). In the coccinelid beetle (Ladybird), assassin bugs and Mantis, both adult and larvae or nymphs kill and feed on their prey, while only larvae are involved in Neuroptera and Odonata. The parasitic insects, especially the endoparasitoids, are found mainly in two insect orders; Diptera and Hymenoptera. These include: Tachinid flies (Diptera: Tachinidae) Braconid Wasps (Hymenoptera: Braconidae) (fig. 6) Cha1cidWasps (Hymenoptera: Chalcidae) Ichneumonid Wasps (Hymenoptera: Ichneumonidae) 9 UNIVERSITY OF IBADAN LIBRARY Fig. 6: The braconid wasps (hymenoptera: braconidae). Braconid wasps use their slender ovipositor to deposit an egg, each within the tissue of their hosts later hatching into a young larva, continues to feed within the host tissue and eventually bursts open the host tissue to pupate outside and with the adult later emerging. In this way, the host is killed. It therefore follows that these groups of entomophagous insects that help in reducing insect pest populations to sub-economic levels can be manipulated for the control of major insect pests through a control strategy called Biological Control. Prof. J.A. Odebiyi enunciated this in his 2004 Inaugurall..ecture as part of the thrust of his work in this University in the control of cassava and mango mealybugs. Soil Nutrient Enrichment Certain soil-dwelling or subterranean insects such as the Springtails (Collembola) as well as dung beetle enrich the soil by degrading plant residue and/or mixing faeces with soil to increase the soil organic matter content which enhances crop growth and subsequent yield increases. This is of great advantage to farmers who most often practise zero-tillage. 10 UNIVERSITY OF IBADAN LIBRARY Edible Insects The larvae of some lepidopterans (butterflies, moths) as well as the larvae and adults of some beetles (palm weevil, 'palm beetle) (figs. 7, Sa & 8b), the winged termites and queen termite, adult crickets, and some grasshoppers (locusts, variegated grasshoppers), are good sources of protein and fat. For instance, travellers along the highways of Delta, Bayelsa States and the creeks of Ondo State witness the hawking of roasted or fried larvae of the palm beetle and the palm weevil, being sold along with 'papa garri' (a crumbed cassava meal). These supply cholesterol-free oils/fats and protein which aid human diet. Fig. 7: Some edible insects. 11 UNIVERSITY OF IBADAN LIBRARY Fig. 8a: Palm beetle. Fig. 8b: Palm beetle larvae. Insects as Aesthetics Indeed insects are very beautiful. The combination of colour display by butterflies, moths as well as other insects such as the beetles, dragonflies, and grasshoppers depict the work of nature. A mounted insect display in the home adds colour and serenity to the immediate environment. There is an insect species given the scientific name Cynthia spp, its common name being 'Painted Lady' and this is without prejudice to ladies who bear Cynthia in this audience. Mr. Vice-Chancellor sir, you know very well that ladies can be very beautiful when they make up but NOT painted. Artists also use insects as designs in the textile industry and they are also used as pendants to add value to dressing. Insect as Source of Wisdom According to the Holy Bible in Proverbs 6: 6-8, 1 quote: "You idler, go to the ant, watch her ways and be wise. She has no master, no steward or overseer. She secures food in summer and stores up provisions during harvest time". In a similar manner, man can learn from termites. The termites construct their mounds or termitaria gradually and reside in them. Thus the saying in Yoruba, Were ni ikan nmole (termites build gradually). They engage in division of labour using their different forms or castes such .as the soldiers, workers, winged reproductives and the queen 12 UNIVERSITY OF IBADAN LIBRARY normally housed in a 'royal chamber' with each performing specific functions for the colony. The soldiers are strictly for defense of the colony in case of any intruder or invasion by unwanted visitors. I dare say that no living creature (not even snakes) can reside in an active termitarium. In fact, the soldiers are the first to rush out to resist any aggression. The queen is for laying eggs and production of offsprings and population