Ibottor of $I)Tlasiopi)P in '^^'^>- ZOOLOGY

CHROMOECOLOGY AND MORPHOMETRICAL STUDIES ON SWARMING ACRIDOIDS

'^ VI y/^ ABSTRACT l/X;^^,,^ '^^-^ THESIS ^/Vv}

SUBMITTED FOR THE AWARD OF THE DEGREE OF IBottor of $I)tlasiopI)p IN '^^'^>- ZOOLOGY

Cy^' BY MD. EHTASHAMUL HASSAN

DEPARTMENT OF ZOOLOGY FACULTY OF LIFE SCIENCES ALIGARH MUSLIM UNIVERSITY ALIGARH (INDIA) 2006 -, *7a

Locusts and grasshoppers have long been regarded as one of the most important groups of agricultural pests belonging to the order Orthoptera. The Orthoptera which includes the dreadfully destructive locusts is on^ of the largest orders oi insects having over 17,250 species known to science globally with more than 900 species have been recorded from India. They are almost all terrestrial and, though usually capable of jumping actively, relatively few strong fliers are known, all belonging to the family acrididae. Locusts and grasshoppers are among the oldest enemies of mankind. Entomologists have made exhaustive studies on the biology, morphology, ecology, control and behaviour of these insects. They have long been regarded as one of the most important groups of agricultural pests cause extensive damage to different crops. Both nymphs as well as adults stages are responsible for the damaging the cultivated crops.

Adults of Phlaeoba infumata are medium sized, with the head elongated, face oblique and frontal ridge raised between median ocellus. The ground colour is brown to carbon black. They are distributed in India (Bihar, Orissa and Uttar Pradesh), East Nepal, Myanmar, Thailand, Malaysia and China. Both adult and their hoppers are a serious pest of economically important grasses and a number of agricultural crops like wheat, sugarcane, millet, maize, and rice, which make them as polyphagous in nature. Apart from these it was found to be a serious pest of bamboo in China

%Ui^^id and recently assumed to be a pest of a number of medicinal plants. They are found in long grasses. Adults and nymphs have tendency to aggregate during the hottest part of the day in bushes and crop fields.

The adults of Oedaleus abruptus are medium sized, with distinct 'X' marking on pronotum, inhibit short and dry grasses, and it was found to be a periodically major pest of millet with 3-4 overlapped generations a year. Life cycle is completed from 44-82 days depending upon the experimental conditions provided, cause damage to a number of cultivated crops in North India and also reported from different geographical areas throughout the globe specially from African countries, therefore has polyphagus status. Khan and Aziz (1973) observed Oedaleus abruptus as an important polyphagous pest with two generations a year, and being found throughout the year except during very hot and cold periods.

Both these species are found in a mixed population with each other at all stages. The life stages in laboratory conditions have shown that under constant conditions there can be 3-4 generations in a year. However, the number of generations may be reduced to 2-3 in natural populations. Adults are found reproductively active in rainy season and in the months of September, October, March and April, but do remain active in winter season also but without reproductive activities. The tendency of swarming is more prominent in P. infumata as compared to O. abruptus.

The present work is a preliminary attempt to evaluate their biology, ecology, and population studies with especial reference to

XULJ^AS swarming behaviour in North India. Chromoecoloygy and morphometrical parametres were chosen as tools to evaluate the magnitude of swarming behviour and extent of phasepolymorphic behaviour by these two grasshoppers by providing them ecological conditions gradient. Two different temperatures {27±1°C and 37±1°C) and number of hoppers per jar (isolated and crowded conditions), combined to each other, thus four different conditions were obtained. Newly emerged hoppers of both grasshoppers reared at two different temperatures under isolated and crowded conditions. During each instar morphometrical measurement, colour changes and their bionomics were recorded and analysed using suitable treatments.

The population of hoppers and adults were dense where grasses like Cyperous rotundus, Seteria glauca, Paspalum distichum, Andropogon adoratus, Cynodon dactylon were dominating the vegetation. The fluctuation in numbers of grasshopper is attributed to the other ability of these grasses. The early nymphal instars have preferred Cynodon dactylon while the advance stages enter in the crop fields. The food preference in nymphs and adults and successful completion of cycle has been recorded typically in these grasshoppers. It was found that the distribution of these species remains directly proportional to the preferential value of the food plants available.

The population size of these grasshoppers with reference to seasonal variations on monthly basis of all stages through random sampling has been estimated. Both these pests form a common population in North Indian grassland and cultivated crops. Their life-cycles are of similar nature except flight potential and vigorous

ftiti:>i^ activities of hoppers in P. infumata is the unique features to that of O. abruptus which was found mostly active when remain crowded otherwise show least activities. It was found that the populations were high twice a year firstly during March - May and secondly during August-October with reproductive activities. Khan and Aziz (1973b) made similar observations on seasonal variation in population of hoppers and adults of Oedaleus abruptus. Seasonal variations govern the population structure in both the species.

Different processes of life cycle were observed under different ecological conditions and it was found that the least period was taken at high temperature under crowded condition while at low temperature prolonged biological processes were observed which suggest preference of higher temperature and higher density for faster rate of development.

Copulation in these two species is preceded by e.laborate courtship with special reference to long copulating time and the mechanism involved in oviposition is of acridian pattern. The egg laying period was shorter at high temperature under crowded condition. Similarly the size of egg-pods, length and breadth of egg and average number of eggs per egg-pod per female was higher at high temperature under crowded condition. The viability of egg- pods and eggs was found to be affected by high temperature but influence of crowded condition was not observed. Moist soil was found to be most preferred for egg laying in these two grasshoppers. Diapause was not observed at any stage of life cycle under any experimental condition, but prolonged developmental period was observed at each stage of life cycle, depending upon experimental conditions.

'A^i* The development of hoppers in these two species is of great interest with reference to number of hopper instars and it was found that under all ecological conditions only five hopper instars were observed in both sexes. The process of alar rudiment reversal was found similar in both grasshoppers, which was observed after third instars in both sexes. Similarly, development of external genitalia, development of posterior margin of pronotum and formation of eye stripes were studied in detail.

The range of colour in acridoids as a whole is in shades of brown (from pale-straw to black), gray and green, while bright red, blue or yellow appear normally on the hind wings and often on the hind legs, but seldom on other parts. Chromoecology of these grasshoppers was studied under different ecological conditions to evaluate magnitude of swarming behavior. To study the colour patterns, only head and pronotum were taken into account as they exhibit prominent colour components. Colour changes were observed visually, using hand lens and under light microscope which not only magnified the object (hopper's colour components) but also magnified the concept of colour changes during postembryonic development. Colour variations were recorded in the form of colour plates under all ecological conditions and deciphered using colour dictionary by Maerz and Paul (1950). Both grasshoppers exhibited wide range of colour patters under stress of different experimental conditions. Bold patterns mainly dark brown and black colours were observed at high temperatures under crowded condition in case of Phlaeoba infumata, while in case of Oedaleus abruptus ground colour varied from green to brown. Other components like dark spots and srteaks also exhibited considerable variation under stress of different experimental conditions. Biologically, the most important difference between the crowded and isolated population was higher activities and tendency of aggregation characterised by higher feeding potential and band formation at high temperature under crowded condition.

For morphometrical studies sixteen body parts were measured using vernier dial caliper. Ratios and their differences of body parts were calculated, and a number of parameters were found to be significant, especially length of hind femur, length and height of pronotum, length of hind tibia and length of elytron at high temperature under crowded conditions, Chandra (1983) observed a small concentration of Oedaleus senegalensis in Rajasthan desert, which is of great importance in desert locust populations in relation to habitats. Rafeeq 86 Rizvi (1989) and Razak 85 Rizvi (1989) have made some observations on the suspected gregariousness in Oedaleus senegalensis and Acrida exaltata, respectively. Zolotarevsky (1930), in his work on Locusta migretoria capita (Sauss.), used the same measurement, but reverse the ratios, and introduced two new measurements; maximum widths of head and pronotum height. Subsequently, Maxwell- Darling (1934), in his work on Schistocerca gregaria, used four measurements: elytron length (E), hind fumer length (F), ponotum length (L) and the minimum pronotum width, also different ratios were taken to ensure phase polymorphism in locust. Among these ratios the elytro-femoral ratio (E/F) has been most commonly used in studies on the morphological phase variation in different locust species and in recording the morphometries characters of populations. Ten differences between means of ratios under isolated and crowded population of different body parts were calculated, and a number of differences between means of ratios were found to be significant but only at high temperature in female population. At low temperature only one ratio was found to be significant, which is an indicative of hidden swarming tendency of these two grasshoppers. Means, standard errors, standard deviations and mean differences of body parts of adults of both P. infumata and O. abruptus under isolated and crowded conditions. Similar observation was made by Rizvi and Basit (1990) for North-Indian grasshoppers and found occasionally swarming tendericy.

The observation certainly leading to accept the fact that both the species are having distinct behaviour of gregarization, swarm- forming and having the ability of mass active behaviour with local migratory instincts. These observations are a substantial addition to the knowledge of polymorphism in Acridoids.

On the basis of the present studies it can convincingly be concluded that the species under study are very aggressive in behaviour, can assume new dimensions of behaviour, may become gregarious and migratory under suitable ecological conditions. The life-cycles of these species, hitherto unknown, have been made known. A complete eco-ethological and chromoecological profile, under all considerations, has been completed and the entire research work will be an addition to Entomology in general and to acridologists in particular. # l^

CHROMOECOLOGY AND MORPHOMETRICAL STUDIES ON SWARMING ACRIDOIDS

V -.^,--- •. f //- -//':• I / ^"^ r' -. "i >^. N THESIS

SUBMITTED FOR THE AWARD OF THE DEGREE OF Bottor of $^tlo)BiopIiP IC IN ^ _ N ZOOLOGY ^ /

' JV BY MD. EHTASHAMUL HASSAN

DEPARTMENT OF ZOOLOGY FACULTY OF LIFE SCIENCES AUGARH MUSLIM UNIVERSITY ALIGARH (INDIA) 2006 f684y I External: 2700920/21-3430 ''^°""M Internal: 3430 DEPARTMENT OF ZOOLOGY ALIGARH MUSLIM UNIVERSITY ALIGARH—202002 INDIA

CHAIRMAN D.No /ZD Dated..h:..?,.:,:^.±

OE^ricAi^

This is to certify that the work embodied in this thesis entitled "Choromoecology and morphometrical studies on swarming Acridiods" is the original research work carried out by Mr. Md. Ehtashamul Hassan initially under the supervision of Mr. S. K. A. Rizvi, and completed under my supervision. The present work is a distinct addition to the existing knowledge on the morphometries and chromoecology of acridoids.

I allow Mr• Md Ehtashamul Hassan to submit this thesis to the AUgarh Muslim University for the award of the Ph.D. Degree in Zoology of the Aligarh Muslim University.

Dr. Mohammad Hayat Dedicated to My Grand Parents And Parents CONTENTS ^^^^^ Page No.

