ELEVENTH DR. S. PRADHAN MEMORIAL LECTURE September 9, 2019
Predaceous Ladybirds: A Reproductive Perspective
By
Prof. Omkar Professor Omkar received his graduation, post graduation and Doctorate degrees from DDU, Gorakhpur University. He continued his Post doctoral research at his alma mater. After a brief stint Ladybird Research Laboratory, Department of Zoology, at Central Tasar Research and Training Institute , Central Silk Board at Ranchi he began his teaching University of Lucknow, Lucknow-226 007 (India) career at the Department of Zoology, University of Lucknow from December 1986 and took over the charge of Head, Department of Zoology, University of Lucknow (2016-2019). He heads the only Ladybird Research laboratory in India. His research on biology, ecology and behaviour of ladybird beetles have resulted in landmark contributions on ladybird predation, life history traits, reproductive attributes and optimization of rearing factors with over 36 pioneering studies spanning, multiple matings, promiscuity, optimal number of matings, costs and benefits of reproduction, ageing, developmental rate polymorphism, sexual selection and mate familiarity all of which have helped in standardizing mass multiplication of ladybirds. He was awarded a grant under Centre of Excellence in Biocontrol of Insects Pests, Govt. of Uttar Pradesh, and is part of IUCN in the Ladybird Species Specialist Commission besides running various research projects funded by state and central funding agencies. He has visited School of Biological Sciences, University of East Anglia, Norwich, UK (2006) and Czech Academy of Sciences, Ceske Budejovice, Czech Republic (2016) under Royal Society and Czech Academy of Sciences-INSA Bilateral Exchange of Scientists programme, respectively. He has supervised 23 Ph. D. students and has 270 articles in reputed international and national journals to his credit. He has authored 11 books, three of which have been published by Academic Press, Elsevier Inc. USA and Springer Nature, Singapore. Prof. Omkar has contributed significally as Member, Chairman of many academic and administrative bodies at University of Lucknow and other universities; Coordinator, UGC-SAP (DRS-II), DST (FIST), and DST-PURSE, University of Lucknow and UGC nominated member of advisory committee for UGC-SAP (DRS–II) at Panjabi University, Patiala; Member of Selection Boards of multiple universities and state commissions as subject expert and examiner of more than 40 universities in 14 states of our country; Chief Editor of Journal of Applied Bioscience, Associate Editor of International Journal of Tropical Insect Science (Springer Nature), and referee of several international and national journals and funding agencies. DIVISION OF ENTOMOLOGY Recipient of several awards among which Saraswati Samman (2017), Govt. of UP, Prof. TN Ananthakrishnan Foundation Award, Chennai, (2012) are noteworthy. Also an elected Fellow of ICAR-INDIAN AGRICULTURAL RESEARCH INSTITUTE National Academy of Sciences, India, (FNASc), Zoological Society of India, The Entomological Society of India, etc. and President, International Society of Applied Biology, Lucknow, NEW DELHI & ENTOMOLOGICAL SOCIETY OF INDIA Organizing Committee
PATRON Dr. A.K. Singh APEX LIFE SCIENCES DDG ( Extension ) ICAR & Director ICAR-IARI
CONVENER Dr. Rakesh Sharma Head, Division of Entomology
MEMBERS Dr. H.R. Sardana Director , NCIPM
Dr Surinder Kumar Principal Scientist, NCIPM
Prof. Subhash Chander Add:- 204, Magnum House-I, 2nd Floor, Jt. Secretary, Entomological Society of India Commercial Complex, Karam Pura, New Delhi-110015 Dr J.P. Singh PAN: AEAPG7108R, GSTIN: 07AEAPG7108R1Z9 General Secretary, Entomological Society of India AN ISO 9001:2015 CERTIFIED COMPANY Tel:- 011-49423591, Mob:- 9212544059, 9868163134 Dr. Debjani Dey Email:- [email protected] Principal Scientist
Dr. Kirti Sharma Principal Scientist
Dr. Sachin Suroshe Principal Scientist M/S. Allenovate (STAND FOR YOUR ALL KIND OF RESEARCH) Dr. Suresh Nebapure Specialized in;- NGS Sequencing ,FAME Analysis, PLFA Analysis, Identification of Scientist microbes,GC-MS, LC-MS etc. Dr. Sagar, D. Also deals with;-- All laboratories chemicals. Consumables, glassware and plastic ware Scientist Contact Person:- Kundan Kumar(Ph.D.MBA,M.SC.,PGCAP) Shri Sushil Kumar Mob:-9311493935, 8130724526 AAO (Member Secretary) E-mail:[email protected], [email protected]
3033AB, 1st FLOOR, GALI NO -21 RANJEET NAGAR, NEW DELHI-110008 E-mail:[email protected], [email protected] ContactsUs : 9311493935, 8130724526 ELEVENTH DR. S. PRADHAN MEMORIAL LECTURE September 9, 2019
Predaceous Ladybirds: A Reproductive Perspective
By
Prof. Omkar Ladybird Research Laboratory, Department of Zoology, University of Lucknow, Lucknow-226 007 (India)
DIVISION OF ENTOMOLOGY ICAR-INDIAN AGRICULTURAL RESEARCH INSTITUTE NEW DELHI & ENTOMOLOGICAL SOCIETY OF INDIA
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Management: Potential and Limitations
2010 Dr. Hari C. Sharma Applications of Biotechnology for Insect Pest
Previous Dr. S. Pradhan Memorial Lectures Benefits
Analyses of their Global Adoption, Safety and
1998 Dr. K.N. Mehrotra Sustainable Plant Protection: Science and Technology
Dr. T.M. Manjunath A Decade of Commercialized Transgenic Crops : 2005 Dimensions Previous Dr. S. Pradhan Memorial Lectures
1999 Dr. R.C. Saxena Neem for Ecological Pest and Vector Management
Regime and Environmental Conservation
1998 Dr. K.N. Mehrotra Sustainable Plant Protection: Science and Technology Dr. S.N. Puri Challenges in Pest Management in New WTO
2003 2000 Dr. Ajai Mansingh DimensionsIntegrated Pest and Pesticide Management :
A Holistic Approach and Biosecurity
1999 Dr. R.C. Saxena Neem for Ecological Pest and Vector Management Dr. M.B. Malipatil Importance of Systematics to Biologicqal Control 2001 2001 Dr. M.B. Malipatil Importance of Systematics to Biological Control
and andEnvironmental Biosecurity Conservation
A Holistic Approach 2003 Dr. S.N. Puri Challenges in Pest Management in New WTO
2000 Dr. Ajai Mansingh IntegratedRegime Pest and Pesticide Management : Dr. Ajai Mansingh Integrated Pest and Pesticide Management :
2000 A Holistic Approach
2005 Dr. T.M. Manjunath A Decade of Commercialized Transgenic Crops :
and Environmental Conservation Analyses of their Global Adoption, Safety and
2001 Dr. M.B. Malipatil ImportanceBenefits of Systematics to Biologicqal Control Dr. R.C. Saxena Neem for Ecological Pest and Vector Management
1999 and Biosecurity
2010 Dr. Hari C. Sharma Applications of Biotechnology for Insect Pest
Dimensions Management: Potential and Limitations
2003 Dr. S.N. Puri Challenges in Pest Management in New WTO Dr. K.N. Mehrotra Sustainable Plant Protection: Science and Technology
1998 2015 Dr. J. S. Bentur RegimeGall Midge Resistance in Rice : An Update
2016 Dr. N. K. Krishna Kumar Sap Sucking Pests and Vectors: Similarities and 2005 Dr. T.M.Previous Manjunath Dr. S.A Pradhan DecadeDissimilarities of MemorialCommercialized in Management Lectures Transgenic Crops : 2018 Dr. Chandish R. Ballal Analyses Challenges of totheir and OpportunitiesGlobal Adoption, for Biological Safety and BenefitsControl Practices in India
2010 Dr. Hari C. Sharma Applications of Biotechnology for Insect Pest Management: Potential and Limitations
Printed by: New United Process, New Delhi-110028, 9811426024
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28 Dr. S. Pradhan May 13, 1913 - February 6, 1973
3 4 Dr. S. Pradhan - A Profile
Dr. S. Pradhan, a doyen among entomologists, during his 33 years of professional career made such an impact on entomological research and teaching that Entomology and Plant Protection Science in India came to the forefront of agricultural research. His success story would continue to enthuse Plant Protection Scientists of the country for generations to come.