build-up. The queen is at the core of the termitarium heavily defended. This semi-fortress can be likened to the British Monarchy System of government in which the Queen is the ceremonial head and heavily defended by British troops. The workers reconstruct damaged termi- tarium and forage for food and feed the young ones as well as the queen that is sedentary in its royal chamber (fig. 9). The winged termites fly out to colonise a new area in case of population explosion. Fig. 9: The royal chamber of a termitarium and a queen termite. Insects as Weapons of War or Biological Warfare There are instances in which insects can be deployed for errands to punish stubborn people or to show resentment against bad behaviour. According to the Holy Bible, out of the ten (10) plagues Yahweh unleashed to torment Pharaoh 13 UNIVERSITY OF IBADAN LIBRARY for failure to release the Israelites from bondage, three (3) were of insect origin. These were the use of mosquitoes in the 3rd plague in Exodus 8: 13-14. The 4th plague was the use of horseflies in. Exodus 8: 20, 1 quote: ."Yahweh did this and dense swarms of horseflies invaded Pharaoh's house and the houses of all his people and devastated the whole country". The 8thplague was the use of locusts in Exodus 10: 13-15, and 1 quote: "So Moses stretched out his staff over the land of Egypt. All that day and night Yahweh brought an .east wind over the land and in the morning the east wind brought locusts. They devoured all the vegetation . . . . . . . . .. . .. . . . . . . . . . . .. and nothing green remained in the land of Egypt". Also, in Nigeria during the First Republic and according to an eye witness account by my very good friend, Professor S.O. Bada (then Mr. Bada), while the then Premier of the Old Western Region, Chief S.L. Akintola of the Nigerian National Democratic Party (NNDP) was on a campaign tour in 1965 at Ikare-Akoko; and as he mounted the podium, a swarm of bees besieged the campaign arena followed by a heavy thunder- storm. Everybody scampered for safety and so ended the campaign. The punitive measure might not be unconnected with the disagreement between Chief S.L. Akintola (the Premier of Western Region) and Kabiyesi Oba Amusa Momoh III, the then Olukare of Ikareland. Chief Akintola had earlier reduced the salary of Oba Momoh to a penny (one kobo) per annum for supporting the then Ooni of Ife (Oba Adesoji Aderemi) and the Action Group (AG) Party. His counterpart, Oba Samuel Akinsanya, the then Odemo of Ishara-Remo in the present Ogun State also had his salary reduced to a penny per annum by Chief Akintola for the same offence. The Harmful Insects This is a group of insects that feed on plants/plant products, animals/animal products and man, resulting in damage and/or 14 UNIVERSITY OF IBADAN LIBRARY death. This covers a, vast area of entomology which may be compartmentalized into the following: (i) Agricultural entomology (ii) Forest entomology (iii) Veterinary entomology (iv) Medical entomology (v) Forensic entomology Agricultural entomology covers aspects of insects infesting and causing damage to cultivated crops and reared animals in the field or in storage. Forest entomology is the study of insects infesting tree species in the forest as well as household forest products. I must state here that our Department has been responsible for the teaching of this course in the Department of Forest Resources Management right from its inception in 1963 as a department in this Faculty. In some developed parts of the world, such has grown into. a full- fletched department; especially at the University of British Columbia, Vancouver, Canada where I was a visiting scientist during my Sabbatical Leave at the University of Ottawa, Ottawa, Ontario, Canada in 1993. Veterinary entomology is the study of insects in relation to transmission of livestock diseases such as trypanosomiasis (sleeping sickness) by tsetse fly (Glossina palpalis, G.. morsitans) as well as, myasis transmitted by Cochliomya hominivorax in cattle. Medical entomology deals with insect vectors trans- mitting diseases such as yellow fever, malaria fever, dengue fever, typhoid fever: dysentery, cholera, diarrhoea, river blindness (onchocerciasis), sleeping sickness (trypa- nosomiasis) and their vector control. While Anopheles sp transmits malaria parasite, Culex sp. transmits yellow fever and dengue fever. Onchocerciasis is transmitted by the black fly, Simulium sp. and cholera, dysentery, and diarrhoea are transmitted by the housefly, Musca domestica. Forensic entomology is the scientific discipline that helps interpret information on homicide cases using insects -as 'silent witnesses' in order to provide evidence or clue which cannot be obtained from normal classic pathology (Usua 2004). 