ACKNOWLEDGEMENTS

1. INTRODUCTION 1

2. REVIEW OF LITERATURE 8

3. MATERIALS AND METHODS 23

4. OBSERVATIONS

(A) Population Studies 27

(B) Biology 30

I. Adult 30

(a) Prc-Copulation Period 31

(b)C o p u1 a t i o n 31

(d)Posl-Oviposition 33

II. Egg-pod and Egg 33

III. Incubation 33

IV. Hatching and Moulting 34

V. Development of Hoppers 35

I. Description of colour patterns in

Phlaeoba infumata 44

II. Description of colour patterns in

Oedaleus ahriiptus 54 (D) Morphometries 63

5. DISCUSSION 68

6. SUMMARY 78

7. REFERENCES 84 LIST OF TABLES

1. Effect of two temperatures on Copulation period, Egg-laying period and moulting period of Phlaeoha infumata and Oedaleus ahnipliis fed on Zea mays. 2. Summary of life history of hoppers and adults at 37±rC and 70±5% R.H. fed on Zea mays. 3. Summary of life history of hoppers and adults at 27±1°C and 70±5% R.H. fed on Zea mays. 4. Number of egg-pods laid by Phlaeoba infumata and Oedaleus ahruptus in different types of soils. 5. Measurements of Eggs, Egg-pods and Hatchlings of Phlaeoha mfumata and Oedaleus abruptus. 6. Effect of two different temperatures on the incubation period and hatching of eggs of Phlaeoba infumata and Oedaleus abruptus at 70±5% R.H. 7. Effect of isolated and crowded conditions on the fecundity of females of Phlaeoba infumata and Oedaleus ahruptus. 8. Effect of isolated and crowded conditions on the survival percentage, daily rate of development and growth index of Phlaeoha infumata and Oedaleus ahruptus at 27±1°C fed on Zea mays. 9. Effect of isolated and crowded conditions on the survival percentage, daily rate of development and growth index of Phlaeoba infumata and Oedaleus abruptus at 37±I°C fed on Zea mays. 10. Reversal of alar rudiments in hopper instars of Phlaeoha infumata and Oedaleus ahruptus (L, rudiments lateral; R, rudiments reversed; Ad, Adult). 11. Application of Dyar's law on tire hoppers of Phlaeoba infumata at 27±l"C fed on Zea mays. 12. Application of Dyar's law on the hoppers of Phlaeoba infumata at 37±1°C fed on Zea mays. 13. Application of Dyar's law on the hoppers of Oedaleus abruptus at 27±1°C fed on Zea mays. 14. Application of Dyar's law on the hoppers of Oedaleus abruptus at 37±1°C fed on Zea mays. \5. Phlaeoba infumata: 1st Instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. \6. Phlaeoba infumata: Ilnd Instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. \1. Phlaeoba infumata: lllrd Instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. \S. Phlaeoba infumata: IVth instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. \9. Phlaeoba infumata: Vth Instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. 20. Phlaeoba infumata: Adult Male: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. 21. Phlaeoba infumala: Adult Female: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. 22. Oedaleiis abniptiis: 1st Instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. 23. Oedaleiis abniptus: llnd Instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul. London. 24. Occlak'iis nbniplus: illrd instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. 25. Oedaleiis abniplns: iVth Instar: ('olour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. 26. Oedaleiis abniptus: Vtli Instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. 27. Oedaleiis abniptus: Adult Male: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. 28. Oedaleus abniptus: Adult Female: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. 29. Measurements (cm) of body parts and rate of increase in various inslars of Phlaeoba infumata under isolated conditions fed on Zea mays at 27±1 °C (Male) 30. Measurements (cm) of body parts and rate of increase in various instars of Phleaoha infumata under isolated conditions fed on Zea mays at 27±l C (Female) 31. Measurements (cm) of body parts and increase in various instars of Phleaoba infumata under crowded conditions fed on Zea mays at 27±l "C (Male) 32. Measurements (cm) of body parts and increase in various instars of Phleaoba infumata under crowded conditions fed on Zea mays at 27±l C (Female) 33. Measurements (cm) of body parts and rate of increase in various instars of Phlaeoba infumata under isolated conditions fed on Zea mays at 37±l C (Male) 34. Measurements (cm) of body parts and rate of increase in various instars of Phleaoba infumata under isolated conditions fed on Zea mays at 37±l "C (Female) 35. Measurements (cm) of body parts and increase in various instars of Phleaoba infumata under crowded conditions fed on Zea mays at 37±1 °C (Male) 36. Measurements (cm) of body parts and increase in various instars of Phleaoba infumata under crowded conditions fed on Zea mays at 37±1 °C (Female) 37. Measurements (cm) of body parts and rate of increase in various instars of Oedaleus abruptus under isolated conditions fed on Zea mays at 27±1 °C (Male) 38. Measurements (cm) of body parts and rate of increase in various instars of Oedaleus abruptus under isolated conditions fed on Zea mays at 27±1 °C (Female) 39. Measurements (cm) of body parts and increase in various instars of Oedaleus abruptus under crowded conditions fed on Zea mays at 27±l °C (Male) 40. Measurements (cm) of body parts and increase in various instars of Oedaleus abruptus under crowded conditions fed on Zea mays at 27±1 ''C (Female) 41. Measurements (cm) of body parts and rate of increase in various instars of Phlaeoba infumata under isolated conditions fed on Zea mays at 37±1 "C (Male) 42. Measurements (cm) of body parts and rate of increase in various instars of Oedaleus abruptus under isolated conditions fed on Zea mays at 37±I "C (Female) 43. McasLircmcnls (cm) of body parts and increase in various inslars of Ocdaleiis uhniplus under crowded conditions fed on Zea mays at 37±1 °C (Male) 44. Measurements (cm) of body parts and increase in various instars of Oedaleiis abnipliis under crowded conditions fed on Zea mays at 37±1 "^C (Female) 45. Means, standard errors and standard deviations of crowded and isolated adults of Phlaeoba infumata Brunner fed on Zea mays at 27±1 °C 46. Means, standard errors and standard deviations of crowded and isolated adults of Phlaeoba infumata Brunner fed on Zea mays at 37±l °C 47. Difference between means of body part measurements of crowded and isolated adults of Phlaeoba infumata at 27±l"C fed on Zea mays (Male). 48. Difference between means of body part measurements of crowded and i.solatcd adults of Phlaeoba infumata at 27±l"C fed on Zea mays (Female). 49. Difference between means of body part measurements of crowded and isolated adults of Phlaeoba infumata at 37±1°C fed on Zea mays (Male). 50. Difference between means of body part measurements of crowded and isolated adults of Phlaeoba infumata at 37±1°C fed on Zea mays (Female). 51. Means, standard errors and standard deviations of crowded and isolated adults of Oedaleus ahruptus Thunberg fed on Zea mays at 27±1 °C 52. Means, standard errors and standard deviations of crowded and isolated adults of Oedaleus abruptus Thunberg fed on Zea mays at 37±l °C 53. Difference between means of body part measurements of crowded and isolated adults of Oedaleus abruptus at 27±l C fed on Zea mays (Male). 54. Difference between means of body part measurements of crowded and isolated adults of Oedaleus abruptus at 27±l''C fed on Zea mays (Female). 55. Difference between means of body part measurements of crowded and isolated adults of Oedaleus ahruptus at 37±l"C fed on Zea mays (Male). 56. Difference between means of body part measurements of crowded and isolated adults of Oedaleus abruptus at 37±l"C fed on Zea mays (Female). 57. Mean ratios in crowded and isolated conditions for adults of Phlaeoba infumata at 27± I C fed on Zea mays. 58. Mean ratios in crowded and isolated conditions for adults of Phlaeoba infumata at 37±l''C fed on Zea mays. 59. Differences between means of ratios in crowded and isolated adults of Phlaeoba infumata at 27± I C fed on Zea mays. 60. Differences between means of ratios in crowded and isolated adults of Phlaeoba infumata at 37±1°C fed on Zea mays. 61. Mean ratios in crowded and isolated conditions for adults of Oedaleus abruptus at 27±l C fed on Zea mays. 61. Mean ratios in crowded and isolated conditions for adults of Oedaleus abruptus at 37±l C fed on Zea mays. 63. Differences between means of ratios in crowded and isolated adults of Oedaleus abruptus at 27± I C fed on Zea mays. 64. Differences between means of ratios in crowded and isolated adults of Oedaleus abruptus at 37±i ^C fed on Zea mays. LIST OF FIGURES