The Beginning Shyam Sunder Lal Pradhan had a humble beginning. He was born on May 13, 1913, at village Dihwa in Bahraich district of Uttar Pradesh. He came from a middle class family. His father, Shri Gur Prasad Pradhan, was a village level officer of the state Government having five sons and three daughters.
After initial studies at a nearby village school, Pradhan completed his primary education from Lucknow and passed High School Examination of U.P. Board from Govt. High School, Bahraich, in 1928, with distinction in Mathematics. He then moved over to Radhaswamy Educational Institute at Agra from where he passed Intermediate Examination of U.P. Board in 1930, in first Division and was awarded a Gold Medal. Thereafter, he joined Lucknow University, where he topped the University in B.Sc. degree examination in 1932 and obtained his M.Sc. degree in 1934. He joined as Research Scholar in the same University under the able guidance of the eminent Zoologist, Prof. K.N. Bahl. He worked on functional morphology of insects for which he was awarded D.Sc. degree in 1938 and his thesis was adjudged the best among those submitted to Lucknow University during the year. Later, in 1948, he was also awarded Ph.D. degree from the same university for his research in insect toxicology carried out at Rothamsted Experimental Station, United Kingdom.
The Career After completing his D.Sc. degree, Dr. Pradhan initially worked on sugarcane pests for a brief period under a scheme of Indian (then Imperial) Council of Agricultural Research at Gorakhpur, U.P. Thereafter, he joined Indian (then Imperial) Agricultural Research Institute in 1940 as Assistant Entomologist at the Institute’s sub-station at Karnal (Haryana), where he did his famous work on insect ecology. In 1946, Government of India sent Dr. Pradhan to Rothamsted Experimental Station, U.K. for training in insect toxicology. After his return from U.K., he established the country’s first school of insect toxicology, in 1948, at the Division of Entomology, IARI, New Delhi. With the creation of Post-Graduate School at the Institute, Dr. Pradhan became the first Professor of Entomology in 1958. He became Head of the Division in 1962 and continued till February 5, 1973 when he left the mortal world, a few months before his scheduled retirement from service.
Contributions to Science and Society Dr. Pradhan was a visionary and scientist par excellence. Having versatile interests, he contributed significantly to various aspects of entomological research, teaching, scientific documentation and popularisation of Entomology and Plant Protection. His numerous contributions in fundamental and applied aspects are not feasible to be enlisted in single article. However, an attempt has been made to bring forth some of his most significant contributions in various spheres of entomological research.
5 The Researcher and Innovator Dr. Pradhan’s research interest ranged from fundamental to applied aspects of Entomology. He was the first to identify and describe “gnathal glands” in coleopterans. His earlier work on morphology of insects led to the understanding of the homology of male genitalia, regeneration of mid-gut epithelium and function of Malpighian tubules in coccinellid beetles. His work on coiling and uncoiling mechanism of proboscis in Lepidoptera was yet another important finding to fill a lacuna in the then existing knowledge. These studies still find a place in standard books of Entomology.
Having deep understanding of insect-crop-environment interactions, keen observation and proficiency in mathematical derivations, Dr. Pradhan was an insect ecologist whose class was not only evident in this field but his work in various other fields of applied Entomology also bore the stamp of an accomplished ecologist. His contributions on effect of abiotic and biotic factors on insect life, population dynamics of pests, assessment and sampling of pest damage and crop losses are highly significant.
His classical work on effect of temperature on the dynamics of development of any cold blooded organism brought him immediate world recognition. He gave 2 equations depicting relationship between temperature and development at constant and variable temperatures, which were later called as Pradhan’s equations,
-ax2 1. For constant temperature: Y = Y0e
where,
Y0 = highest developmental index,
Y = Developmental index at required temperature t,
x = T-t (T being temperature at Y0), and e is a constant (2.718282).
2. For variable temperature: Y (t1t2) =
where,
Y (t1-t2) = Average value of developmental index corresponding to temperature fluctuation
between t1 (max.) and t2 (min.) at which the values of x are x1 and x2, respectively.
The validity of these equations have been verified in several laboratories world over. Among the various temperature - development relationship equations evolved by different workers, ‘Pradhan equations’ have been found best and provided most accurate relationship. These equations evoked so much interest globally among leading development biologists that several full papers were published on his innovation. The deduction of these equations provide a glimpse of Dr. Pradhan’s mathematical acumen and insight of biophysical aspects of growth and development, a rare combination of abilities in a biologist.
Based on these equations Dr. Pradhan developed ‘Biometer’, a ready reckoner for reading the insect development at any temperature and for estimating the amount of development or the number pf
6 generations in any given period and under any range of temperature fluctuations by counting the number of certain cells. He tested the hypothesis by preparing biometers of several important crop pests.
Dr. Pradhan was a pioneer in insect toxicology research. One of his most important contributions was on temperature-toxicity relationship, where he showed that the susceptibility of insect to insecticides depended on the combined effect of temperature prevailing before, during and after the treatment. He was the first to demonstrate that insect response to DDT varies with post-treatment temperature. Dr. Pradhan and his illustrious student, late Dr. Ranga Rao proposed a physical theory on the mode of action of DDT based on electrophysiological evidences in 1966.
His contributions to fundamental aspects of insect toxicology included relationship between particle size of suspensions and insect mortality; the role of cement and wax layers of insect cuticle in penetration of insecticides and suitability of various diluents in insecticide formulations. He was the first to report insecticide resistance from the country and developed a strong section for work on this aspect. He demonstrated that the resistance to insecticides had two components in external (resistance to the entry of poison into insect system) and internal (resistance to the poison that has gained entry into insect system) and that these 2 components of resistance differ markedly in different species. A regular bioassay of relative toxicity of insecticides to different pest species was initiated under his leadership in order to (a) screen out most effective insecticides against different pests and (b) to detect development of resistance to different insecticides in various pests.
Dr. Pradhan evolved effective chemical control measures against various crop pests through bioassay and field trials. His ecological leanings were evident even in this field where the chemical control recommendations were directed against weaker spots in pest life cycle, timed suitably and adapted in a manner which was safe to non-target organisms and natural enemies. His critical analyses of various key pest problems of different crops for effective pest control strategies can be gauged by the biotic circuit and chronological strategy of pest control based on ecological timing and integration’ of different control strategies against sugarcane pests.
Under his leadership, work on insecticide residues was started in the Division using bioassay technique. Dr. Pradhan was one of the earliest proponents of the concept of integrated pest management (IPM). His address at the International Seminar on Integrated Pest Control, organised by him in 1969, provides a lucid account of the philosophy and feasibility of IPM strategies for different crops. His concept of double screening of natural enemies due to direct toxic effects of insecticides and indirect stress from reduced host density was a revealing analogy of IPM.
His approach to solve problems associated with grain storage on the basis of ecological requirements of grain and different stored grain pests is an essential reading material for everyone interested in storage pest management. Based on this analysis Dr. Pradhan and coworkers developed a cheap, scientifically fabricated and effective grain storage structure named as Pusa Bin and later its larger version, Pusa Cubicle.