15 UNIVERSITY OF IBADAN LIBRARY .bIel1.,.Ger'insecets,pecially blowflies, can be used in medico- legal investigations since they are the first arthropods to visit a corpse within a few hours following death (Catts and Goff 1992). Mr. Vice-Chancellor, sir, kindly permit me to refer to a classic unfortunate incident in Tafawa Balewa (Postgraduate) Hall in 1998 when I was the Hall Warden. A student had died overnight in his room in B Block with the door locked. It was the ,swann of blowflies hovering on die window net and door m!lfellJI2~r1oUr toW21.J.ng morrn.ng that provi'ded m. r1o: rmatio• n that rrr 2 Emmemnnzs uwas arrn.s. hivou 2·~rJ. s In t e room. Beoh, ld on breaking the ~ "i~B8n,me student was already stone dead. The swarm of .~Rs/rfJX-ffl~f;s forensic indicator of corpses (Greenberg 1991). ru, ni 5 11. . , i}~ y,tto Agncultural Entomology W~tSJfn ..,..ea of insect study that deals with insect/host Plan,t i~l'lliliffi with respect to infestation of cultivated crops andlO}'ll~t~~5>Ck~,hedamage c~used and their control in ord~r to obtai rill~ YIelds. Here IS my account of stewardship spanning {~ period of over thirty two (32) years in this University. ";', I. Since the beginning, man by his inquisitive nature on life sustainability has from time to time explored the environment fOf sekaion and domestication, at: Dlants, and animals for ••••• doGf.tingami 9IDtlnm:. 1FIhe utIuntF rrOlm nomadic: life on' • C Ering fruits, ~,lluutblganimals:. to)dhealmtary form' ~ apiculture whereby nearb~ land is tiHkdf ffurgmwing crops and rearing animals indeed! lIIIlIikedthe origin' of organisms known as pests. The practice of growing crops or rearing animals together within a given area attracts these insects which consequently multiply in large. numbers to become the pests that destroy the crops and animals. Left alone in nature, these organisms, insects inclusive, interact in such a way that their population is in equilibrium with other living things (plants, animals) in the environment over a long period of time and thus establishes a General Equilibrium Position (GEP). Man's tampering with this near-stable environment by cultivating the land for crop and animal productions has 16 UNIVERSITY OF IBADAN LIBRARY displaced this equilibrium with the attendant rise in the number of the organisms or pests to reach the economic threshold and economic injury level (fig. 10). - - - - - - =t:••, - - ECOnoftlk:lnjuryw.,e(I - - -£tonornic ThnithoW I /' : I , I 11t---jl-:t-T+-++~t-I:-J4-/':.J., Gtnt~8JuiUbriu", l. Palit;a" (GEP ) 1••• r..•• • eor$ ) Fig. 10: A theoretical pest population rising to economic threshold and economic injury level. The economic threshold is the density at which control measures should be determined to prevent an increasing pest population from reaching economic injury level. In other words; it. is an .aetion thresihOlHat!wffiilthone should kick-start control', meaamYSS.However; economic: injury level is the lowest gcslt lJIIlPllati<»l1 tttatl will: cause, economic damage while.:tft' •••• lii.Lctamage·is;tH~diljurythan will justify the cost of artifi'Ciirnamntol measure and for it to take effect. Bactors affecting the Development of Insects to Pest status The following factors: wdlithrow more light into how insects rise to a pest status. (1) The selection of crops for palatability and nutritive value by man for growth, development and healthy living is also carried out by insects for the same 17 UNIVERSITY OF IBADAN LIBRARY purpose. Therefore insects compete very well with man for food. (2) The practice of monoculture whereby one .crop species is grown over a large hectarage has provided abundant and closely grown plants readily available as food