1. Distribution of Phlaeoha infiimata Brunner & Oeclaleiis ahriiplus Thunberg 2. Taiisil Koil Aligarii 3. Meteorological data ofAligarh during 2002 4. Meteorological data ofAligarh during 2003 5. Meteorological data ofAligarh during 2004 6. Natural population fluctuation of Phlaeoba infumata Brunner 7. "Natural population fluctuation of Oedaleus abruptits Thunberg 8. Nymphal duration and range variables of Phlaeoba infumata under isolated and crowded conditions at 27±l °C fed on Zea mays. 9. Nymphal duration and range variables of Phlaeoba infiimala under isolated and crowded conditions at 37±l °C fed on Zea mays. 10. Nymphal duration and range variables of Oedaleus ahruptus under isolated and crowded conditions at 27±l °C fed on Zea mays. 11. Nymphal duration and range variables of Oedaleus abruptus under isolated and crowded conditions at 37±1 °C fed on Zea mays. 12. Pre-oviposition, Oviposition and Post-oviposition periods and longevity of adults of Phlaeoha infumata under isolated condition at 27±1°C fed on Zea mays. 13. Pre-oviposition, Oviposition and Post-oviposition periods and longevity of adults of Phlaeoba infumata under crowded condition at 27±1°C fed on Zea mays. 14. Pre-oviposition, Oviposition and Post-oviposition periods and longevity of adults of Phlaeoha infumata under isolated condition at 37±1°C fed on Zea mays. 15. Pre-oviposition, Oviposition and Post-oviposition periods and longevity of adults of Phlaeoba infumata under crowded condition at 37±1°C fed on Zea mays. 16. Pre-oviposition, Oviposition and Post-oviposition periods and longevity of adults of Oedaleus abruptus under isolated condition at 27±I°C fed on Zea mays. 17. Pre-oviposition, Oviposition and Post-oviposition periods and longevity of adults of Oedaleus abruptus under crowded condition at 27±1°C fed on Zea mays. 18. Pre-oviposition, Oviposition and Post-oviposition periods and longevity of adults of Oedaleus abruptus under isolated condition at 37±1°C fed on Zea mays. 19. Pre-oviposition, Oviposition and Post-oviposition periods and longevity of adults of Oedaleus abruptus under crowded condition at 37±1°C fed on Zea mays. 20. Application of Dyar's law to Phlaeoba infumata Brunner under isolated and crowded conditions at 27 ± I "C & 37 ± 1 °C fed on Zea mays. 21. Application of Dyar's law to Oedaleus abriiptus Tiiunberg under isolated and crowded conditions at 27 ± 1 °C & 37 ± 1 °C fed on Zea mays. 22. A diagrammatic representation showing formation of eye stripes during hoppers development in Oedaleus ahruptus. 23. A diagrammatic representation showing development of posterior edge of pronotum during the successive hopper instar in Phlaeoha iiifiinuita and Oedaleus abruptus. lA. Phlaeoha infumata - Showing development of alar rudiments and reversal of wings during hopper development. a.Oedaleus abruptus - Showing development of alar rudiments and reversal of wings during hopper development. 26. Phlaeoha infumata Brunner. growth of external genitalia of male. 27. Phlaeoha infumata Brunner, growth of external genitalia of female. 2S. Oedaleus abruptus Thunberg, growth of external genitalia of male. 29. Oedaleus ahruptus Thunberg, growth of external genitalia of female. 30. Scheme of Morphometries 3 1. Instarwise increase in the Total Body Length in Phlaeoha infumata and Oedaleus ahruptus under isolated and crowded conditions at 37 ± 1 "C & 27 ± 1 °C fed on Zea mays 32. Instarwise increase in the Width of Head in Phlaeoha infumata and Oedaleus abruptus under isolated and crowded conditions at 37 ± 1 °C & 27 i I "C fed on Zea mays 33. Instarwise increase in the Pronotum Length in Phlaeoha infumata and Oedaleus ahruptus under isolated and crowded conditions at 37 ± 1 °C & 27 ± 1 °C fed on Zea mays 34. Instarwise increase in the Pronotum Height in Phlaeoha infumata and Oedaleus abruptus under isolated and crowded conditions at 37 ± I °C & 27 ± I °C fed on Zea mays 35. Instarwise increase in the Length of Hind Femur in Phlaeoha infumata and Oedaleus abruptus under isolated and crowded conditions at 37 ± 1 °C & 27 ± 1 "C fed on Zea mays 36. Instarwise increase in the Length of Hind Tibia in Phlaeoha infumata and Oedaleus abruptus under isolated and crowded conditions at 37 ± 1 °C & 27 ± 1 °C fed on Zea mays 37. Instarwise increase in the Length of Elytron in Phlaeoha infumata and Oedaleus abruptus under isolated and crowded conditions at 37 ± I °C & 27 i 1 °C fed on Zea mays 38. Instarwise increase in the Length of Antennae in Phlaeoha infumata and Oedaleus ahruptus under isolated and crowded conditions at 37 ± 1 °C & 27 ± 1 °C fed on Zea mays 39. Showing breeding areas of: A. Oedaleus abruptus B. Phlaeoha i nfu m a t a. 40. Rearing cages in the laboratory. 41. Phlaeoha infumata Brunner: A. Female B. Male 42. Oedaleus abruptus Thunberg A. Showing different types of colours under natural condition. B. Showing dark patters under stress of high temperature and crowded condition. 43.Phlaeoba infumata Brunner sliowing: A. Adult female showing dark spots and lateral streaks. B. Ilnd instar hopper showing dorsal streaks on head and pronotum. 44. Phlaeoba infumata - 1st Instar (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimen1;al conditions at high temperature and low temperatures. 45. Phlaeoba infumata - Ilnd Instar (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimental conditions at high temperature and low temperatures. 46. Phlaeoba infumata - Ilird Instar (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimental conditions at high temperature and low temperatures. 47. Phlaeoba infumata - IVth Instar (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimental conditions at high temperature and low temperatures. 48. Phlaeoba infumata - Vth Instar (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimental conditions at high temperature and low temperatures. 49. Phlaeoba infumata - Male (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimental conditions at high temperature and low temperatures. 50. Phlaeoba infumata - female (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under dilTerent experimental conditions at high temperature and low temperatures. 51. Oedaleus abruptus - 1st Instar (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimental conditions at high temperature and low temperatures. 52. Oedaleus abruptus - Ilnd Instar (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimental conditions at high temperature and low temperatures. 53. Oedaleus abruptus - Ulrd Instar (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimental conditions at high temperature and low temperatures. 54. Oedaleus abruptus - IVth Instar (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimental conditions at high temperature and low temperatures. 55. Oedaleus abruptus - Vth Instar (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimental conditions at high temperature and low temperatures. 56. Oedaleus abruptus - Male (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimental conditions at high temperature and low temperatures. 57. Oedaleus abruptus - Female (Dorsal and Lateral views of Head and Prothorax) showning colour spectra under different experimental conditions at high temperature and low temperatures. 58. Vernier dial caliper JJB

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Aoddiifie daM?rid aarim ine coarde of^a^or4 ao/ie.

fi^^. ^4iad4amac .^ddanj chapteR'i INTRODUCTION

For thousand of years, grasshoppers and locusts have been among the most destructive pests of agricultural crops and there is no stretch of land free from them. Grasshopper species compete with the human for plant resources all over the world (Dempster, 1963) and hence threatens human prosperity and survival. An insect species may not be harmful at low population densities but when the number increases under certain conditions, they can cause great harm and become pests (Youdeowei 86 Service, 1986). The degree of harm caused by insect pests varies considerably with the particular species of insect concerned by frequency of occurrence and density of pest population, the prevailing circumstances and the nature of the host.

Moreover, our country is based on agriculture, and about 57% of its population is engaged in agriculture one way or other. The problem of grasshoppers and locusts seems meager nowadays, may become detrimental in neat future, if unchanged. The devastation caused by the grasshoppers is not vigorous like locusts but its persistency, omnipresence and hidden tendency to become a locust, not only attracted acridologists but compel them to take necessary steps for their control and containment.

The Orthoptera which includes the dreadfully destructive locusts is one of the largest orders of insects having over 17,250 species known to science globally with more than 900 species have been recorded from India. The term 'locust' is applied to those grasshoppers that exhibit physiological, morphological and behavioural changes from solitarious phenotype to a gregarious phenotype (Dempster, 1963; Uvarov, 1966; Farrow, 1990). In addition to this, some of the workers also defined the term 'locust' as species of acridids which are capable under certain conditions of forming large swarms which move over wide areas causing great devastation of natural and cultivated vegetation where they feed. Uvarov (1921; 1928) has proposed a theory that each species of locust can exist in two main forms ('Phases'), which differ structurally and biologically. These are the gregarious phase (Phasis gregaria] and the Solitary phase (Phasis Solitaria) and the two are often so distinct as to have been regarded by earlier taxonomists as separate species. Intermediates (Phasis transiens) also occur during the transition of population from one extreme to another.

The Orthopterans are distributed throughout the physiographic zones of world but their distribution largely depends upon the vegetations prevailing in grass fields, forests and agricultural lands. Temperatures, seasonal precipitations of rain fall and soil conditions are some important factors, which also determine the distribution of these grasshoppers. India provides a unique habitat and most suitable climatic conditions as humid grasslands in north India.

Phlaeoba infumata commonly called as nursery locust is moderate in size, with a brown to black ground colour. They have been reported from India (West Bengal, Arunachal Pradesh, Bihar, Delhi, Himachal Pradesh, Orisa, Rajasthan and Tamilnadu), East Nepal and Myanmar. Both adult and hopper stages have been reported as a pest of Bamboo leaves and also found damaging to rice (Oryza sativa), maize [Zea mays), and sugarcane [Saccharum officinarum) (Roffey, 1979). During surveys in and around Aligarh, it was found to be attacking a number of cultivated crops like guava {Psidium guajava), jowar {Sorghum vulgare), bajra {Pennisetum typhoideum), millet (Pennisetum typhoides) and groundnut (Arachis hypogaea) thereby causing severe damage to these economically important crops. Besides, medicinal plants under cultivation in the area were also damaged severely at seedling stage inflicting considerable loss to the pharmaceuticals. In the last few years it has become a major pest of these plantations in and around Aligarh.

Oedaleus abruptus is a polyphagous pest causing damage to a number of cultivated crops similar to that caused by P. infumata in and around Aligarh. Adults are moderate in size, with filliform antennae. Majority of populations show green body colour with a prominent 'X' mark on the dorsum. They have been reported from India (West Bengal, Andhra Pradesh, Orisa, Rajasthan, Tamil Nadu), East Nepal, Myanmar, China, Indo- China, Pakistan, Sri Lanka and Thiland. In both grasshoppers five nymphal instars were found with higher activities at latter hopper instars.

The present work is a preliminary attempt to evaluate their biology, ecology and behaviour. Chromological changes and morphometrical analysis were chosen as tools to estimate the magnitude of behavioural changes and extent of phase polymorphic behaviour exhibited by these acridoids under optimum ecological conditions. The biological parts are related to the life-cycle and various stages while the ecological part gives information regarding population studies and relations with various environmental factors. The morphometrical part deals with the body part measurement during various behavioural changes shown under different ecological conditions and chromoecology of these grasshoppers is studied at different life stages under different experimental conditions to unfold hidden swarm forming tendencies.

The grasshopper population observed was maximum during July-October, probably associated with maximum vegetative growth and favorable climatic conditions during that period (Dwivedi, 1977; Tandon and Khera, 1978; and Hazra, 1984). Since the bionomics, ecology, colour pattern and behaviour of these two very important pests is not known in detail except for a few occasional observations made by several workers, therefore, comprehensive study of biology, ecology, colour patterns, and their behaviour is urgently needed with special reference to the Indian sub-continent.

Shorter life processes were observed at high temperature under crowded condition. Fecundity and rate of development were higher at higher temperature. During hopper development a number of morphological and chromoecological changes of great importance were observed which solved a number of problems, especially the number of hopper instars and their identification sexwise.

t^ ^1.^ Laboratory studies have shown that nymphs reared in isolation show some characteristics of the soHtary phase exhibited by estabhshed locusts. While the crowding together of many young nymphs results in increased activity, which in turn, is associated with the development of black pigments and other characteristics, attributes of the gregaria form of locusts. Further, the high level of activity of the latter is probably promoted by the higher internal body temperature of the black crowded nymphs, which absorb more radiant heat than do the green or brownish nymphs of the same. The factors responsible for gregarization and swarm formation have been investigated. Such studies have been very exciting and may prove to be of great importance in the phase forming behavior of acridoids in general.

The range of colours in acridoids is less extensive than in butterflies or beetles, and as a group they appear rather uniformly coloured in shades of brown (from pale-straw to black), grey and green; such bright colours as red, blue or yellow appear normally on the hind wing or often on hind legs, but seldom on the other parts. Effect of environmental conditions on these grasshoppers led to remarkable changes in their colour patterns therefore, colour of all stages of these two grasshoppers was recorded as proposed by E.J. Clark (1943). Ground colour is due to pigments present in the integument. Streaks and dots are most important components of colour especially during hopper stages, as they remain fewer in adult stage. The description of each colour form is made by formulae, composed of symbols. Clark's notation systems have been so far tested only on a few British grasshoppers and may need modification for more general use. A much more precise method of recording colours and patterns has been developed for an objective investigation of colour variation in hoppers of Schistocerca by Stower (1959), depending on the instars and environmental factors, which become the source of inspiration for this arduous task.

The colour of each region was determined exactly by comparing the colours with that given in the plates. Color Dictionary by Maerz and Paul (1950), which not only make it possible to record their corresponding names and indices but also rectified errors in the visual perception made during observations.

Only head and thorax were taken into account as they exhibit prominent colour patterns. Changes in colour patterns were recorded under light microscope to differentiate minor changes exhibited under different ecological conditions.

Morphometrical studies were supposed to be most reliable parameter since time to identify phases in locust. The solitary phase is characterized in its nymphal instars by being variable in colour, green, grey or brown and similar to the colour of its environment, in the adult state, the pronotum is longer and crested, while hind femur is relatively long compared with the fore wing. Under crowded conditions, the nymphal coloration is a bold pattern mainly of black and dark brown and the adult has a shorter, saddle-shaped pronotum and a relatively shorter hind femur. Biologically, the most important differences under these two conditions are the higher activity and swarm forming tendencies. This is manifested in the nymphs by their habit of living in large bands during hot part of the day and march from place to place.