Dr. Pradhan was the pioneer in ‘neem research’. He demonstrated the strong insect antifeedant activity of neem kernel suspension during early sixties, when botanicals were given a low priority. It has revitalised
7 the entire research work on neem leading to purification of bitter principles. He also demonstrated its success in field trials and advocated its use by farmers in protecting their crops during locust invasions.
He propounded the famous Biotic theory on the periodicity of locust cycles which explains the distribution and abundance of locusts both in time and space, and showed the way for the locust cycles to be nipped in the bud. He presented a paper ‘Biological control of Acridid Pests’ at the International Conference on Current and future problems in Acridology at London in 1970.
Under his leadership, studies on host plant resistance were intensified during sixties and a number of good sources of resistance to key pests of sorghum and maize were identified and later used for breeding pest resistant varieties. Also the pioneering work on evaluating world germplasm of wheat for their reaction to major storage pests was undertaken by Dr. Pradhan with co-workers which showed that host grain resistance could be used for alleviating post harvest losses in this important cereal.
The Educator Dr. Pradhan was a gifted teacher and those whom he taught, many of them becoming eminent entomologists themselves, remember the lucidity and clarity of his lectures. Dr. Pradhan guided the research work of 65 students (45 for diploma of Associate, IARI; 1 for M.Sc. and 19 for Ph.D. degrees) in the discipline of Entomology. His popularity as a teacher could be illustrated from the fact that besides his specialised lectures in fields of ecology, toxicology and pest management, the basic course of Introductory Entomology wherein he taught dominance of insects attracted students of Entomology and other disciplines. Being the first Professor of Entomology, Dr. Pradhan drew up the curricula for M.Sc. and Ph.D. courses of the Institute.
The Visionary and Crusader Dr. Pradhan was first to visualise the country’s need for integrated pest management instead of the conventional method of chemical control alone, which though successful at the time had started showing adverse effects. Also he emphasized that the crop protection research should have its rightful place among overall agricultural research efforts of the/country.
In his later years, he published several papers and delivered talks at different fora to bring forward the causes of accentuation of pests and need for intensification in crop protection research. His last published paper In tropics protection research more needed than production research is a treatise on crop protection component of overall crop production scenario.
His forceful arguments for adopting Integrated Pest Management in order to fully realise the production potential of new high yielding strains of different crops, had their impact and the country adopted IPM as a national strategy, though after his demise.
The Writer Dr. Pradhan’s scientific papers as well as popular articles all bore his characteristic clarity of thought and acumen of presenting complex subjects in simple form. Dr. Pradhan published about 200 research papers in various Indian and foreign journals, besides several popular articles in Hindi and English. He was
8 almost a regular contributor in annual meetings of Indian Science Congress. His book on Insect Pests of Crops, published by National Book Trust in 1969, and Agricultural Entomology and Pest Control, which was published by Indian Council of Agricultural Research in 1983, would always remain valuable to agricultural scientists. His chapter on Entomology: New Dimensions of Pest Control in Plant Protection Section of the Agricultural Year book of ICAR entitled New vistas in crop yield provides an exhaustive view of needs and feasibilities of managing agricultural pest problems of the country. He also wrote a chapter on Ecology of arid zone insects excluding locusts and grasshoppers in Human and Animal Ecology published by UNESCO.
The Vanguard Besides heading the Division of Entomology of the premier national Institute for more than a decade, Dr. Pradhan was also President of Entomological Society of India for four terms. The two positions and his towering personality made him the natural leader of the country’s crop protection specialists. Dr. Pradhan was a founder member of Entomological Society of India. The Society took great strides under his stewardship. The membership of the Society increased and the Indian Journal of Entomology grew in circulation and stature. The Society celebrated its Silver Jubilee in 1964 and a National Seminar was organised under his patronage. Again in 1969, he organised the International Seminar on Integrated Pest Control as President of the Society. Dr. Pradhan was Chairman of Entomology Committee of ICAR for a number of years and guided the planning and implementation of national policies on entomological research, teaching and extension.
Recognitions Dr. Pradhan was regarded as an authority on insect ecology, toxicology and integrated pest control, not only in India but internationally. In one such recognition, he was invited by UNESCO as one of the selected six authorities to write different chapters in the book on Human and Animal Ecology. He felt that since the invitation was because of his work at IARI, the honorarium received for writing the review article was used by him to create an endowment for award of a gold medal every year to the most outstanding post-graduate student of Entomology at IARI in the name of his father Shri Gur Prasad Pradhan. Dr. Pradhan was invited to present his Biotic theory of locust cycles to a select gathering of acridologists and locust control workers at Porton, U.K. in 1970, He was a member of FAO panel of experts on integrated pest management. Dr. Pradhan chaired three important sessions in the 14th International Congress of Entomology, held at Canberra, Australia,, in August 1972. Soon after he was invited to Hawaii to advise on the preparation of syllabus for a special course on integrated pest control. Dr. Pradhan was posthumously awarded Dr. P.B. Sarkar Endowment Prize for the triennium 1971-74 for outstanding research contribution that lead to enhanced food production in India.
Indian scientists would always remain inspired by Dr. Pradhan’s work, who was rightly called ‘Father of Modern Applied Entomology’ by Late Prof. K.N. Mehrotra, Ex-Head, Division of Entomology, ICAR- IARI.
9 10 Predaceous Ladybirds: A Reproductive Perspective
Omkar
Ladybird Research Laboratory, Department of Zoology, University of Lucknow, Lucknow-226 007 (India) [email protected]
Introduction Family Coccinellidae of Coleoptera is an ancient and successful group of insects that evolved in the lower Permian period, about 280 million years ago. The members of this group are commonly termed as Ladybird beetles, Ladybeetles or Ladybirds. The “Lady” part of their name was initially given to the cosmopolitan ladybird beetle, Coccinella septempunctata and comes from the Virgin Mary, for the resemblance of the scarlet colour of elytra to the cloak in Renaissance era paintings of her, seven spots reflective of the seven joys and sorrows of her life, and the belief that they bring good luck and bountiful harvests (Plate 1). They are the most recognized and loved insects across the world (Plate-2). Ladybirds are most fascinating and various products of human use are designed in their shape. They are quite attractive to the children across the world and have always culturally been associated with good luck charms.
Not only are ladybirds traditionally believed to be good luck symbols but scientifically too, the majority are considered as potential biological control agents owing to their predaceous nature against numerous agricultural and horticultural pests. However, some members of family Coccinellidae belonging to subfamilies Epilachninae and Coccinelinae are phytophagous in nature and are harmful to crop plants as pests.
Life Cycle Ladybirds are holometabolous insects with their life cycle starting from the egg leading to four larval stages of increasing size and complexity. The final larval stage pupates and metamorphoses into an adult (Plate 3). Dixon (2000) grouped ladybirds on the basis of their dietary preference as aphidophagous or coccidophagous. The former have fast development and the latter have slow development which is
Plate 1: Coccinella sepetempunctata
Plate 1: Coccinella11 sepetempunctata
Plate 2: Different types of ladybirds
12 possibly adaptive in nature and in sync with the developmental durations of their prey. But both the groups have similar number of larval instars, except one coccidophagous species that has three larval instars rather than the usual four instars. The aphidophagous ladybirds generally lay eggs in clusters while coccidophagous ones lay eggs singly.