sources, but also for cross infestation. (3) The practice of seed storage in large granaries such as the big silos at one of the Federal Government Strategic Grain Reserves situated along Iwo Road, opposite Dizengoff Company, in Ibadan. This provides a suitable habitat for continual repro- duction and population build-up of stored product insects such as the bean beetle, Callosobruchus maculatus on cowpea; grain weevils, Sitophilus zeamais, S. oryzae on maize and rice, respectively as well as Dermestes maculatus on stored animal products such as smoked fish, stockfish, leather, etc. (4) The transfer of insects across geographical barriers leaving behind their natural enemies will allow them to multiply unchecked until they attain pest status. This is clearly evident in the citrus scale insect, Icerya purchasi, which was imported along with good varieties of citrus from Australia to California in USA leaving behind its natural enemy, Rodolia cardinalis. It Was not until .rhis natural enemy was exported to California to .deal with this scale insect that it got eradicated. This was the first classical case of biological control of insects. Right here in Nigeria, Professor Odebiyi in his 2004 Inaugural Lecture reported the successful control of the Cassava mealybug and Mango mealybug that threatened cassava and mango production in 1979 and 1987, respectively. This was achieved through the release of their natural enemies into the field and they eventually got controlled. 18 UNIVERSITY OF IBADAN LIBRARY Insects in Relation to Crop Commodities and Plant Health It is convenient to group crops according to the commodities they supply and for pest identification and management purposes. Thus, there are groups such as the cereals, legumes, vegetables, fibres, porno logy , floriculture, tree crops and stored produce. Each of these groups is infested by insect pests to varying degrees, some as major pests and others minor pests, all impairing the growth and productivity of such crops culminating in yield and quality reduction. Evidence of research work conducted on major insects attacking some of the crops in each group, as my contribution to knowledge, will suffice. Insect Pests of Cereals The most important cereal crops cultivated in Nigeria include maize (Zea mays), rice (Oryza spp.), guinea corn (Sorghum spp.), millet (Pennisetum typhoides), and wheat (Hordeum triticum). They are a good source of carbohydrates in human diets and animal feed, and supply the needed energy. In table 1, the field insect pests of cereals are the stem borer complex, Eldana saccharina Sesamia calamistis, Busseola fusca, Acigona ignefusalis and Mussidia nigrivenella found to cause damage on farmlands in South-Western Nigeria during the early and late cropping seasons in 1991. E. saccharina and S. calamistis caused the highest percentage stalk length damage (Oigiangbe et al. 1997). However on rice, the lepidopterous stem borer complex was the most damaging, including the pink stem borer, Sesamia calamistis; yellow stem borer, Scirpophaga incertulas; striped borer, (Chilo suppresalis); Maliarpha sp. and a dipterous stem borer, Diopsis thoracica (Stalk-eyed borer) (fig. 11). Their effect is more severe in the late cropping season than in the early season. The two symptoms associated with stem borer attack are the 'window pane effect' (fig. 12) and 'dead heart' in maize but 'dead heart' and 'white head' (fig. 13) in rice. Up to 30% loss of stands has been recorded in severe infestation by these borers. Rice grains especially at the dough stage are attacked in the 19 UNIVERSITY OF IBADAN LIBRARY field by grain suckers, Aspavia armigera and Stenocoris elegans (Joda 2000; Alamu and Ewete 2014). A survey across fifteen (15) states in Nigeria, namely; Abia, Anambra, Delta, Ebonyi, Ekiti, Enugu, lmo, Lagos, Nassarawa, Niger, Ogun, Ondo, Oyo, and Plateau, revealed three (3) species of Aspavia and their natural enemies (Joda 2000). Table 1: Mean Number of Stemborer Species per Stalk during the First and Second Cropping Seasons in South Western Nigeria Borer species . Season N No of CV(%) t-value Stemborers Eldana First 850 0.77 56.71 0.1241ns saccharina Second 330 0.71 22.82 Seasamia First 850 0.51 83.14 4.2193* calamitis Second 330 0.81 35.61 Acigona First 850 0.11 256.28 1.8229ns ignefusalis Second 330 0.21 20.90 Busseola fusca First 850 0.01 207.45 2.5505* Second 330 0.08 55.73 Mussidia First 850 0.00 nigrivenella Second 330 0.35 52.69 ns= Not significant at P= 0.05, *= Significant at P ::;0.05 20 UNIVERSITY OF IBADAN LIBRARY l, 2J?