The effective control of locusts and grasshoppers, chemical or biological is intimately connected with the detailed ecology and biology of the species, and depends on a comprehensive knowledge of salient features in the bionomics and behaviour of an acridoid. Very little information is available about the biology and behaviour of these two species especially in northern India though they are assuming the status of pests on the major crops. The present study was carried out in order to fill a gap in our knowledge of the ecology, biology and behaviour of these pests. A complete description of all stages of development may be of practical utility, but without a thorough understanding of the relations between acridoids and their environment, effective control measures are not possible. chapteR-2 REVIEW OF LITERATURE

The studies on the locusts and grasshoppers have been carried out since the early twenties. Notable contributions in the field of Acridology concerning the present studies have been made by many acridologists but mostly dedicated to locusts. Very few references on these two grasshoppers are available thereby indicating that least attention was paid especially in the field of cromoecology.

The various modes of copulation and oviposition have been studied by several workers with specific observations made among others, by Katiyar (1952, 1955, 1956b) in Eyprepocnemis roseus, Autarches punctatus, Oedaleus abruptus and Gastrimargus transuersus, Norris (1954) and Hunter-Jones (1960) in Schistocerca gregaria, Hafez and Ibrahim (1958) in Acrida pellucida, Pickford and Gillott (1972) in Melonoplus sanguinipes, and Iqbal and Aziz (1974) in Spathostemum prasiniferum. Uvarov (1966) gave a detailed account of the common mode of copulation in acridoids, the 'riding mode' in Schistocerca gregaria. Jhingaran (1944) recorded another mode of copulation called the lateral mode' in Heteracris capensis. Katiyar (1952) recorded a third mode of copulation 'hanging mode' in Parahieroglyphus bilineatus. Katiyar (1956b) also observed an intermediate mode of copulation between lateral and hanging mode in Oedaleus abruptus and Gastrimargus africanus. Popov (1958) observed in Schistocerca gregaria that copulating females continue to feed, crawl and jump during mating process. Pickford and Gillott (1972) in Melanoplus sanguinipes reported that female aggressiveness; female density and production of chemical attractant by the females are factors, which have a great influence on the selection of a mate. Pickford (1974) studied the reproductive behaviour of the clear winged grasshopper, Camnula pellucida. Gregory (1965a, 1965b) in Locusta and Norris (1954) in Schistocerca gregaria observed the presence of more than one spermatophore in each female of these species which indicate the necessity of repeated copulation. Hunter-Jones (1960) reported that the last copulation in Schistocerca gregaria is the effective one. Singh and Dhamdhere (1984) observed the mating behaviour in rice grasshopper while Riede (1987) made a comparative study of pre-mating and mating behaviour in 25 species of South American grasshoppers belonging to 9 subfamilies.

The type of soil plays an important role in the act of oviposition. It was observed that the females preferred to lay eggs in moist soil. In cases, where the soil is extremely dry female fails to oviposit at all (Norris, 1968; Edwars and Epp, 1965) and sometimes the female dies without egg-laying (Katiyar, 1956a). Katiyar (1955) in Aularches punctatus observed deposition of the eggs in sandy loam soil. Joyce (1952b) reported that the female of Oedaleus senegalensis prefers to lay eggs in wet soil. Pradhan and Peswani (1961) observed egg-laying in Hieroglyphus nigrorepletus in the roots of various shrubs. Chandra et al. (1973) made detailed observations on the selection of soil type for oviposition in the migratory locust (Locusta migratoria) and studied the development of eggs deposited in the different soil types. The detailed process of oviposition has been described by Katiyar (1955, 1956a) in Aularches punctatus and Parahieroglyphus bilineatus, Hafez and Ibrahim (1958) in Acrida pellucida and Iqbal and Aziz (1974) in Spathostemum prasiniferum. 10

Norris (1950, 1952) studied the effect of crowding on the pre- oviposition period and interval of successive egg-laying in Locusta migratoria migratorioides and Schistocerca gregaria respectively. Antoniou and Hunter-Jones (1956) studied the effect of crowding on the fecundity of Eyprepocnemis capitata. Hunter-Jones and Ward (1959) studied the incubation period, instar hoppers and polymorphic behaviour in Gastrimargus africanus giving some effect of crowding on the sexual maturation of females, egg-laying rate, fecundity and viability of eggs. Lautie (1979) observed the fecundity of the females of Locusta migratoria in the absence of fertile males and Baloch (1980) made an observation on the pod crowding affecting viability of eggs in Locusta and Schistocerca. Pickford (1976) studied the embryonic growth and hatching of eggs of Melanoplus biuittatus in relation to oviposition date. Richards and Waloff (1954) studied the arrangement of eggs in the egg-pods of Stenobothrus lineatus and Mecoseth grossus. Ewer (1977) studied the function of the plug of egg-pods. However, Moriarty (1969) observed the decline of egg-laying capacity with the ageing of the female in Chorthippus bruneus. Fresa (1971) observed that the grasshoppers Dichroplus elongates, D. pratensis and D. punctatus were unable to oviposit and die without egg-laying due to the fungus Entomophthora grylli. Petty (1973) has mentioned that the hatching success is associated with the sand structure. An excellent contribution was made by Mahmood and Qazi (1989) in the field of Biological control. They have studied density and parasitization of egg pods of Phlaeoba.

The variation in the size and shape of the egg-pods associated with the available moisture, food of the parent grasshopper, number of egg-pods laid previously as described by Norris (1950) while Hilliard (1959) attributed it to the soil. Khalifa (1957) recorded 11 development of eggs with special reference to the incidence of diapause in the eggs of Eyprepocnemis plorans.

Bernays (1971b) provided some details about hatching process with reference to temperature, moisture and food for Schistocerca gregaria. Temperature, moisture and food have been reported to play an important role in successful hatching (Dempster, 1963). Agrawal and Rizvi (1982) observed an early emergence due to early monsoon in Hieroglyphus nigrorepletus, which is a diapausing species.

Resistance to desiccation among grasshoppers with reference to water loss on the hatchability of grasshopper eggs was studied by Salt (1952) while Pickford (1966b) observed the rate of mortality in extreme dry conditions. Hunter-Jones (1964) observed detrimental effect of high level of soil moisture, which may be due to restricted supply of oxygen. Shulov and Pener (1961, 1963) studied the incubation period in relation to moisture content. Shulov (1956) in Anacridium aegyptium observed the development of eggs extending for more than two months from normal time due to water deficiency. The mortality rate with extreme soil moisture level was recorded by Shulov and Pener (1961) and Harjai and Sikka (1970). Hunter-Jones (1964) in Schistocerca gregaria observed the eggs neither hatched in water-logged soil nor in almost dry soil. The eggs in dry sand die within two days but the water content of the sand in between these two extremes made no difference. Donaldson (1970) observed the difference between the top and the bottom eggs and between the resulting hoppers and adult populations of two strains of Locusta migratoria migratorioides.

The incubation period of eggs in Chrotogonus has been mentioned as inversely proportional to temperature and moisture but moisture affects the viability of developing eggs as noticed by Grewal 12

and Atwal (1968). Church and Salt (1952) studied the normal development of Melanoplus biuittatus even at 12°C, while Hunter- Jones (1964) observed the reduction in hatching percentage at temperature extremes. Parihar and Pal (1978) attached significance to temperature on the development of eggs of surface grasshopper and Chapman and Page (1979) observed the mortality of Zonocerus variegates in Southern Nigeria. Hewitt (1979) laid stress on temperature and precipitation in the environment and their effects on the development of rangeland grasshoppers. Shulov (1952b) observed higher relative humidities as detrimental to one-day-old eggs. However, Shulov (1970) has given the importance of humidity in the development of eggs of Nomadacris septemfasciata and Locusta migratoria migratorioides. Bernays (1972) emphatically attributed the water content of the soil as an important factor in determining the size of hatchling in Schistocerca gregaria. Uvarov (1966) and Bernays (1971a) made a comprehensive study of the form and activity of vermiform larva of Schistocerca gregaria. Papillon et al. (1980) could find hormonal imbalance in Schistocerca gregaria with changing temperature. El-Ibrashy et al. (1985) have made some very useful observations on metabolic effects of juvenile hormones in the female desert locust.

Studies on the effects of different degrees of temperatures and level of humidities on different stages of acridoids have been worked out by many workers like Grewal and Atwal (1968); Parihar (1971); Khan and Aziz (1973a, 1974c); Iqbal and Aziz (1973); Majeed and Aziz (1980a, 1980b) and Ali (1982). These workers have attributed temperature and humidity as an ecological factor responsible for the rate of development and hopper duration periods. Grewal and Atwal (1968) observed decrease in hopper duration of Chrotogonus trachypterus with the increase in temperature and relative humidity. 13

Pradhan and Peswani (1961), while working with Hieroglyphus nigrorepletus, observed that the rearing at 32±1° C was more favourable than at 26±1° C at which the attainment of the adult stage has taken double the time and the development did not proceed at 20° C and 40° C. Similar observations are made by Parihar (1971); Khan and Aziz (1973a, 1974a, 1974c); Iqbal and Aziz (1973) in Poekilocerus pictus, Oedaleus abruptus, Eyprepocnemis alacris and Spathostemum prasiniferum, respectively. Abou-Elela and Hilmy (1977) observed the effects of photoperiod and temperature on the developmental stages of Acrotylus insuhricus.

The effects of different levels of temperature and humidity on the development were studied by Grewal and Atwal (1968) with reference to pre-oviposition and oviposition periods in Chrotogonus trachypterus. Iqbal and Aziz (1973) recorded that the gonads mature earlier at 35° C and the female did not oviposit at 18°C and 45° C in Spathostemum prasiniferum.

In acridids the number of nymphal instars may vary from species to species and even in individuals of the same species (Joyce, 1952b and Katiyar, 1961). However, Hunter-Jones and Ward (1959) in Gastrimargus africanus observed variation in the number of hopper instars in male and female of the same species.

The distribution of grasshoppers in relation to vegetation and physical factors has been observed by Smith (1950); Abushama and Elhag (1971); Iqbal and Aziz (1975) and Moonis and Aziz (1977, 1980). Similar observations were made by Majeed and Aziz (1981b) while working on Gastrimargus transversus with reference to the effect of different food plants on the development of hoppers and their survival. Observations made by Ratan (1978) on the utilization of food by Acrida exaltata are of an informative nature. Khan (1974) 14 in his Ph.D. thesis on bionomics and life-history of some acrididae made preliminary observations on Oedaleus abruptus and its nutritional behaviour and Majeed (1978) in his Ph.D. thesis on ecological factors affecting Gastrimargus transversus had also made some observations on the importance of food plants as an ecological factor. Workers like Williams (1954) and Misra (1962) made some sporadic observations on the food patterns in grasshoppers. Abushama and Elhag (1971) have studied the distribution and food plants selection near Khartoum while Toye (1974) was able to record the feeding and locomotary activities of Zonocerus variegatus and Aziz and Aziz (1985) recorded plant selection pattern in Oxya uelox. Useful studies were made by Bailey and Mukherji (1976) on the feeding habits and food preferences of Melanoplus bivittatus. Mulkern et al. (1969) had given comprehensive account of food habits of grassland grasshoppers of North Central Great Plains. Berbays et al (1974) observed the inhibitory effects of seedling grasses on fecundity and survival of Locusta migratoria migratorioides, Nomadacris septemfasciata, Chortoicetes terminifera, Melanoplus sanguinipes and Schistocerca americana. Misra (1962) observed that Camnula pellucida was able to discriminate nutritionally favourable plants from unfavourable ones. Rizvi and Aziz (1967) recorded the damage to vegetable and medicinal plants caused by Oxya velox. The intra species food preferences were studied by Ba-Angood and Khidir (1975) in Schistocerca americana and Iqbal and Aziz (1975) used twelve different food plants for food preferential values of different stages of Spathostemum prasiniferum.