Plate 3: Life cycle of Coccinella septempunctata
Food of Ladybirds Plate 3: Life cycle of Coccinella septempunctata Food greatly influences the growth, development, survival, reproduction and offspring development and survial in ladybirds. The diet of predaceous ladybirds includes aphids, coccids, psyllids, diaspids, pentatomids, aleyrodids, and other insects and acarines. However, non-predaceous ladybirds feed on fungi, pollens, honey dew, etc. The dietary breadth in predaceous ladybirds is an outcome of the seasonal abundance and the synchrony of their potential prey. Moreover, attributes like morphology, chemistry and behaviour of the prey, the efforts involved in reaching the prey, the host plant architecture, and the level of threats or challenges imposed by intraguild predators and other natural enemies also affect the dietary breadth of predaceous ladybirds. While some ladybirds are stenophagic and have a narrow prey range, the others are euryphagic and depend on wide prey range. The former are further termed as specialists as they feed on monospecific or few prey species. However, the latter are termed as generalists or polyphagous, based on their broad dietary habits.
Numerous ways such as analyzing prey-predator coexistence (Mills,1981), gut or excreta of predators (e.g. Putman, 1964; Agarwala and Ghosh, 1988), serological essays (Hagley and Allen, 1990), radioisotope
13 labelling of prey (Southwood, 1966), ELISA testing (Crook and Sunderland, 1984), double antibody sandwich immunodot (Stuart and Greenstone, 1990), molecular gut content analysis using DNA half life (Greenstone et al., 2007), polymerase chain reaction of DNA of prey remains (McMillan et al., 2007), and chromatography (Sloggett et al., 2009) have been suggested for identifying the dietary range and suitability of ladybirds
However, the most reliable indicator of prey suitability is quantitative data on developmental and reproductive parameters of predators on being fed different prey (Hodek and Honek, 1996; Omkar and Srivastava, 2003a; Bind and Omkar, 2004; Omkar and Bind, 2004; Omkar and James, 2004a; Pervez and Omkar, 2004a; Omkar and Mishra, 2005a; Omkar et al., 2009; Bista et al., 2012; Bista and Omkar, 2013, 2014, 2016; Bista et al., 2017).
Differential responses have led to the classification of ladybird food, which continues to evolve with emerging information. The current classification has been a result of modifications by Hodek (1959, 1962, 1967), Hodek and Honek (1996) and Michaud (2005). Food is broadly grouped into accepted and rejected prey. The accepted prey is further subdivided into essential (supports development and reproduction) and alternative (supports survival not the development or reproduction). Michaud (2005) subdivided essential prey, into optimal (better than conspecific egg diet), adequate (at par with conspecific eggs) and marginal (poorer than conspecific eggs; development but considerable mortality). Another level of toxic prey was proposed which straddles the lower extreme of alternative and parts of rejected. The rejected category does not entirely fall under the ambit of toxic as sometimes the prey is either at inaccessible locations or simply too well defended to be consumed (Hawkes, 1920; Telenga and Bogunova, 1936; George, 1957) (Plate 4). Further, ladybirds accept some prey species which worsen their life-history parameters, although they are not toxic but they are considered as ‘problematic prey’. Moreover, the prey species selected by ovipositing females as food for their larvae are termed as ‘nursery prey’, and they are the species of prey on which the larvae are likely to develop the best in terms of survival and growth. This classification of food while being adequate and well formed is not rigid and absolute in nature.Food associations of ladybirds (Coleoptera: Coccinellidae) have been extensively studied because of their economic value as biocontrol agents.
Ladybirds have a tendency for prey specialization, which could be both diet- and habitat- related. The concept of prey specialization elucidates that ladybirds reared on suboptimal diets for few generations specialize and condition themselves for the suboptimal diet. These conditioned ladybirds perform better on a suboptimal diet than those on the optimal diets. Moreover, the switching of prey after few generations of rearing on either suboptimal or optimal diets causes deterioration in their performance and fitness (Rana et al., 2002). Further, the prey specialization in ladybirds has been argued as a function of their size, or the size and density of their prey. The body size of ladybirds provides an important trade-off determining their dietary breadth and the prey specialization (Chaudhary and Mishra, 2015a). Specialist ladybird species prefer the prey species that closely matches their size, have higher capture efficiencies and can easily reproduce at lower prey densities for longer duration of time. However, the generalist species adopt a one-size-fits-all strategy, which results in their lower capture efficiencies, making them difficult to sustain at lower prey density. Moreover, unlike the generalist species, whose adults frequently move between patchy prey habitats, the specialist species stay in the prey patches and lead a more sedentary
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Plate 4: Different prey of ladybirds life due to their greater tolerance of lower prey densities. They even start to reproduce earlier than the generalist species and also reproducePlate 4 in: Different the later stages prey of of declining ladybirds prey patch. Thus, their sedentary, stubborn and non-dispersing behaviour makes them the better biocontrol agents.
Predatory Responses The analysis of how a ladybird responds to varying pest populations and how it affects pest management can be understood in terms of the functional response. It is the predator’s feeding response against the increasing prey density, and can be analytically explained by Holling’s type I (linear), type II (curvilinear), and type III (sigmoidal) responses (Figure 1). Ladybirds usually exhibit a type II functional response,
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Figure 1: Holling’s Type I, II and III curves where there is an initial increase in the rate of prey consumption with increase in prey densities up to a certain level and this rate decreases with a further increase in prey density, i.e. the increase is not linear but rather curvilinear (Omkar and Srivastava, 2001, 2002, 2003b, c; Omkar and Pervez, 2004a; Pervez and Omkar, 2005; Omkar and Pervez, 2011; Pandey et al., 2014; Kumar et al., 2014). This happens due to satiation, as there is a threshold of prey consumption and the prey density dependent curve reaches an asymptote. The data obtained from this also helps calculate the attack rate and handling time of predator. Predators with high attack rate and less handling time are more likely to succeed at biocontrol.
Numerous factors affect the functional response outcomes of ladybird predators. These include, generation time ratio of predator and prey, prey preference (Omkar et al., 1997; Omkar and Bind, 1998), prey switching, size disparity between prey and predator, prey density, predatory stage, walking speed of predator and prey, gender, intrinsic rate of increase of prey and predator, consumption rate, prey patchiness, predator patch allocation time, host plant, abiotic factors, and intra- and interspecific predator competition. Moreover, the functional responses of ladybird predators are interlinked with their numerical responses. The numerical response of a predator is its tendency to increase its number with increasing prey density; and can be both aggregative and reproductive numerical responses. In response to increasing prey density, predaceous ladybirds show aggregative numerical response by increasing the cumulative prey consumption; however, the rate of prey consumption decreases curvili nearly due to mutual interference (Omkar and Pervez, 2004a; Pandey et al., 2014). The reproductive numerical response is a consequence of the functional response in predaceous ladybirds; because the females lay high number of eggs at higher prey densities, which is a direct implication of their functional response. A predator with a high tolerance for increasing predator density and increased reproductive capacity with increasing prey densities is likely to be more successful in biocontrol.
16 Effects of abiotic factors on life attributes of ladybirds Temperature is the most crucial abiotic factor affecting ecological, functional, and behavioural attributes of predaceous ladybirds. It sets the limits of biological activity in form of low and high temperature thresholds. The developmental rate is almost zero at lower development threshold, which increases with temperature, reaches a peak value, and then decreases rapidly as the high thermal threshold is achieved. Not much lower development threshold variation occurs in ladybirds with similar dietary habits and this is also a reason for their successful establishment in different habitats and countries. At the optimum temperature, around 25–30 °C in most aphidophagous and coccidophagous ladybirds, the peak of oviposition is attained at the youngest age of the ladybird demography(Omkar and Pervez, 2002; Pervez and Omkar, 2004b) (Figure 2). Thereafter, with further rises in temperature this peak is delayed and shortened. Development of immature stages is known to increase with increase in temperatures, but high temperatures are known to be detrimental to reproduction and survival (Srivastava and Omkar, 2003; Omkar and Pervez, 2004b; Omkar and James, 2004b; Pervez and Omkar, 2004b; Omkar and Kumar, 2016). While in aphidophagous and coccidophagous ladybirds optimal temperature lies between 25-30°C, in acarophagous ladybirds it is 30–35 °C, resulting in high values for the demographic parameters, like the net reproductive rate and the intrinsic rate of increase in population for the latter.