~qh9UtI I:) zuonrmuge.I q noegiq .fTf;'Xi b'JL/t" J?Orti 'Al! Fig. 11: Diopsis tho'flitiC'f'(s~-iym1x>ter. bO,,'3.J II D:52U '1:)11£1~1rl1 .:).1 'l :.J rDl!uolg bus »q b-l1; fll.;'Jd L'Oc .t;:.Jq'IJO::l q j ~")tJqlf1')~fl(Y) 1L)dJ ~ filJ! :;J,lIJ; !2;:}(] bi~il ern :31 'J:nrlJ 5(5 d .znitrh 1'JIIOrt OJ n'e}1w5!1 i:bdow pan£errewffbect' bsy::m15p2tomlfI5o:t.ns· UNIVERSITY OF IBADAN LIBRARY Fig. 13: The 'white head' of stem borer attack in rice. Insect Pests of Legumes Leguminous crops which include groundnut, cowpea, soya bean, pigeon pea, sword bean and bambara groundnut supply the most needed plant protein in the diet of human beings and livestock. Groundnut is processed into both the groundnut oil and groundnut cake, the latter used in animal feed. However, cowpea, soya bean and pigeon pea are grown for human consumption for the needed plant protein. Field insect pests - attacking them have been well documented and studied. Of the field pests at the vegetative and reproductive phases of these three legumes, pests of the reproductive phase mainly flower thrips, Maruca pod borer and the pod sucking bugs, mainly the coreids and alydids, cause over 50-70 percent seed yield loss. The coreid bugs include Anoplocnemis curvipes, CZavigraZZatomentosicoZZis while the alydids are Riptortus dentipes and Mirperus jacuZus. The most studied of these bugs are C. tomentosicoZZis and R. dentipes in terms of their biology and damage caused on cowpea (Aina 1972; 1975a; Egwuatu and Taylor 1977; Akingbohungbe 1977). However, Ewete and Niba (1994) showed that Mirperus jacuZus completed its development (first instar - adult) in 19.8 days on Ife Brown cowpea variety and infestation of pods 2 weeks and 3 weeks after pod development caused 85.6-99.0 percent and 61.0-66.8 percent seed damage, respectively, at a 22 UNIVERSITY OF IBADAN LIBRARY population of 8 bugs per peduncle (table 2). Similarly, Ewete and Joda (1996) showed that Riptortus dentipes at a population of 8 bugs per plant caused 99.0-100.0 percent seed damage on soya bean varieties TGX 536- 02D and TGX 849- 294D but 90.5 percent seed damage on TGX 996-28E (table 3). Table 2: Percentage Seed damage (± S.E.) with varying Densities of Mirperus jaculus on Cowpea at three different Pod Ages Insect Number Age of Pods 1 Week 2 Weeks 3 Weeks o (Control) 7.62 ± 1.38 b 7.90 ± 2.14 c 8.02 ± 2.40 b 2 100.00 ± 0.00 a 85.60 ± 4.99 b 61.00 ± 12.95 a 4 100.00 ± 0.00 a 93.00 ± 4.52 d 64.80 ± 6.60 a 8 100.00 ± 0.00 a 99.00 ± 1.00 a 66.80 ± 13.85 a S.E. 1.8 7.49 19.82 CV% 2.34 10.49 39.5 Means followed by the same letters are not significantly different at 5 % level (Duncan's multiple range test): mean of 5 replications. Table 3:Degree of Seed damage (%) on Soybean caged with varying Densities of ~iptortus dentipes Insect Number Soybean Varieties· TGX536.02D TGX 996.28E TGX 849 294D o O.Oa 2.6a . l.4a 2 82.5b 29.9a 66.3b 4 98.5c 83.8b 80.0b 8 99.2c 90.5b l00.0c S.E 5.20 10.09 4.41 CV (%) 3.21 8.44 3.08 Means followed by the same letters are not significantly different at 5% level (LSD) Insect Pests of Vegetables Vegetables (leafy and fruit), supply minerals and vitamins needed for good and healthy growth of man and his animals. ·A good example of fruit vegetables are okra, tomato, pepper, garden eggs, eggplant, cucumber, water melon, et cetera. 