Smith et al. (1952) and Barnes (1955) studied the survival, fecundity and growth of the grasshoppers. Iqbal and Aziz (1977) observed the effect of different food plants on the development and reproductive potential of Spathostemum prasiniferum. The findings 15 made by Toye (1973) and Bernays and Chapman (1973) on locusts are to be mentioned.

Barnes (1965) studied the effects of different food plants in terms of diet quality and its respective bearings on the health of the grasshopper. Manchanda et al. (1980) observed a new phenomenon related to the effects of host plant on the morphometries and phase status of Schistocerca gregaria while Roonwal (1982) attached a biological significance to the pigmentation in the grasshoppers.

Effect of crowding on the development of grasshoppers and changing behaviour were studied by Norris (1950, 1952) in the African migratory locust and desert locust, using crowded and isolated conditions but could not detect any difference in nymphal duration. Schmidt and Albutz (1999) Identified phases in desert locust, Schistocerca gregaria, by experimental breeding method. Schmidt (2001) excellently examined effect of density on adult Locusta migratoria cinerascens on its behavior, reproduction and morphology in successive generations. Antiniou and Hunter-Jones (1956) found crowded conditions detrimental to the survival of Eyprepocnemis capitata especially in the first instar hoppers. Burnett (1951) studied the life-cycle of Nomadacris septemfasciata in relation to solitary phase while Staal (1961) observed the development of Locusta migratoria migratorioides in relation to crowded conditions. Hunter-Jones and Ward (1959) found rearing density having no effect on the adult morphometries in Gastrimargus africanus. The density among adults of Gastrimargus africanus has no relevance with sexual maturation. This is in contrast to the observations made by Norris (1950, 1952) and Hunter-Jones (1958) in locusts. Norris (1962a) made observations on the density and grouping effect on sexual maturation, feeding and activity in caged Schistocerca 16 gregaria. Gassier (1972) observed the influence of rearing conditions (Isolated and Crowded) on the fecundity of females of Locusta migratoria migratorioides. Papillon (1972) also studied the influence of crowding of adults on their fecundity and polymorphism in their progeny while working on Schistocerca gregaria. Khan and Aziz (1974b) observed the effect of crowding on the hopper developmental periods on Oedaleus abruptus and Eyprepocnemis alacris under controlled conditions. Rizvi et al. (1975) observed the effect of crowding on the nymphal duration of Hieroglyphus nigrorepletus. Majeed and Aziz (1977, 1981c) observed the effect of crowding on the fecundity and viability of eggs and development of different stages under different density at constant temperature and relative humidity. Moonis and Aziz (1978) studied the effects of crowding on the development and fecundity of Trilophidia annulata.

The studies on the phase polymorphism and morphometries of locust were made by several workers. Dirsh (1951, 1953); Roonwal and Nag (1951) and Misra et al. (1952) gave several measurements and ratios for characterizing phases in locusts. Key (1950) made a valuable critique on the phase theory of locusts. Roy (1960) reported Desert Locust in India, New Delhi. Blackith (1957) experimented polymorphism in some Australian locusts and grasshoppers. Stower et al. (1960) also studied the morphometries in desert locust. Ellis (1951) studied marching behaviour of hoppers while Denis et al. (1976) recorded morphometrical changes in Locusta migratoria in relation to density. Ellis (1962) noted behavioural differences of locusts in relation to phase and species. Tanaka (1982) studied the crowding effects on the migratory locust in Japan. Chandra (1983) observed a small concentration of Oedaleus senegalensis in Rajasthan desert, which is of great importance in desert locust populations in relation to habitats. Basit et al. (1984) reported 17 morphometrical changes of significant nature while rearing Gastrimargus africanus. Bellinger and Pienkowski (1987) investigated developmental polymorphism in red-legged grasshopper in USA. Rafeeq and Rizvi (1989) and Razak and Rizvi (1989) have made some observations on the suspected gregariousness in Oedaleus senegalensis and Acrida exaltata, respectively. Ahluwalia, Sikka, and Venkatesh (1976) reported swarming behaviour of Oedaleus senegalensis Krauss.

Environmental conditions play an important role in the grasshopper population structures. It is correlated with food availability, ecological niche and changing temperature with photoperiod. Favourable ecological conditions are attributed to vigorous biological activities. Comprehensive work in this field has been done on different acridoids notably by Richards and Waloff (1954); Katiyar (1955, 1956b); Mac Carthy (1956); Edwards (1960); Pickford (1960, 1966a); Chapman (1962); Riegert and Pickford (1963); Lea (1969); Smith (1969); Nakhla (1970); Pick and Lea (1970) Khan and Aziz (1973a, 1973b); Qayyum and Atique (1973) Descamps (1975); Majeed and Aziz (1975); Moonis and Aziz (1977) Haq and Aziz (1978); Duranton and Lecoq (1980); Ali (1982); Julka et al. (1982); Serjeev and Li (1982); Agrawal and Rizvi (1982); Basit et al (1983); Hazra et al (1984); Materu (1984) and Mital and Chandra (1984). Khan et al (2003) recorded damage in some medicinal plants by the nursery locust, Phlaeoba infumata. These workers have investigated the ecological and biological aspects of various grasshoppers in relation to their habitats, environmental conditions and zoogeographical conditions. Paranjape (1985) gave special emphasis to behaviour analysis to feeding and breeding in orthopteroid insects while Singh et al. (1985) made some field observations on the seasonal abundance of Atractomorpha crenulata. 18

Notable contribution was made by Haider, 1986 in the population ecology of acridids with special reference to the seasonal abundance and sex ratios of Phlaeoba panteli.

The work on the population fluctuations of locusts and grasshoppers, their pattern of distribution and abrupt changes due to competitive relationships had been done by El-Minshawy et al. (1978); Ting et al. (1978); Chapman et al. (1979). Mulkern (1980) and Farrow (1982) investigated population dynamics of Australian plague locust with reference to analysis processes, while in India, Parihar (1983) stressed seasonal variations in population of Pyrgomorpha bispinosa. Haider (1986) studied the population ecology of three acridids in West Bengal. Hugueny and Louveaux (1986) observed specifically aridity gradient and latitudinal variations in size in populations of Calliptamus barbarus while Hewitt and Onsager (1988) studied effect of sagebrush removal and legume interseeding on rangeland grasshopper population. A new approach has been adopted by Johnson and Adla (1988) in studying spatial and temporal computer analysis for grasshoppers in Alberta and is considered a new technology for rapid mapping of variables and their use in computer models.

Agrawal and Rizvi (1982) observed the emergence of Hieroglyphus nigrorepletus just after winters in North India when the species was found throughout India as univoltine and only seen in monsoon season. Jago (1963) describing life-histories Eyprepocnemis plorans in different seasons of the year. Phipps (1968) made observations on ecological distribution and life-cycle of some tropical African grasshoppers. Khan and Aziz (1973b) and Majeed and Aziz (1978) studied the seasonal variation in the population of the hoppers of Oedaleus abruptus and Gastrimargus transversus, 19 respectively. Descamps (1975) made observations on acridid population of the state of Veracruz in relation to climatic conditions.

Some good work on the life-history, life-cycle (laboratory as well as field conditions) and biological characteristics of various species had been produced by Pener and Shulov (1960) for Calliptamus palaestinensis; Dudley (1961) for Locusts; Bhatia and Singh (1965) for Schtstocerca gregaria; Riegert (1967a) for Camnula pellucida; Antoniou and Hunter-Jones (1968) for Eyprepocnemis plorans omatipes; Ibrahim (1970) for Pyrgomorpha conica; Ba-Angood (1976) for Cyrtacanthacris tatarica; Roonwal (1976) for Hieroglyphus nigrorepletus; Moonis and Aziz (1977) for Trilophidia annulata; Antoniou (1978) for Humbe tenuicomis; Baloch (1978) for Ailopus thalassinus; Haq and Aziz (1978, 1979) for Acrotylus humbertianus; Lecoq (1978) for Sudansese acridid; Duranton et al. (1979) for Catantops; Parihar (1979) for Pyrgomorpha; Gunnarsson (1980) for locusts; Ibrahim (1980) for Heteracris; Lecoq (1980) for West African acridids; MacFarlane and Thorsteinson (1980) for Melanoplus bivittatus; Onsager and Hewitt (1982) for rangeland grasshopper; Capinera and Sechrist (1984) for Colorado grasshoppers; Holmberg and Hardman (1984) for six species of Canadian grasshoppers; Chapman et al. (1986) for Zonocerus variegatus; Waloff and Pedgley (1986) for South American locust; Whitman (1986) for Taenipoda eques and Xu and Lu (1987) for Phlaeoba angustdoris. Cherill and Begon (1989) laid special stress on the timing of life-cycle in a seasonal environment with reference to temperature dependence of embryogenesis and diapause in Chorthippus bruneus.

In the last two decades work on acridoids have been carried out with unsequential concepts. Species have been chosen as a model not to be investigated with special reference. Gupta and Vats 20

(1980) have worked out the food consumption, assimilation and tissue growth in Phlaeoba infumata. Das et al. (2002) could record the effect of food plants on the growth rate and survivability of Hieroglyphus banian therein. Similarly Das et al. (2001) had dealt with fecundity and fertility of Oxya fuscovittata.

Nath and Haldar (1992) while working on food preferences have given satisfactory results and they have given a touch to mortality rate in acridoid with environmental cues. Dealing with nutritional ecology, Waldbauer and Friendman (1991) have published a good work on self-selection of optimal diets by insects, which may be applicable to acridoids as well.

Some data on the histological location of pigments in pattern morphs of Chortoicetes terminifera were given by Byrne (1962). Vorontzovskii (1928) in his study of parallel colour variation in forty- eight species of grasshoppers established thirty-two categories of patterns and called them varieties, with suggestive Latin names. Ramme's (1951b) diagrammatically illustrated in colour for twenty- one European species, were not denoted by names, but by one-letter symbols.

Key (1954) studied pattern variation in Chortoicele and Austroiceles, established twelve 'forms', while a complex combination of colour and pattern was proposed by E. G. Clarck (1943). The description of each form is by a formula, composed of symbols, which denote both the general and the detailed distribution of colours. Clark's notations system was simplified by W.Richards and Waloff (1954). A much more precise method of recording colours and atterns has been developed for an investigation of colour variation in hoppers of Schistocerca by Stower (1959), depending on the instars and the environmental and inherited factors. 21

Okay (1948 and 1954) made useful investigations on the changes in colour patterns among the Orthopterans. Zolotarevsky (1930) provided evidence that some features of the colouration of transient adults of Locusta migratoria capita reflected the sequence in its development, depending upon the direction of the density change (i.e. on the history of the population). Some sporadic reports have been published on colouration associated with microhabitat or as behavioural pigmentation depiction or as a cryptic behaviour presentation by Colvin and Cooter (1995), Eterovick et al. (1997), Islam (1998), Konno (1998), and Sword and Simpson (2000). But recently Badruddin et al. (2003) and Khan et al. (2003) have experimentally proved that the acridid colouration under abiotic and biotic factors can be used as bioindicators for the environmental changes.

The first report by Badruddin et al. (2003) on Choroedocus illustris regarding chromo-ecological studies as bio-indicators in polymorphic behaviour appear to be important in the field of the colour patterns of hoppers as initiated by stower (1959) in desert locusts.