Not only temperature, the light also affects the life attributes of ladybirds. Various variables of light, like intensity, quality (wave length), and duration of exposure (photoperiod) significantly affect the development, reproduction and progeny fitness (Mishra and Omkar, 2005; Omkar et al., 2005; Omkar and Pathak, 2006) (Figure 3). Ladybirds have a wide range of tolerance limits to these variables. They
Figure 2:Age specific fecundity curves of P.dissecta at different temperature
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0.75
0.7
0.65
0.6
0.55
0.5 Individual Fitness 0.45
0.4 Long day Continuous day Short day
Photoperiods Figure 3: Influence of photoperiods on individual fitness levels in C. saucia are primarily diurnalFigure insects 3: Influence and depend of photoperiods on visual cues on individual and presence/absence fitness levels inof C.light saucia to undergo. various essential activities, like mating, moulting, and pupation. Most ladybirds are highly sensitive to light, particularly its photoperiod and wavelength. Short day lengths with intensities of 1500 lux are beneficial for the reproductive activities. The likelihood of female ladybirds accepting the males increases in the dark because females are unable to evaluate visual criteria to select male and thereby mating rejection displays are also minimized. However, prolonged light days could have a negative effect on the physiology of ladybirds.
There exists a photoperiod-dependent bimodal or two-peak pattern in the development of certain ladybirds, where the first peak represents fast developing and the second shows slow-developing individuals in the same cohort of eggs (Mishra and Omkar, 2012; Singh et al., 2014, 2016a, b). The slow-developing individuals are generally more in numbers in short day lengths; however, long day lengths promote fast developing adults having heavier body masses and more capable of producing quantitative progeny (Singh et al., 2016b). Selection over generations of these differentially reproducing adults in response to various abiotic and biotic factors can enhance these developmental as well as associated predatory responses (Siddiqui et al., 2015a, b; 2017).
Cannibalism and Intraguild predation in ladybirds
A guild is formed by the association of predators that share a common food resource. However, when the ladybirds exploit a common food resource (extraguild prey) within a guild, they often attack each other. In this struggle, one becomes dominant (intraguild predator) and overpowers the other (intraguild prey) (Plate 5). This exploitative competition of food resource is more advantageous to small-sized ladybirds as they have lesser food demands. As the density of extraguild prey decreases or the density of intraguild prey increases, the frequency of intraguild predation increases. On the contrary, an increase in extraguild
18 Plate 5: Interaction between aphidophagous ladybirds within a guild prey density lowers the possibilities of intraguild predation. The relative size and stage, mobility of species, aggressive strategies, mandibular structure, degree of feeding and habitat specificity, defense strategies, and density of extraguild prey determine the outcome of intraguild predation (Omkar et al., 2004, 2005, 2006a; 2007; Pervez et al., 2005, 2006) .
Majority of the ladybirds attack, prey upon, and displace other members of their family in a limited food resource struggle. Invasion and establishment of aggressive species following displacement of indigenous ones may be an outcome of such interactions. The Harlequin ladybird, Harmonia axyridis (Pallas) is an invasive species that has a competitive advantage over the indigenous species, such as Coccinella septempunctata, Coleomegilla maculata DeGeer, Hippodamia variegata Goeze, and Adaliabi punctata L., due to its vast prey range, and higher predation and foraging potential. This invasive species frequently indulges in either interference competition or the apparent competition. Thus, it either interferes with other competitors or may compel the inferior indigenous species to become specialist predators of less preferred prey in nature (Pervez and Omkar, 2006).
Ladybirds also struggle with other group of insects outside the family for the common food resource. They usually co-occur with chrysopid (Neuroptera: Chrysopidae) and syrphid (Diptera: Syrphidae) larvae
19 and share the limited food resources (Omkar and Mishra, 2003, 2016). However, intraguild interactions between them are asymmetrical which could have positive, negative, or neutral impacts on pest biocontrol. The ladybird larvae, pupae, and adults all use chemical substances containing volatile hydrocarbons/ alkaloids to deter predators.
Ladybirds and Biocontrol of Pests The term biological control (=biocontrol) was introduced by Smith (1919) for the “topdown” action of natural enemies/biocontrol agents (viz., predators, parasitoids and pathogens) in maintaining the pest population density at a lower level than what may have occurred in their absence. Although several stories exist regarding the successful utilization of ladybirds as biocontrol agents, their use in biocontrol came into existence when the Vedaliabeetle, Rodolia cardinalis (Mulsant) was selected to control the population of scale insect, Icerya purchasi on citrus in California (USA) in the year 1889. Thereafter, numerous ladybird species were successfully used in the biocontrol of aphids, scale insects and mealybugs. The impact of ladybirds in terms of successful pest biocontrol is largely dependent on their voracity, prey specificity, intrinsic rate of increase, and the mean generation time ratio between prey and predator. Interestingly, the size of ladybirds attacking similar kind of prey does affect the biocontrol with the large sized ladybirds being the better biocontrol agents. However, if the prey type is different, the small sized ladybirds with specialization on that particular prey type are more promising biocontrol agents than the large but less specialized ladybirds.
A need for aphid biocontrol led to the introduction of 179 aphidophagous ladybird species in North America since 1900, but only 18 have successfully established. A few aphidophagous ladybirds took many years to establish. However, many established after accidental introduction, including Coccinella septempunctata, Harmonia axyridis, and Propylea quatuordecimpunctata (L.). The introduction and invasion of certain aphidophagous ladybird species has been implicated in the decline of some native species in the USA and elsewhere. A flightless form of Harmonia axyridis was also produced using a chemical mutagen followed by selective breeding. The innundative releases of such flightless adults against aphids on glasshouse cucumbers were highly successful. However, in huge agriculture fields they were not successful because of their impaired foraging on account of being flightless. However, these adults had lower reproductive fitness and had fewer offspring despite ovipositing for a longer period.
Aphidophagous ladybirds are generally not considered as better biocontrol agents largely due to significant differences in their intrinsic rates of increase and mean generation time ratios, although the relative development rates of aphidophagous ladybirds are lower than those of aphids. However, aphid biocontrol could be benefitted if the prey is targeted early in the season, i.e. prey suppression initiation could be done when the aphid colony is young. The coccidophagous ladybirds are also successful biocontrol agents of both coccids and diaspids. Chilocorus nigritus (Fab.) is a highly successful and effective generalist predator of numerous species of Diaspididae with equal effects on some species of Coccidae and Asterolecaniidae (Omkar and Pervez, 2003). The Indian ladybird, Cryptolaemus montrouzieri Mulsant, is also a generalist predator of various scales and mealybugs and has been commercially exploited in both classical and augmentative biocontrol programs.
Moreover, certain specialist ladybirds belonging to genus Stethorus are potential biocontrol agents of
20 tetranychid mites, especially at their high density. Similarly, the ladybird, Clitostethus oculatus (Blatchley) is credited for the biocontrol of whitefly, Aleurodicus dispersus (Russell) in Hawaii and India. While the specialists are better biocontrol agents than the generalists because of their selective feeding and persistence in the target prey habitats; the invasion of generalists in their resource space is an issue of serious concern as they become intraguild prey or they are forced to emigrate.