23 UNIVERSITY OF IBADAN LIBRARY Some of the leafy vegetables are Amaranthus spp (Tete), Celosia argentea (Soko), Telfairia occidentale (Ugwu or iroko), Basella spp (Amunututu), Corchorus olitorius (ewedu), Vernonia amygdalin a (bitter leaf) and Cabbage. They are infested by defoliators, mostly beetles, larvae of butterflies and moths as well as the grasshoppers, while their fruits or grains are attacked by hemipterous bugs. Ewete (1978) identified six (6) major groups of insects infesting all parts of okra plants namely; foliage-feeders, root-feeders, stem-feeders, flower bud-and flower-feeders and fruit- feeders. Among these pests are three (3) major defoliators, Podagrica sjostedti, Podagrica uniforma , Sylepta derogata (leaf-roller); six (6) major green and dry fruit-feeders, Dysdercus (superstitiosus) voelkeri (fig. 14), Oxycarenus hyalinipennis, O. gossypinus, (fig. 15), Earias biplaga and Earias cupreoviridis as well as two (2) major stem-feeders, Hypolixus nubilosus and Alcidodes crassirostris. Ewete et al. (1980) showed that the problem of poor seed viability was solved in okra by timely harvesting of okra fruits when they are 28 to 35 days old on the plant and the seeds attain high germination (table 4). The okra fruits harvested at this stage showed ripening colour and splitting along the ridged fruit wall and with seeds already turned black but not yet dried. This tended to avoid excessive feeding by seed bugs (Dysdercus sp) which aggregate on the fruits (fig. 1) when allowed to remain completely dried on the plant before harvesting for seed production as practised by the local farmers. 24 UNIVERSITY OF IBADAN LIBRARY Table 4: Comparison of Treatment means on Okra Seed Germination (Summary of 1974/75 Experiments) Days after Mean seed germination (percent) in okra flower opening Early season Late season Early season 1974 1974 1975 14 o c 1.7 d 0 d 21 13.0 b 33.3 b 14.3 c 28 28.3 a 68.7 a 49.7 b 35 38.3 a 65.0 a 72.7 a 42 36.0 a 65.7 a 55.3 b 86 o c 16.0 c 0.3 d (Completely field-dried) Means followed by the same letters are not significantly different at P = 0.05. Fig. 14: Dysdercus superstitiosus, a major green and dry fruit-feeder. 25 UNIVERSITY OF IBADAN LIBRARY ( Fig. 15: Oxycarenus gossypinus, a major green and dry fruit-feeder. In another study on insect pests infesting the leafy vege- table, Amaranthus spp, the two major defoliators identified were Hymenia recurvalis and Psara bipunctalis while Cletus fuscescens and Aspavia annigera were the major grain or seed feeders. The defoliators infest Amaranthus from 3-4 weeks after planting (WAP) and skeletonises the leaves at 4-5 WAP as the leaves are ready for harvesting and consumption. What a right time to strike and readily deprive the farmers of their harvest! In our study on the survival of C. fuscescens on five species of Amaranthus, the wild Amaranthus (A. spinosus) is as good as four other cultivated amaranths (A. cruentus (grain), A. cruentus (leafy), A. dubius (leafy), A. hypo- chondriacus (grain))· in supporting the biology and high survival of Cletus fuscescens (table 5), and hence acting as a major link in their infestation (Ewete and Ukwela 1991). Thus a first step in the management of this pest is to destroy suspected wild alternative host plants around the farm during its cultivation. 26 UNIVERSITY OF IBADAN LIBRARY Table 5: Nymphal Survival (%) of Cletus fuscescens on Five Species of'Amaranthus Nymphal instars Food sources I II III IV V Mean nymphal survival Amaranthus hypochondriacus (grain) 88.9 90.0 80.6 86.2 84.0 85.9a Amaranthus spinosus (weed) 92.5 68.6 86.4 92.2 80.8 84.1 Amaranthus dubius (leafy) 92.3 83.3 70.0 76.2 87.5 81.9a Amaranthus cruentus (leafy) 83.3 82.5 60.6 75.0 93.3 78.9a Amaranthus cruentus (grain) 76.1 68.6 91.7 68.2 80.0 96.9a Means followed by the same letters are not significantly different at 5% level. 27 UNIVERSITY OF IBADAN LIBRARY Insect Pests of Fibre Crops The well known fibre crops are cotton, kenaf, roselle, sisal, et cetera. Cotton is grown for its lint used in sericulture and cotton seed for its oil and cake for livestock feed. The bast fibres of kenaf and roselle are used in making jute bags and ropes. However, cotton, kenaf and roselle belong to the family Malvaceae as okra. They are infested by similar insect pests, the seed bugs, Dysdercus spp and Oxycarenus spp, inclusive. Extensive research work had been conducted on Dysdercus spp (Cotton-stainer) infesting cotton and okra, while Oxycarenus spp was neglected. This prompted my doctoral study on the ecology and biology of Oxycarenus gossypinus and the damage caused on okra as well as on the alternative host plants, cotton, kenaf and roselle. On its biology, Ewete and Osisanya (1988) showed that the egg incubation period was 6.98 days and the mean developmental period (first instar - adult) was 16.7 days, and there were 5 nymphal instars whose morphometric measurements and immature stages were also reported (Ewete 1984) (fig. 16). Ewete and Osisanya (1985) also showed that the nymphal developmental period on four food sources (seeds) - okra, kenaf, roselle and cotton averaged 15.5, 15.7, 15.8 and 18.1 days, respectively (table 6). This implied that okra, kenaf and roselle should not be grown in the vicinity of cotton to avoid early infestation that will reduce cotton yield. 