During the present studies, experiments have been conducted and observations were made on the colour patterns, such studies may be of great significance and of being bioindicators especially in the case of the grasshoppers which might be of ethological importance to acridologists.

Therefore, looking at the available literature, the present work may prove to be a complete investigation on these two important grasshopper species of the sub-continent. 22

Rizvi et al. (2003) and Khan et al. (2003) have initiated investigations on the biology, biological control and ethology of acridoids. Therefore, the present author has selected to investigate various biological, bioethological and chromoecological aspects of two grasshopper species. chapteR'3 MATERIALS AND METHODS

In order to maintain stock in the laboratory large number of mature adults and immature stages of Phlaeoba infumata and Oedaleus abruptus were collected from different areas of Aligarh (Lat. 27 ° 34' 30" N and Long. 78° 4' 26" E) (Fig. 2). They were kept in insect breeding cages, each measuring 54 cm x 40 cm x 30cm. Three sides of the cages were made of wood while the front side was further divided into two parts, the upper and the lower. The upper part was fixed and made of glass panel, measuring 31x 31cm, while the lower part, measuring 31 x 12cm, formed a wooden window for cleaning the fecal matter (Fig. 40).

A false-floor of wire gaze was provided at the level of glass plate inside the cage. Six perforations, each measuring 4.3 cm in diameters were provided inside the cage to support egg-laying tubes.

The metallic tubes measuring 11cm in length and 4cm in diameter, filled with sterilized sand and moistened with distilled water (8 ml distilled water for every 100 gm of sand) were inserted into the perforations for ovipositions.

These cages were not thermostatically controlled but changing the number and wattage of the electric bulb in the cage could roughly regulate the heat, which served two purposes; heat and photoperiod for normal growth and development. Each cage was provided with a number of sticks for perching and moulting and also for basking. A petridish of water covered with perforated 24

zinc sheet was kept in each cage and refilled as often as necessary, to keep the humidity at the desired level.

In order to obtain eggs, adult males and females of these grasshoppers were also placed in the glass jars, each measuring 21cm X 19cm. Each jar was provided with a cardboard having a circular hole, 5cm in diameter, in the center. A beaker, measuring 5.4 cm X 5 cm, with sterilized sand (wet) was placed inside the hole of the cardboard. The egg-pods were moistened daily according to the requirement.

The hoppers thus hatched were transferred in glass jars (15x20cm) with the help of especially designed aspirator and fed twice daily with fresh leaves of maize (Zea mays) as per the experimentally designed. The open ends of the jar were covered with muslin cloth held with rubber band. Four different combinations were made by using two ecological parameters viz. temperature and crowding. Two of them were reared at 27±1°C both under isolated and crowded conditions. Similarly rest of the two groups reared at 37±1°C both under isolated and crowded conditions. In isolated condition only two individuals, one from each sex was taken, but in case of crowded condition 25 pairs were placed in a glass jar. Two more replicas of each four combinations were run in parallel to replace the experimental jars, if secondary infection leading to mortality. On every stage of development, the morphometries was done for the developmental rate and differences between various biological stages.

The daily temperature, relative humidity and rainfall in the field was recorded from the weather station. Department of Physics, A.M.U. Aligarh, which is hundred yards away from our field 25

laboratory. The incubators were set at 27±1°C and 37±1°C with 70 ± 5% R.H. for above conditions. The egg-pods were moistened daily according to the requirement.

Whole work is divided into four parts, which are as follows.

A. Population studies.

B. Biology.

C. Colour patterns.

D. Morphometries.

Population studies were made by random sampling of these grasshoppers in and around Aligarh thrice a month for three years (2002-2004). Population fluctuations for both hoppers and adult stage were made separately on monthly basis.

Second part is dedicated to their biology in which different life processes were observed at different life stages. During adult stage; pre-copulation period, copulation period, oviposition period, preference of soils, and longevity of adult were studied under different experimental conditions. Dimension of egg pods, length of froth and eggs were measured using vernier dial caliper. At hopper stages; moulting, time taken during each instar, development of alar rudiments, posterior margin of pronotum, formation of eye stripes, and total hopper duration were studied in detailed. Average fecundity was calculated under isolated and crowded conditions. Growth indices were calculated and compared in different experimental conditions. Dyar's low was applied on these grasshoppers using maximum width of head. 26

Third part of the observation deal with the most important aspect of the chromatic changes. The colour changes under stress of different experimental conditions were studied and presented in the form of colour plates. Only head and thorax (both dorsal and lateral) were taken into account to simplify the work. Poster colours were used to prepare the colour plates. Name and codes of colours exhibited by these grasshoppers were recorded using the Dictionary of Colour by Maerz and Paul (1950).

In morphometrical studies sixteen body parts were measured using vernier dial caliper (least count, 0.05 mm) for each instar. Measurements of body parts were taken after five days of moulting of a given instar to neutralize age factor. Rate of increase, differences between means of body parts, differences between means of ratios of different body parts, standard error and standard deviations were calculated at two different temperatures (27±l°Cnad 37±1°C), under isolated and crowded conditions. The data thus recorded were analysed statistically. chapteR'4 OBSERVATIONS

A. POPULATION STUDIES

The population studies of Phlaeoba infumata, and Oedaleus abruptus with reference to seasonal variations in a year are based on numerical strength of all stages of both sexes in the field. In addition to this, special importance was paid to their small-scale movements in breeding grounds. Khan and Aziz (1973b) observed seasonal variation in population of hoppers and adults of Oedaleus abruptus. Phlaeoba infumata is found throughout year and almost all stages could be seen during extreme climate. Normally they are abundant in short grasses like Cynodon dactylon Pers., and during rainy season they enter into tall grasses.

The local open savanna in Aligarh consists of the following grasses: Cyperus rotundus Lin., Paspalum distichum Linn., Seteria glauca (L.) Beauv., Andropogon adoratus Linn., Panicum psilopodium Trin., and Cynodon dactylon Pers., most of them are preferred by P. infumata. Two complete generations in a year with third overlapping generation was recorded. Population of adults and final instar hoppers were of considerable size during the months of September and October, while they dwindled in the month of December and January. The seasonal variations in the population of these species were made on the basis of monthly records for three consecutive years; 2002, 2003, and 28

2004 shown in Figs. 6 and 7. The relative abundance in different months of the year in relation to temperature, humidity, rainfall and meteorological data are shown in Figs. 3, 4 and 5.

These two grasshoppers have never been found totally absent from breeding grounds, but occasional or abrupt depletion in the population has been observed due to sudden changes in the extreme environmental conditions. The sex ratios in the nymphal instars during 2002 -2004 were recorded as 1:1 in first instar hoppers. The ratio oscillates in subsequent nymphal instars but finally in the adult stage favored to females than males, may be due to high mortality among males. In some areas the male population was exceeding the female but very rarely, which generally exceeds in September. Male population dwindles in October onwards, while the female population increases and stabilized in the month of October and November. This may be attributed to the post copulatory exhaustion resulting in higher mortality among males. The stability of sex or survival potential of female sex was more pronounced than the male.

During the years 2002-2004, a comprehensive study for each hopper instar was made to explore the environmental factors influencing movements of these species in and around Aligarh. Plot marking technique was adopted with special reference to available grass plot in the habitat which were divided into three categories namely: 29

1. Grasses between 15-20 inches.

2. Grasses between 5-10 inches.

3. Bare ground with an occasional tuft of long grass.

These plots were marked alternatively and the number of hoppers and adults entering in each area was recorded by random sampling in these plots. It was found that initial hopper instars preferred plot with small grasses, later instars preferred plot with moderate grasses, and adult stage mostly preferred plot with long grasses.

The small-scale movements of the adults were confined to long and short grasses with intermittent and abrupt flights. The movements of these two grasshoppers found to be affected by the day length. It was also critically observed that the male adults entering tall grasses leaving short while females were entering into short grasses as compared to tall grasses. All observations are based on visual inspections during the course of experimentation. 30

B. BIOLOGY

Observations were made on different biological aspects of life cycle under different experimental conditions throughout their life stages to ascertain its preference to the normal growth and development.

I. ADULT

Phlaeoba infumata Brunner shows sexual dimorphism, medium size and robust body, slender; antennae with the joint flattened, ensiform and more filliform: head elongate, face oblique, rugosely punctured or with irregular low regositise. Frontal ridge raised between and without impression near the median ocellus, narrowly sulcate, slightly widened towards the clypeus, in the female sometimes less distinct near the clypeal margin. Elytra and wing well developed and with anterior and posterior margin nearly parallel. Wings about as long as the elytra, apex rounded; hind femora short or more slender, keels finely, but sparsely serrate, keenlobes obtusely pointed. Hind tibiae straight and rounded with 11-12 spines on outer and inner margin; inner spars a little longer than the outer ones. Hind tarsi short; first joint about as long as the two other together.

Male medium size, antennae reaching beyond the hind margin of pronotum, slightly ensiform, fastigium of vertex short, lateral margin parallel, apex obtuse. Pronotum with distinct lateral and parallel margin subgental plate with apex pointed, cercus slander. Female larger than male, antennae more ensiform (Fig. 41). 31

On the other hand, Oedaleus abruptus is medium sized with rugosity of integument and pitted. Fastigium of vertex angular with truncated apex. Antennae filiform and longer than head and pronotum together. Eye more or less oval, pronotum constricted anteriorly and junction between prozona and metazona. Median carina linear, crossed by only posterior sulcus. There is distinct 'X' marking on pronotum which is covered on both arms of prozona and metazona and. Posterior margin of metazona more or less rounded. Cercus conical, apically acutely rounded, subgenital plate short, subconical with rounded apex. Ovipositor short with robust valve, excurved vertically. O.abruptus mainly exhibits three forms of colours in natural population (Fig. 42).

(a) PRE-COPULATION

Pre-copulation period in P. infumata observed was 6-11 days at 37±1°C under isolated condition and 4-10 days under crowded condition while it was found to be 8 - 18 days under isolated and 8 -16 days under crowded condition at 27±1°C. Similarly, pre- copulation period of O. abruptus observed was 4-12 days and 4-10 days at 37±1°C and 4-12days and 4-11 days at 27±1°C under isolated and crowded conditions, respectively. About 10-20% prolonged pre-copulation period was observed at low temperature under isolated and crowded conditions. Similar observed was made for O. abruptus under all experimental conditions (Tables 2 and 3).

(b) COPULATION

The act copulation in both the grasshoppers is usually preceded by more or less elaborate courtship behaviour, which involve brisk movement of antenna in both sexes. Sometimes 32

maxillary palps are touched by each other and last for about an hour. Male sits on the back of the female, clasping her firmly by the first two pairs of legs, the third pair playing no direct part during the whole process, the male abdomen is curved down, its tip is brought under that of the female abdomen is curved, the aedagus extruded and hooks of the epiphallus grasp the female subgenital plate. The aedagus is then introduced between the ventral ovipositor valves of the female. Dorso-lateral or back-to-back postures were observed rarely or at the verge of copulation, as an indication of separation. Copulation period varied from 90-765 min. and 97-248 min. in Phlaeoba infumata and Oedaleus abruptus respectively (Table 1).

The oviposition behavior is characterised by searching and probing for suitable conditions. Since most acridoid laid their eggs in the soil, the digging of a hole is an essential part of the oviposition process. Female raises its body on the first two pairs of legs and arches the abdomen so that it is approximately vertical. Digging results due to alternate opening and closing of the dorsal and ventral valves. As the closed valves are pushed in and then opened widely they force the particles of soil sideways and somewhat upward; this is followed by closing of the valves and a new thrust of the ovipositor. Process of digging continues until the hole is sufficiently deep. Such extension is possible because the abdominal intersegmental membranes in the female are folded and very much more elastic than in the male.