Reproductive aspects of ladybirds Reproduction in ladybirds is tuned in respond to the various environmental conditions in order to maximize the reproductive success. Following are the few aspects of reproduction which has been studied in ladybirds:
Sexual Maturation Sexual maturation is the development of reproductive organs such as ovaries in females and accessory glands in males. Maturation is generally correlated with age (Adam, 2000; Brent, 2010). In ladybirds, activation of the follicular tissue of testes starts during pupal stage (Hodek and Landa, 1971; Hodek and Honek, 1996). The rate of sexual maturation is directly related to availability of resources (Papaj, 2000). However, females may also rely on the environmental conditions for the better investment in the egg production and sexual behaviour (Aluja et al., 2000). In Propylea dissecta Mulsant, sexual maturation was age specific (Omkar and Pervez, 2005). The age of sexual maturation in male and female ladybird was recorded 7.12 and 9.33 days, respectively, which proved the protandrous nature of ladybird (Figure 4; Omkar and Pervez, 2005). Another study on P. dissecta by Shahid et al. (2016) revealed that the mating influences the ovarian development (Plate 6) and it is accelerated by the number of matings. Results also showed that the number of follicles in each ovariole increased with the age of virgin and mated females upto the age of 3 days (Plate 7).
Mating Behaviour Reproduction is one of the most important activities of an organism and the efforts involved in the progeny production are directly related to the organisms evolution and extinction. Mating behaviour is an important aspect of the reproduction which leads to the continuation of the generations. It includes
Figure 4: PercentFigure mating 4: Percent incidence mating incidence amongst amongst males males andand females females of different of different ages of pale ages of pale morph of P. dissecta.wmorph of P. dissecta.
21
Plate 6: Effect of mating status on the ovariole development in P. dissecta.
Plate 6: Effect of mating status on the ovariole development in P. dissecta. Plate 6: Effect of mating status on the ovariole development in P. dissecta.
Plate 7: Age specific ovariole development in unmated females ofP. dissecta. the sequence of processPlate 7: whichAge specific commence ovariole from development the courtship in unmated through females insemination of P. dissecta. to the termination of mating. Courtship has been viewed as a set of pre-copulatory signals by which a male seeks to gain access to the female, presumably by satisfying a set of female choice criteria (Alexander et al., 1997). Insemination andPlate sperm 7: Agetransfer specific are ovariole other developmentimportant processes in unmated enabling females of male P. dissecta. insects to fertilize eggs with their sperm (Choe and Crespi, 1997). In ladybirds, courtship includes six steps i.e. approach, watch, examine, embrace, mount and attempt (Omkar and Pervez, 2005) (Plate 8). Embrace is basically an appeasement act while examine helps to recognize the mate. Chemical cues play an important role to initiate the mate attraction.
Ladybirds generally possess two types of matings i.e. active and quiescent mating. Active matings include bouts and shakes and it occurs in large beetles (C. septempunctata. and Coccinnella transversalis Fab.) while quiescent matings occur in small beetles (P. dissecta and Menochilus sexmaculatus Fabricius).
22 Once the mating has been initiated, it can last for several minutes. Large beetles mate for shorter duration as compared to the small beetles (Omkar, 2004). In C. septempunctata the mating duration was recorded (69.20±0.99 minutes) (Bista et al., 2012) while in P. dissecta it was recorded 275.40±12.23 minutes (Omkar and Pervez, 2005) and in M. sexmaculatus it was 133±2.8 minutes (Bind, 2007).
Mating behaviour gets varied with multiple matings which is a widespread phenomenon (Jennions and Petri, 2000; Hosken and Stockley, 2003; Ronkainen et al., 2010). Both males and females are capable of mating more than once (Dunn et al., 2005; Diana et al., 2008). These matings could be with same partner (monogamous) or with different partners (polygamous) (Srivastava and Omkar, 2005a; Omkar and Mishra, 2005b, 2014; Haddrill et al., 2008; Wenninger and Hall, 2008). Though multiple matings occurs widely in insects, there are several costs and benefits associated with it. In terms of benefits, it was reported that polyandrous females have higher reproductive fitness than the monogamous females (Arnqvist and Nilsson, 2000; Simmons, 2005). In other words, matings provide direct benefits in terms of increased fecundity and fertility. In C. septempunctata effect of multiple matings was recorded on their behavioural patterns and reproductive attributes (Omkar and Srivastava, 2002: Bista et al., 2012) (Table 1). Another study on Cheilomenes sexmaculata and C. transversallis also revealed that the multiple matings increased the fecundity and the percent egg viability (Omkar, 2004) (Figure 5). While promiscuity enhances reproductive output significantly, the increase is more significant when clubbed with mate choice (Srivastava and Omkar, 2005a, b; Omkar and Mishra, 2005b, 2014). While there are many benefits associated with multiple matings and promiscuity, increase in number of matings has a detrimental effect on the longevity of males and females (Mishra and Omkar, 2006) (Figure 6). Since, there is the occurrence of multiple matings which are beneficial for reproductive success, there are optimal number of matings which are required for the maximum reproductive output. Study on C. sexmaculata and P. dissecta revealed that 95% of maximum fecundity was obtained after 13.25 and 12.95 matings in C. sexmaculata and P. dissecta, respectively; and 8.95 and 11.25 matings were required for 95% maximum per cent egg viability in C. sexmaculata and P. dissecta , respectively (Omkar et al., 2006b)(Figure7).
Table 1: Effect of multiple matings on behavioural patterns and reproductive attributes of C. septempunctata.
Table 1: Effect of multiple matings on behavioural23 patterns and reproductive attributes of C. septempunctata.
Figure 5: Influence of number of matings on the fecundity and hatching of two aphidophagous ladybirds.
Table 1: Effect of multiple matings on behavioural patterns and reproductive attributes of C. septempunctata.
Figure 5: Influence of number of matings on the fecundity and hatching of two aphidophagous ladybirds. Figure 5: Influence of number of matings on the fecundity and hatching of two aphidophagous ladybirds.
Approach Watch Examine
Embrace Mount Copulatory Attempt
Plate 8: Courtship behaviour of ladybirds Plate 8: Courtship behaviour of ladybirds 24
140
120 Male
100 Female
80
60 Longevity (in days) 40
20
0 0 5 10 15 20 25 30 35 40 45
Reproduction (No. of matings)
Figure 6: Influence of multiple matings on the longevity of male and female ofP. dissecta.
Figure 6: Influence of multiple matings on the longevity of male and female of P. dissecta.
Figure 7: Gompertz curves depicting the effect of number of matings on fecundity and fertility of two Figure 7:ladybirds, Gompertz and curves the identification depicting of the optimal effect number of number of matings of matingsrequired foron 95%fecundity and 50% and fertility ofmaximum two ladybirds, theoretical and fecundity the identification and fertility in a of lifetime. optimal number of matings required for 95% and 50% maximum theoretical fecundity and fertility in a lifetime. 25
Plate 9: Range of polymorphism in Menochilus sexmaculatus
Ageing Reproductive output in ladybirds is known to vary with age. Studies have shown that increase in female age leads to increase in fecundity upto a particular age after which it declines (Mishra and Omkar, 2004; Srivastava and Omkar, 2004; Omkar and Singh, 2005; Omkar et al., 2006c. 2010; Mishra and Omkar, 2006). The age at which a ladybird has peak oviposition usually varies with species and can range between 20-40 days of age. Egg viability also responds to change in male age and follows a triangular function as in the case of fecundity. However, fecundity and egg viability normally do not respond to male and female age, respectively. Though both fecundity and egg viability have triangular functions, these are not in synchrony with the female usually peaking before the male. This leads to asynchrony in peaks if adults of the same age are paired together. Thus age differences that facilitate synchronization of these aging trajectories leads to enhanced reproductive output (Omkar and Mishra, 2009). Aging in ladybirds is also known to influence offspring development and survival (Singh and Omkar, 2009).