28 UNIVERSITY OF IBADAN LIBRARY ~ I·Om., Fig. 16: Eggs and larval instars of Oxycarenus gossypinus distant. 29 UNIVERSITY OF IBADAN LIBRARY Table 6: Mean Duration (days ± SE) of Nymphal Development in O. gossypinus reared on Dry Seeds of four cultivated Malvaceae (range in parenthesis) Nymphal instars Mean period* Food source of nymphal First Second Third Fourth Fifth development. Okra 3.27±O.18 3.80±O.26 2.0±O.O 2.40±O.13 4.0±O.O 15.47±O.39b (2-4) (3-5) (2) (2-3) (4) (13-18) Cotton 4.0±O.O 3.67±O.28 3.0±O.O 2.30±O.15 5.17±O.3J 18.14±O.48" (4) (3-5) (3) (2-3) (4-6) (16-21) Kenaf 3.0±O.O 3.63±O.1l 2.63±O.11 2.14±O,10 3.74:tO.1O 15.74±O.23b (3) (3-4) (2-3) (3-4) (3·4) (13-17) Roselle 3.13±O.ll 3.0±O.O 3.0±O.O 2.83±O.O8 3.86±O.O7 15.S2±O.lSb (2-4) (3) (3) (2-3) (3-4) (13-17) Means followed by the same letters are not significantly different at 5% level *Mean periods of nymphal development are significantly different (F = 3.18; PI! 0.05). 30 UNIVERSITY OF IBADAN LIBRARY Stored Product Insects Storage of agricultural products forms an integral part of farming practice whereby farmers produce in excess of family needs and the rest could be stored for various reasons. The reasons for storage include the continued sustenance of life or for future planting or until the produce commands higher price, or against a period of any disaster (drought, flood, war or insurgency as in Boko Haram). Such a storage condition provides an excellent condition for a field-to-store insect developing into a pest status. This is particularly true of insects infesting stored cereals (maize, rice, guinea com, wheat) such as Sitophilus zeamais, S. oryzae, S. granarium; stored pulses (cowpea, pigeon pea, soya bean) such as Callosobruchus maculatus; stored animal produce (smoked fish, stockfish, leather) such as Dermestes maculatus; stored cocoa beans such as Ephestia cautella; stored Cola nut such as Balanogastris kolae (Ivbijaro 1975); Sophrhorhinus insperatus, S. gbanjaensis (Daramola 1973) to mention a few. The control of this group of insect pests has been centred on the application of fumigants such as methyl bromide, phos- phine gas marketed as phostoxin or quickphos, and actellic dust (pirimiphos-methyl). Effective as these synthetic insecticides are, they inflict side-effects on humans and the environment through the depletion of the ozone layer, and consequently their ban. This has prompted the search for alternatives with the searchlight beamed at investigating the use of plant natural products or insecticides of plant origin. Isman (1994) highlighted five botanical insecticides registered for use in USA namely; Pyrethrum from Chrysanthemum cinerariaefolium (flower); rotenone from Derris elliptica and Lonchocarpus sp. (roots and tubers); ryania from tropical shrub, Ryania speciosa (stem wood); Sabadilla from South American Veratrum (seeds) and azadirachtin from neem, Azadirachta indica (seed). My interest in the use of natural product insecticides in controlling insects was1foused during my one-year UNIVERSITY OF IBADAN LIBRARY sabbatical leave in 1993 at the Department of Biology, University of Ottawa, Ottawa, Ontario, Canada, with two scientists, Professors J.T. Amason (botanist) and BJ.R. Philogene (entomologist) as my hosts under the 1992 Canadian Fellowship of CIDAINSERC. Similarly, I was a visiting scientist in 1997 at the Scottish Agricultural College, University of Edinburgh, Edinburgh, Scotland with Dr. Andy Evans as my host under the 1996 Royal Society Third World Fellowship to test the efficacy of the stem-bark extracts of five Khaya species against the cabbage defoliator. In a study on the biological activities of four traditionally used Nigerian plants, Ewete et al. (1996) showed that ethanolic extracts of