Some observation like period of oviposition and preference of soil were recorded (Table 4). The most preferred soil was mud as 33 compared to sand and Badarpur (coarse grains used in constructions). These observations are very useful as it is an indicative of egg-laying sites of these grasshoppers. Though, these grasshoppers are hypodephic in nature but sometimes epidephic and epiphytic oviposition behaviour were also recorded. The time taken for egg-laying process observed was 7-10% more at high temperature condition than at low temperature under isolated. Same observations were made for cage bred female individuals as they deposit eggs on wood surfaces or wire mesh; most probably due to lack of suitable conditions, age and health status of female (Table 2 and 3).

(d) POST-OVIPOSITION

Post-oviposition period was found to be 7-23 days for Phlaeoba infumata, while 2-7 days for Oedaleus abraptus under different ecological conditions with shortest post-oviposition period was found to be at high temperature under crowded condition (Tables 2 and 3).

II. EGG-PODS AND EGGS

The size of egg-pods decreased gradually and latter egg-pods were about 20-30% smaller in size as compared to first deposited egg-pod after copulation. More or less same observations were made for the length and width of eggs-pods and eggs under all ecological conditions (Table 5).

III. INCUBATION

The incubation period of Phlaeoba infumata and Oedaleus abruptus was studied at two different temperatures. The most 34 preferred and productive temperature was found to be at 37±1°C, and prolonged incubation period was observed at 27±1°C, which is about 50% more in case of Phlaeoba infumata and about 25% more in case of Oedaeleus abruptus. Significant differences were not observed under isolated and crowded conditions (Table 6).

IV. HATCHING AND MOULTING

The excess of watering of the egg pods result in severe mortality to the eggs. High temperature beyond 45°C and low temperature below 19°C were found to be the upper and lower limits for embryonic development and hatching of young one. Percentage of hoppers hatched in case of Phlaeoba infumata at low temperature was 61.16% and 73.98% at high temperature. Oedaleus abruptus show low hatching percentage, 58% and 66.07% at low and high temperatures respectively. Khan and Aziz (1973a) shown Influence of different levels of temperature and moisture on the development and hatching of the eggs of Oedaleus abruptus. The development of eggs per day was higher at high temperature in both grasshoppers. Similar observation was made for average number of eggs per egg pods. The comparative accounts of hatching and molting periods are given in Table 6.

Effect of isolated and crowded conditions on the fecundity of females in both the species was observed. It gives a clear picture about the ability of the female to produce offspring and its vale (birth rate) and hence is of utmost importance in the biological studies of the grasshoppers. The average fecundity among grasshoppers under study was found higher under isolated condition (Table 7), in order to assure their survival under drastic environmental conditions. Effect of isolated and crowded conditions 35 on the survival percentage, daily rate of development and growth index of both grasshoppers were calculated at two different temperatures. Growth index of P. infumata was found to be higher under crowded condition, but in case of O. abruptus it was lower under same condition (Tables 8 and 9).

An accidental observation was encountered, when few fresh leaves of maize left in BOD incubator near the egg laying tubes, stimulated hatching and it was observed that hatching was 200% more when same practice was made to ensure the influence on the hatching. Same observation was made with Cynodon dactylon, but no significant difference was found. Further investigations are needed in this direction to understand its relationship to the host plants.

V. DEVELOPMENT OF HOPPERS

The first instar hoppers, immediately after the intermediate moult, are called hatchings. It has a relatively short and compressed abdomen, but this extends at the end of instars. Similar changes occur during each successive instar. The changes in size are, however, due not to grow, but to the expansion of the intersegmental membranes.

(a) Development of alar rudiments.

The development of tegmina in acridoids has already been considered as an especial feature for distinguishing between hoppers instars. In the first instar, the lower posterior angles of the mesonotum and metanotum do not show any differences but in the second, they become some what extended and punctured, or 36 rugulose; in the subsequent instars they appear as rounded- triangular lobes directed obliquely downwards, with distinct traces of raised longitudinal ridges, which are trachea, later to become axillary veins. After Ill-instar, striking change occurs in both pairs of rudiments that turn on their axes, so that the outer surface become inner and rudiment of the tegmina become covered by those of the wings, both lying dorsally and directed obliquely upward. The mechanism of the reversal is based on the lower of hypodermal cells of the rudiment growing at the greater rate than the upper, thus pushing the rudiment upwards. (Ivanova, 1947). The treachation of the rudimentary tegmina and wings is more clearly discernable after the reversal and reflect their final venation. Further development consists only in growth of the rudiments towards the body axis in V-instars. The phenomenon of the wings reversal was exclusively observed after Ill-instars for both sexes in both the grasshoppers. Influence of crowded condition and high temperature on wing reversal was observed unlike some of the acridoids, where reversal may occur after the lllrd to last hopper instars. Reversal of alar rudiments is given in Table 10. Development of alar rudiments of both grasshoppers are given in Figs. 24 and 25.

(b) Development of external genitalia.

Development of terminal abdominal segments and observed in these grasshoppers are more or less similar with the slight differences. Comparative descriptions on the changes of the terminal abdominal segments during the development in both sexes are as follows:

In 1-lnstar male subgenital plate was semi-circularly excised at apex in both the grasshoppers, with short obtuse lobes covering 37 only base of paraprocts. Subgenital plate narrowed to apex, which has a shallow excision and reaches middle of paraprocts in II- Instars which become broadly parabolic apex, reaching beyond middle of paraprocts in Oedaleus abruptus but in Phlaeoba infumata excision was deeper. Subgenital plate in IV-lnstars was narrowly parabolic and reaching almost to apex of paraprocts. In Phlaeoba infumata subgenital plate was elongate-parabolic, much longer than paraprocts, pointed at apex but in Oedaleus abruptus it was comparatively shorter.

In I-instar female upper ovipositor valves were short, triangular sub-acute, separated by a broader excision. Lower ovipositor valves represented by a transverse fold (distinct only towards end of the instar) at bases of upper valves.

In 11-instars upper ovipositor valves were longer, subacute, separated by a broader excision. Lower valves were broadly triangular, separated by acute angular excision in both grasshoppers.

In Ill-instars upper ovipositor valves were broader, separated by a broader excision. In Phlaeoba infumata upper ovipositor valves were narrower, separated by a narrower excision. Lower valves were almost as long as upper valves. Left and right ovipositor valves in IV-instars were separated along its middle line. Upper and lower valves were longer than paraprocts. Upper and lower valves sclerotised and well-developed ovipositor is formed in last hopper instars (Fig. 26 - 29). 38

The changes in posterior margin of pronotum during hopper development can also be used to identify hopper instars especially in initial stages when alar rudiments and changes in last abdominal segments are not prominent or difficult to percieve. General and comprehensive changes during hopper development of these two grasshoppers are as below.

I-instar hoppers exhibited more or less right angle to acute posterior margin of pronotum as its median carina is much shorter than lateral

ll-instar hoppers are characterised by obtuse posterior margin of pronotum as median carina become elongated.

lll-instar hoppers exhibited shallow or concave posterior margin of pronotum because median carina and lateral carinae show minor difference thus curved posterior margin was produced, which is pushed inward (Fig. 43b).

IV-instar hoppers median carina and lateral carinae are almost equal in length making posterior margin almost straight or slightly pushed outward as convex margin.

V-instar hoppers show greater median carina than lateral making more convex margin or obtuse.

More or less similar observations were found in both the grasshoppers except during IVth and Vth instars, which exhibited more acute hind margin of pronotum in case of Oedaleus abruptus 39 as compared to Phlaeoba infumata. Median carina is little longer than lateral carinae that form an obtuse posterior edge (Fig. 23).

(d) Formation of eye stripes.

Formation of eye stripes observed carefully at each hopper instar under light microscope in both the grasshoppers. In case of Phlaeoba infumata mainly two types of eye stripes were observed; a dark stripe along the anterior margin of compound eye or with a transverse stripe along the horizontal diameter of compound eye. But in case of Oedaleus ahruptus more prominent and numerous eye stripes was observed. Changes during hopper development at each stage was depicted and diagrammatically presented. Scattered patches were observed through out the compound eye in I-instars, which arranged in Il-instars making discontinues eye stripes but only toward the anterior margin of compound eye. In successive instars these discontinuous eye stripes merged to produce continuous eye stripes. Anterior stripes completed first than posterior, thus number of continuous and discontinuous vary at each instar, which can be used to distinguish hopper instars (Fig. 22).

Effect of ecological conditions on the formation of eye stripes was less pronounced and only delayed eye stripes formation was observed at 27±1°C, reduction in eye stripes was not observed under any ecological conditions. In case of Phlaeoba infumata, only one or two eye stripes were observed about middle of the compound eye along horizontal diameter of eye.

The most important aspect of hopper development as observed by many acridologists is the variation in the number of 40 hopper instars and hopper duration under varying ecological conditions. The present study shows no such variation in the number of hopper instars but the temperature along with isolated and crowded condition has certain impact on hopper duration, i.e. both males and females have five hopper instars in all experimentally set ecological conditions in the laboratory; however, the hopper duration varied quite considerably. In case of Phlaeoba infumata minimum male hopper duration was observed to be 58.000±1.135 days (crowded conditions) at 37±10C and the maximum was 82.90011.433 days (isolated conditions) at 27±10C, where as minimum female hopper duration was observed to be 57.500±1.258 days (crowded conditions) at 37±1°C and maximum as 85.60011.266 days (isolated conditions) at 27lloc. Similar observation was made for Oedaleus abruptus but its hopper duration was relatively shorter than Phlaeoba infumata. It was noticeable the hopper duration in both males and females decreased under crowded condition and at 37110C. (Tables 2, 3).

Moreover, the nymphal instars of both the grasshoppers show variations under different experimental conditions. Initial hopper instars were taken 8-12 days, but among later instarslO-17 days were observed to moult (Figs. 8, 9 and 10, 11).

The moulting from one hopper instar to the next is essentially a biological repetition. The hopper climbs a plant or twig and hangs head downward thus utilizing the gravity in the process but it was notable that impartial moultings do occur, when there is over crowding in the cages. Such impartial moultings lead to various deformities, which are manifested in various morphological changes 41 but having no adverse effect on the reproductive potential. This is a new observation hitherto unknown.

The Dyar's law was applied in both grasshoppers and found progressive development at successive formation of instars. The measurements of head width of the successive instars were made separately in both sexes and within the same sex.

The head width of each instars increases in a geometrical progression. In case of Phlaeoba infumata the average ratio of increase was higher at high temperature under crowded condition for both male and female populations. In case of Oedaleus abruptus the average ratio of increase was higher at low temperature under isolated condition for male population, but average ratio of increase was found to be higher at high temperature under isolated condition in female population. The calculated head width is found close to the observed head width. Majeed and Aziz (1979) successfully applied Dyar's Law in different hopper instars of Gastrimargus transverses. These figures are enough to determine the instars and eliminate any possibility of missing an ecdysis in the life-cycle. Although the above figures are not completely identical in nature but these are sufficiently close to infer that the increase in head width follows Dyar's law (Tables 11-14).