Mate choice in ladybirds Mate choice is a central process of sexual selection which includes the process of choosing mates as a result of active competition. Traditionally, females have been regarded as the choosier sex while males are the enticers. However, increasing evidences now show that males can be choosy too. Evolution of mate choice was attributed to the parental investment theory by Trivers (1972). The profits of mate choice can be direct or indirect. Direct gains include more fecundity and percent egg viability and indirect benefits include the genetic quality of the offspring (Kokko et al., 2003).
Mate choice is well documented in several Coleopterans such as Tribolium castaneum (Herbst) (Arnaud and Haubruge, 1999; Fedina and Lewis, 2008), bark beetle, Ipspini (Say) (Reid, 1991), oak ambrosia beetle, Platypus quercivorus (Murayama) (Kobayashi, and Ueda, 2002), grain borer, Prostephanus truncates (Horn) (Birkinshaw and Smith, 2001), hide beetle, Dermestesagittari De Geer (McNamara et al., 2004), burying beetle, Nicrophorusquadripunctatus Kraatz (Suzuki, 2009), seed beetle, Callosobruchus chinensis L. (Maklakov and Arnqvist, 2009), dung beetle, Onthophagus sagittarius Fabricius (Watson and Simmons, 2010).
The elytra of ladybird beetles vary in the localization and extent of melanin, a dark pigment (Majerus, 1994). Temporal and geographic variations also contribute for the fluctuations in the frequency of melanic individuals (Timofeeff-Ressovsky, 1940; Creed 1975; Honek, 1975; Bengtson and Hagen, 1977; Majerus and Zakharov, 2000; Honek et al., 2005; Wang et al., 2009; Michie et al., 2010, 2011). The first study of mate choice was conducted on C. septempunctata where melanics were more preferred over the typical (Srivastava and Omkar, 2005b). Polymorphism is one of the striking characteristic of ladybirds (Plate 9). In P. dissecta, females readily mate with typical darker morphs than the pale morphs (Mishra and Omkar, 2014). Study by Dubey et al. (2016) revealed the combined effect of temperature and morphs on the mate choice. At low temperature (15°C) melanics were more preferred as mates while at high temperature no biasness was observed towards any particular morph. Another factors which modulates mate choice are body size (large size mates were preferred), age (middle age males were preferred), food (individuals reared on the abundant diet were more preferred) (Dubey et al., 2016). Extending the studies, mate choice was also studied in respect of several other factors. Ladybirds always prefer to mate with the novel mates (Saxena and Mishra, 2018). Males and females coexisted for several days and then
26
Figure 7: Gompertz curves depicting the effect of number of matings on fecundity and fertility of two ladybirds, and the identification of optimal number of matings required for 95% and 50% maximum theoretical fecundity and fertility in a lifetime.
Plate 9: Range of polymorphism in Menochilus sexmaculatus the choice was given where the unfamiliar males were more preferred. Relatedness is another important factor which influences Platethe mate 9: Range preference of polymorphism in ladybirds (Saxena in Menochilus et al., 2016). sexmacula Ladybirdstus avoid mating with related individuals in order to avoid inbreeding.
Mate selection is not only restricted pre-copulatory but post-copulatory mate selection also exists. The post-copulatory mechanisms of sexual selection are more subtle. Post-copulatory selection links the perceived trait with the gametic quality to ensure apt choice of mate via sperm competition and cryptic female choice. Other prominent post-copulatory mechanisms of sexual selection include differential ejaculate quality and quantity, sperm blockers and killers, testicular hijacking, rejection of mating plugs, etc. In ladybirds also post-copulatory mate selection exists.In one of the experiments, prolonged matings in the ladybirds has been attributed to the mate guarding phenomena. At matings of up to 1 minute deformed eggs were found which dissolve within 10 minutes of laying (Plate 10). Up to 2.00 minutes of mating no eggs are found viable, which indicates that spermatophores have not been transferred as of yet. While fecundity increased in spurts with the increase in mating duration, dramatic increase in fecundity at 30 minutes beyond which there was no increase in fecundity (Figure 8) (Chaudhary et al., 2015b). Another strategy of post-copulatory mate selection is last male sperm precedence. Last male sperm precedence theoretically explains that the last mating male will fertilize more eggs than the previous males. To secure this, males opt for various methods such as behavioural mate guarding in Coenagrion puella (Linnaeus) where males remain in direct contact with females carrying them to ovipositing site (Banks and Thompson, 1985) and non contact guarding in Calopteryx maculatum (Waage). In a double
27
Plate 10: Deformed eggs laid by Menochilus sexmaculatus after the mating of 1 minute.
Plate 10: Deformed eggs laid by Menochilus sexmaculatus after the mating of 1 minute
Plate 10: Deformed eggs laid by Menochilus sexmaculatus after the mating of 1 minute.
Figure 8: Fecudity of Menochilus sexmaculatus at different mating durations mating experiment on M. sexmaculatus it was found that the offspring were majorly sired by the second male irrespectiveFigure 8: of Fecudity the morph of (Chaudhary Menochilus et al., sexmaculatus2016) (Figure at9). different mating durations.
Oviposition Dynamics in Ladybirds
Oviposition is a process in which mature eggs are deposited by the female and it is considered a true measure of reproductive success. It involves the sequence of behavioural and physiological events that commence with the movement of the egg through the oviduct and ends with egg laying on a surface that will help inFigure the sustenance 8: Fecudity of offspring of Menochilus (Klowden, 2009). sexmaculatus Oviposition at behaviour different is also mating helpful durations. in relating behaviour with population and community dynamics (Murdoch et al., 1997; Tammaru and Javeis, 2000; Spencer et al., 2002). Thus, oviposition is a key component in investigation of evolutionary life history traits, population dynamics studies, tri-trophic interactions and biocontrol of insect pests. Successful oviposition requires a sequence of behavioural processes: thechoice of habitat, approaching plant, landing decision on the potential host or not, decision to oviposit or not, number of clutch and size of clutch (Jones, 1991; Bernays and Chapman, 1994; Schoonhoven et al., 1998). In some insects, female protect their eggs and neonates by physical guarding (Trivers, 1972; Mappes and Kaitala, 1994; Tallamy, 1994).
The pattern of egg laying is quite striking in insects, While some lay their eggssingly, others prefer to oviposit in batches. In insects, it has been observed that within a clutch some eggs hatch, while others
28
Figure 9: Offspring phenotype of Menochilus sexmaculatus during double mating experiment do not hatch. Some of these eggs are consumed by the ladybird larvae as meal. In coccinellids, broadly twoFigure kinds 9: of Offspring eggs have phenotype been seen ofi.e Menochilus. viable and inviablesexmaculatus eggs. Harmoniaduring double axyridis mating (Pallas) and other experiment. ladybirds produce three types of eggs: apparently infertile eggs, viable eggs and inviable‘ eggs where a larva develops but does not emerge from the egg capsule (Perry and Roitberg, 2006). In M. sexmaculatus, experimentFigure was 9: Offspring conducted phenotype where Petri of dish Menochilus was partitioned sexmaculatus in two chambers during anddouble the aphidsmating were kept in lowerexperiment. chamber (Plate 10). It has been found that the oviposition was modulated by perceiving the prey quantity (Singh et al., 2016a).
Plate 11: Schematic presentation of the experimental arena. Ladybird was kept on the upper chamber which was created by separating the Petri dishes with muslin cloth. Plate 11: Schematic presentation of the experimental arena. Ladybird was kept on the upper chamber which was created by separating the Petri dishes with muslin cloth. Plate 11: Schematic presentation of the experimental arena. Ladybird was kept on the upper chamber which was created by separating the Petri dishes with muslin cloth. 29 Conclusions The predaceous ladybirds have a promising future in the biocontrol of insect pests of agricultural importance. While majority of ladybirds are generalists, some are specialists. There are lots of arguments on prey specialization in predaceous ladybirds. However, the resource partitioning and consistent exposure of a single prey type might have evolved the prey specialization in predaceous ladybirds. Moreover, the prey specialization in ladybirds is generally considered as a function of body size, prey size and prey density.