Piper guineense (Piperaceae), Cedrela odorata (Meliaceae), Dennettia tripetala (Annonaceae) and Aframomum melegueta (Zingiberaceae) in artificial diets significantly reduced larval growth of European com borer (ECB), Ostrinia nubilalis, at a concentration of 1000ppm (0.1%) (table 7). It is noted that O. nubilalis exhibits the same symptom (window pane) as Busseola fusca on maize leaf in the tropics. Further tests on the nutritional indices for habituated 3rd instar larvae with the two most promising plant extracts indicated that P. guineense and C. odorata extracts showed the best potential for development as botanical insecticide. The active compounds, piperine from Piper and gedunin from C. odorata were compared for efficacy with their respective crude extracts against ECB. It was shown that piperine had no marked effect at reducing maximum larval weight but caused larval mortality compared with the crude extract, but gedunin, C. odorata crude extract prolonged larval development and adult emergence of ECB (Ewete et al. 2000) (table 8). This implied that certain compounds exist within the crude extract of Piper which are acting as synergist(s) and this may have accounted for the efficacy. 32 UNIVERSITY OF IBADAN LIBRARY Table 7:The Effects of Plant Extracts on Larval Growth ot O. nubilalis Plant species Weight gain (s.d.) of second instar larva (mg) Concentration 0 10ppm 100 ppm 1000 ppm 0 10ppm 100 ppm 1000 ppm Piper guineense 3.08a 3.85a 3.08a 0.96b (1.2) (1.0) (1.6) (0.3) Cedrella odorata 4.00a 3.60ab 3.21b o.sse (1.0) (1.0) (1.2) (0.3) Xylopia aethiopica 3.78a 3.69a 3.87a 3.84a (1.6) (1.0) (1.6) (1.2) Dennetia tripe tala 5.79a 3.13b 3.28b 2.21b (1.1) (1.4) (1.3) (1.0) Aframomum melegueta 4.72a 4.95a 4.58a 2.76b (1.5) (1.3) (1.5) (0.9) Means followed by the letter in a row are not significantly different in the SNK test (P = 0.05) Table 8:Effects of Gedunin and Piperine on Larval Weight (mg) and Mortality of Ostrinia nubilalis (s.d. in parenthesis) Larval mortality (%) Larval wt. at day 12 at day 8 (% control) Concentration Gedunin Piperine Gedunin Piperine Oppm 6.7 3.3 l00±23.1 lOO±15.7 5ppm 6.7 0 46.0±11.3 79.3±14.1 lOppm 3.3 3.3 45.7±14.0 112.5±13.4 20ppm 3.3 3.3 43.9±15.2 148.5±14.8 40ppm 6.7 23.3 43.3±16.2 103.6±17.1 In another study on the insecticidal activity of the extract of an Asian shrub, Aglaia odorata (Meliaceae), and the purified active principle, rocaglamide, from same plant, Ewete et al. (1996) showed that the extract was a potent inhibitor of larval growth of ECB at dietary concentrations 12.5-lOOppm (fig. 17). The dietary concentrations of 0.1 and 0.2ppm of rocaglamide strongly inhibited larval growth and survival of ECB (table 9, fig. 18). Rocaglamide was more potent than azadirachtin which had earlier been reported to be the most potent natural product ever tested by Amason et al. (1985). 33 UNIVERSITY OF IBADAN LIBRARY - 10.rs;n ::E O'l '~ ~ ~~ ~ Ii:: ("IS c:p E OJ .0 0-1 ..) 5 10 15 20 25 30 35 Larval age (days) --0-- control; -.0-- 12_5 pprn; --0- 25ppm; A 50 pprn; m 100 PPFtl Fig. 17: Effect of dietary administration of A odorata extract on larval growth of O. nubilalis. 34 UNIVERSITY OF IBADAN LIBRARY Table 9: Mean Weight gained and total Mortality of O. nubilalis Larvae exposed to Dietary Concentrations of Rocaglamide over a 15 day period Rocaglamide in diet Mean weight gain Total mortality (ppm) (mg) (%) 0.0 69.71±3.40 a,b 7.4 n=24 0.0125 76.63±5.84 a 13.3 n=25 0.025 84.72±4.78 a 16.6 n=22 0.05 58.46±3.99 b 43.3 n= 19 0.1 31.94±3.16 c 60.0 n=24 0.2 5.3±1.l5 d 85.0 n= 13 Means are ± S.E.M.; means followed by the same letter do not differ significantly using a Student-Neuman-Keus (SNK) test at P 0 0.05. 35 UNIVERSITY OF IBADAN LIBRARY ao,---------~------------------~ 60 C-»E1: .f;i!! 40 t: :I:; 20 o~~~~==~~~----__----~ o 5 10 15 20 Larval age (days) Ia p onnltol; &I O.05ppmj --0- 0.1 ppm: o O.2ppm: Fig. IS-:·Effect of dietary administration of rocaglamide on larval growth of O. nubilalis. With these positive, results- on) the, biolngicalt alldRiWafrRUmt natural: n~.~ wc.m: amnlIIIIttIdIronttllel.Uleaftn1ant· extracns ttDuw'lttdhttDodigJlDilmttiitr.iW.tl>'" ~ am altbrllliltWe tro synttllllt~ ffimigarrts.. Iln atm· flrst' test on the bioactiv.i~