Mating and reproductive studies in ladybirds have provided knowledge on the optimal conditions pertaining to number and quality of mates to produce better progeny both in terms of quantity and quality. Similarly, the information pertaining to age, aging trajectories and age differences between mates not only increases the level of knowledge on ladybird physiology in general but may also help in mass multiplication of ladybirds by allowing mating with optimal age individuals. The intraguild interactions amongst the ladybird species distress the coexistence of ladybirds and displace many indigenous ladybird fauna, the biological invasions of certain dominant species could result in complete disappearance of numerous native ladybird species. Amongst the abiotic factors, temperature has a major impact on the ladybird’s life attributes. Moreover, the optimization of abiotic conditions, like temperature and light, is a prerequisite for the augmentative rearing of ladybirds. There is an utmost need to understand the role of ladybirds in pest management through comprehensive ecological and ethological studies supported with laboratory experimentation, and glasshouse and field studies. The use of biocontrol agents in the suppression of pest populations minimizes the use of pesticides in agriculture. Biocontrol is an ecofriendly technique which is cost effective in the long run and self-perpetuating, and would lead to sustainable agriculture.
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37 Simmons, L.W. (2005). The evolution of polyandry: sperm competition, sperm selection, and offspring viability. Annual Reviews on Ecology and Evolutionary System, 36: 125–146. Singh, K. and Omkar (2009) Effect of parental ageing on offspring developmental and survival attributes in an aphidophagous ladybird, Cheilomenes sexmaculata. Journal of Applied Entomology, 133 500-504. Singh, N, Mishra G and Omkar (2014). Does temperature modify slow and fast development in two aphidophagous ladybirds? Journal of Thermal Biology, 39, 24-31. Singh, N, Mishra G and Omkar (2016a) Slow and fast development in two aphidophagous ladybirds on scarce and abundant prey supply. Bulletin of Entomological Research, 106, 347-358. Singh, N., Mishra, G. and Omkar (2016b).Effect of photoperiod on slow and fast developing individuals in aphidophagous ladybirds, Menochilus sexmaculatus and Propylea dissecta (Coleoptera: Coccinellidae). Insect Science, 23, 117-133. Sloggett, J.J., Obrycki, J.J. and Haynes, K.F. (2009). Identification and quantification of predation: novel use of gas chromatography- mass spectrometric analysis of prey alkaloid markers. Functional Ecology, 23, 416-426. Smith, H.S. (1919). On some phases of insect control by the biological method.Journal of Economic Entomology, 12, 288-292. Southwood, T.R,E. (1966).Ecological Methods, with Particular Reference to the Study of Insect Populations. Methuen, London. P. 391. Spencer, M., Blaustein, L. and Cohen, J.E. (2002). Oviposition habitat selection by mosquitoes (Culisetalongiareolata) and consequences for population size. Ecology, 83(3), 669-679. Srivastava S. and Omkar (2003). Influence of temperature on certain biological attributes of a ladybeetle, Coccinella septempunctata Linnaeus. Entomologia Sinica, 10(3), 185-193. Srivastava, S and Omkar (2004). Age Specific Mating and Reproductive Senescence in Seven Spotted Ladybird, Coccinella septempunctata. Journal of Applied Entomology, Blackwell Wiley Interscience Germany, 128(6), 452-458. Srivastava, S. and Omkar (2005a). Short and long term benefits of promiscuity in a seven-spotted ladybird, Coccinella septempunctata. International Journal of Tropical Insect Science, 25(3), 176-181. Srivastava, S. and Omkar (2005b). Mate choice and reproductive success of two morphs of the seven spotted ladybird, Coccinella septempunctata (Coleoptera: Coccinellidae). European Journal of Entomology, 102, 189-194. Stuart, M.K., Greenstone, M.H. (1990). Beyond ELISA: a rapid, sensitive, specific immunodot assay for identification of predator stomach contents. Annals of the Entomological Society of America, 83, 1101-1107. Suzuki, S. (2009). Mate choice and copulation frequency in the burying beetle Nicrophorus quadripunctatus(Coleoptera: Silphidae): Effect of male body size and presence of a rival. Psyche- Journal of Entomology, doi:10.1155/2009/904604
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39 Organizing Committee
PATRON Dr. A.K. Singh APEX LIFE SCIENCES DDG ( Extension ) ICAR & Director ICAR-IARI
CONVENER Dr. Rakesh Sharma Head, Division of Entomology
MEMBERS Dr. H.R. Sardana Director , NCIPM
Dr Surinder Kumar Principal Scientist, NCIPM
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ELEVENTH DR. S. PRADHAN MEMORIAL LECTURE September 9, 2019
Predaceous Ladybirds: A Reproductive Perspective
By
Prof. Omkar Professor Omkar received his graduation, post graduation and Doctorate degrees from DDU, Gorakhpur University. He continued his Post doctoral research at his alma mater. After a brief stint Ladybird Research Laboratory, Department of Zoology, at Central Tasar Research and Training Institute , Central Silk Board at Ranchi he began his teaching University of Lucknow, Lucknow-226 007 (India) career at the Department of Zoology, University of Lucknow from December 1986 and took over the charge of Head, Department of Zoology, University of Lucknow (2016-2019). He heads the only Ladybird Research laboratory in India. His research on biology, ecology and behaviour of ladybird beetles have resulted in landmark contributions on ladybird predation, life history traits, reproductive attributes and optimization of rearing factors with over 36 pioneering studies spanning, multiple matings, promiscuity, optimal number of matings, costs and benefits of reproduction, ageing, developmental rate polymorphism, sexual selection and mate familiarity all of which have helped in standardizing mass multiplication of ladybirds. He was awarded a grant under Centre of Excellence in Biocontrol of Insects Pests, Govt. of Uttar Pradesh, and is part of IUCN in the Ladybird Species Specialist Commission besides running various research projects funded by state and central funding agencies. He has visited School of Biological Sciences, University of East Anglia, Norwich, UK (2006) and Czech Academy of Sciences, Ceske Budejovice, Czech Republic (2016) under Royal Society and Czech Academy of Sciences-INSA Bilateral Exchange of Scientists programme, respectively. He has supervised 23 Ph. D. students and has 270 articles in reputed international and national journals to his credit. He has authored 11 books, three of which have been published by Academic Press, Elsevier Inc. USA and Springer Nature, Singapore. Prof. Omkar has contributed significally as Member, Chairman of many academic and administrative bodies at University of Lucknow and other universities; Coordinator, UGC-SAP (DRS-II), DST (FIST), and DST-PURSE, University of Lucknow and UGC nominated member of advisory committee for UGC-SAP (DRS–II) at Panjabi University, Patiala; Member of Selection Boards of multiple universities and state commissions as subject expert and examiner of more than 40 universities in 14 states of our country; Chief Editor of Journal of Applied Bioscience, Associate Editor of International Journal of Tropical Insect Science (Springer Nature), and referee of several international and national journals and funding agencies. DIVISION OF ENTOMOLOGY Recipient of several awards among which Saraswati Samman (2017), Govt. of UP, Prof. TN Ananthakrishnan Foundation Award, Chennai, (2012) are noteworthy. Also an elected Fellow of ICAR-INDIAN AGRICULTURAL RESEARCH INSTITUTE National Academy of Sciences, India, (FNASc), Zoological Society of India, The Entomological Society of India, etc. and President, International Society of Applied Biology, Lucknow, NEW DELHI & ENTOMOLOGICAL SOCIETY OF INDIA