Another aspect of crowded and isolated conditions and their impact on various biological processes are shown in for Phlaeoba infumata and Oedaleus abruptus, inferring that a change is found in pre-copulation and post-copulation periods. Both these species are found tending towards behavioural changes when subjected to two different conditions. The deviations are of much importance and are indicative of having gregarious tendency since times in the above grasshoppers. 42

C. COLOUR PATTERNS

There is a characteristic feature in all acridids to show drastic colour variations in response to the environmental conditions as behavioural changes. Keeping this in view, observations were made to know the factors responsible for this colour pattern exhibition and variation therein. Visible colours of insects are due either to the structure of the integuments or the pigments in it. Structural colours of insects are mostly metallic or lusture in appearance and these are very rare in acridoids. The rang of colours in acridoids as a whole is in shades of brown (from pale-straw to black), gray and green, while bright red, blue or yellow appear normally on the hind wings and often on the hind legs, but seldom on other parts. Completely uniform colouration, however, is not a general rule, and disruptive patterns, composed either of different colours or of contrasting shades of the same colour, are common. A different and more comprehensive method of colour notation was used to describe the colour of head and pronotum (both dorsal and lateral views), as these body parts exhibit more prominent colours and patterns in these two grasshoppers.

Broadly there are three main components of the colour patterns in the hoppers of the P. infumata; ground colour, streaks and dark spots. Ground colour varied from light brown to carbon black under different ecological conditions. In case of O. abruptus ground colours varied from light green to brown or even dark brown in colour. Streaks in case of P. infumata are confined on the head and pronotum (Fig. 43a). It may be prominent to faded depending upon ecological conditions. Dorsal streaks were observed about the 43 mid dorsal line of vertex, which are parallel to each other and perpendicular to the mid dorsal line on the vertex. Alternate shorter and longer streaks of vertex give an impression of a characteristic DNA-helical. Streaks were very prominent in case of O. abruptus especially on the dorsum and lateral arms of the pronotum. Dark spots like fine and coarse dots were observed throughout the hopper stages. Maximum number of dots was observed during second hopper instars, but it frequencies decreased during successive instars, and on the onset of adult stage only few dots was exhibited that to the specialized area of the integument. Reduction in the number of darks spots during hopper development suggested accumulation of few fine dots leading to the formation of coarse spots within the hopper instars.

A major difficulty with the existing classification of colour patterns was standardization of colour descriptions for the interpretation and comparisons of the subjective terms used in colour description. The colour types were standardized, coded and named for all hopper instars and adult males and females of P. infumata and O. abruptus by using A Dictionary of Colour by Maerz and Paul, 1950, London.

The colour changes of head and prothorax (pronotum) of these grasshoppers were arrested and drawn on the line diagrams with all possible care (Fig. 44-57), although the colour drawn may not be in close accuracy with the colour of different stages though all possible care were taken to rectify the errors in observations. Name and codes of colour components exhibited under stress of different experimental conditions were given in (Tables 15-28). Minor changes in colour patterns were observed for two sexes at a 44 given hopper stage, moreover female is biologically and ethologically more important than male, therefore female individuals were chosen as model for colour depiction. In case of adult both sexes were considered due to appreciable colour changes under different ecological conditions. The colour patterns observed in these two grasshoppers are not based on individual but a model colour patterns was recorded which the majority exhibits.

I. Description of colour patterns in Phlaeoba infumata

(i) 1st INSTAR HOPPERS

Dorsal view

(a) HEAD

No mark differences were observed in the ground colour under all ecological conditions. A slightly different colour, Italian straw (11D2) and Cockatoo (10H2) were observed at 27±1°C and 37±1°C respectively. Similarly, colour of streaks observed as Bonito FUSCOUS+ (7C7) at 27±1°C, while Natal Brown (7A10) at 37±rC. Colour of dots were similar for both high temperatures and low temperatures. Influence of crowded condition was not appeared.

(b) PRONOTUM

Only Italian Straw (11D2) as ground colour was observed under all ecological condition. Similarly, sunstone (12F12) was observed for dots. 45

Lateral view

(a) HEAD

Pineapple+ (12J2) was most prominent ground colour with a slight change at 27±1°C under isolated condition.

(b) PRONOTUM

Again Pineapple+ was most prominent ground colour in all ecological conditions provided. Colours of dots were same through out the instars under all ecological conditions.

(ii) Ilnd INSTAR HOPPERS

Dorsal View

(a) HEAD

Slight change in ground colour was observed at two different temperatures. Colour was more prominent at 37±1°C (Seminole- 14E10), while less intensified at 27±rC (Airedale-14F6). Colour of streaks were found to be Cub-15Cl for each experimental conditions except at high temperature and crowded condition, where colour was more intense (English Grey-15C2). In case of dots two slightly different colours were observed at tow temperature, while influence of crowded condition was not prominent.

(b) PRONOTUM

Background colour observed was ACORN-15E7 at high temperature. Colour of streaks were almost similar except at 37±1°C under crowded condition which was observed as brown suger-15Hll. Colour of the dots was Sooty Black under all 46 ecological conditions, while Slate Black+ (14C6) was observed at high temperature under crowded condition.

Lateral view

(a) HEAD

Cloudy Amber (12K3) was observed at 27±1°C under isolated condition. Ship skin Moth+ (11C3) was observed in rest of the ecological conditions. There were no mark differences in the colour of compound eye with varied ecological conditions. Colours of the dots observed were similar as mention above in dorsal view of head.

(b) PRONOTUM

The ground colour observed was Bronze Clair (13D2) at 27±1°C under both isolated and crowded conditions. Same observation was made at 37±1°C and isolated conditions. Colour of dots was more or less similar to that of the dorsal view of pronotum.

(iii) Ilird INSTAR HOPPERS

Dorsal view

(a) HEAD

OLIVE GREEN (15L4) and Metallic Green T (15L6) were observed at 27±1°C and 37±1°C respectively. No mark difference in colour was observed under crowded and isolated conditions. Also there were no changes in the colour of streaks under any ecological condition, since GRAPHITE+ (48C7) was observed under all studied ecological conditions. 47

(b) PRONOTUM

Roman Green (15L5) was observed at 27±1°C while Serpentine (14K3) was observed at 37±1°C as ground colours under isolated as well as crowded conditions. Streak were uniformly limestone (15A4), under all ecological combinations, similarly colour of dots were also uniform.

Lateral View

(a) HEAD

Two different ground colours were observed at 27±1°C, both under crowded and isolated conditions. But at high temperature similar colour (Sallow-12E2) was observed, under crowded and isolated conditions. Colour of compound eye was similar, both at high and low temperature under crowded and isolated conditions. Similar observation was made at 37±1°C under crowded condition, which change to Sirocco (14B2) under crowded condition.

Two different colours, Hay-1212 and ACRON-15E7, were observed under crowded and isolated conditions at same temperature (27±1°C). Similar observations were made under crowded and isolated conditions, at high temperature (37±1°C).

(b) PRONOTUM

Two slightly different colours, 01iveshen-13K3 and Cloudy Amber 12K3, were observed at 27±1°C under crowded and isolated conditions, but at high temperature colour was comparatively darker than the low temperature. Colour of dots on the head and 48 pronotum was similar as observed under dorsal view for the same in star.

(iv) IVth INSTAR HOPPERS

Dorsal View

(a) HEAD

Different ground colours were observed at two different temperatures, but similar colour was observed under crowded and isolated conditions at same temperature. Similar observations were made for streaks and dots.

(b) PRONOTUM

Ground colour was similar at low temperature, Acacia (llKl), both under crowded and isolated conditions. Slight darker ground colour was observed at high temperatures under crowded condition. Colour of streaks was similar under all ecological combinations, except at high temperature under crowded condition, and observed colour was Army Br (6A10). Colour of dots was slightly darker at high temperature under crowded and isolated conditions.

Lateral View

(a) HEAD

A gradient of colour changes were observed at all above experimental conditions. At low temperature, colour was less prominent, while at high temperature it was dark. Two different colours of streaks were found at low temperature, but at high temperature same colour was observed under both crowded and 49 isolated conditions. Two different colours were observed at 27±1°C, under crowded and isolated conditions. Similar observations were made at high temperature with slightly intensified under crowded condition. Colour of compound eye was similar, Manon (6A9), at low temperature and at high temperature under isolated condition. Slightly darker colour was observed at high temperature under crowded condition.

(b) PRONOTUM

Ground colour observed at low temperature was sheepskin Moth+(llC3), under both crowded and isolated conditions, while Cloudy Amber (12K3) was observed at 37±1°C. Colour of Streaks was found uniform, except at high temperature under crowded condition. Colour of dots observed was similar to that of the dorsal side of pronotum.

(v)Vth INSTAR HOPPERS

Dorsal View

(a) HEAD

Two different ground colours were observed at low temperature, while at high temperature prominent but similar colour was observed under both crowded and isolated conditions. Colour of streaks observed was Moose (8C10) under each experimental condition, except at low temperature and isolated condition. Colour of dots was uniformly Slag (48A4) under all above combinations. 50

(b) PRONOTUM

Two different colours (Roman Gr-15L5 and CITRINE-14L6) were observed at low temperature and under isolated and crowded conditions. Colour changes were not observed at 37±1°C under crowded and isolated conditions. Colour of streaks was different at two temperatures, but similar, under crowded and isolated conditions. Colour of dots (Brownzesheen-12J5) observed was uniform under all said combinations.

Lateral View

(a) HEAD

The ground colour observed was Prairie+(13F6) at low temperature under crowded condition and also at high temperature under both crowded and isolated conditions. Clour of streaks observed were Pelt (15C4) at high temperature under crowded and isolated conditions. Similar colour was observed at 27±1°C, only under crowded condition. Colour of dots observed was SUNSET (10C4) under most of the combinations except at low temperature and isolated condition. Colour of compound eye was slightly intense at high temperature under crowded condition, while under rest of conditions; similar colour (LIGHT STONE-12J5) was observed.

(b) PRONOTUM

Two different ground colours were observed at two different temperatures, but similar colour was observed under isolated and crowded condition at each temperature. Colour of streaks observed 51 was mainly SUNSET (10C4). Colour of dots observed was similar to that of pronotum.

(vi) ADULT MALE

Dorsal View

(a) HEAD

Similar ground colour was observed at low temperature, under crowded and isolated conditions, but two different colours were observed at 37±1°C under crowded and isolated conditions. Colour of streaks observed was Hair Brown (15A4) at low temperature under crowded condition and also at high temperature under both isolated and crowded conditions. Colour of dots was uniform under all each conditions.

(b) PRONOTUM

Two different colours. Brown Surger (15H11) and Bronze Brown (16C9), were observed at low temperature under isolated and crowded conditions, similar colour was observed at 37±1°C under both isolated and crowded conditions. Colour of streaks was similar to that of the head. Two slightly different colours, BURNET UMBERP-15A12 and COCOA BROWN+, were observed at 27±rC and 37±1°C respectively.

Lateral View

(a) HEAD

The ground colour observed was BEAVER (15A6) at 27±rC, but slightly darker colour (OLD GOLLD-14K5) was observed at 52

37±1°C under crowded condition. Colour of streaks observed were SEAL (8E10) at low temperature, under both isolated and crowded conditions; Java Brown (8J8) was observed at high temperature under crowded condition. Colour of dots was similar observed in dorsal view of head. Colour of compound eye observed was Bronze Clair (13D2) under all experimental conditions, except at low temperature and isolated condition.

(b) PRONOTUM

Two different ground colours were observed at two different temperatures, further difference in colour was observed under isolated and crowded conditions. Colour of streaks observed was similar to that of head and colour of dots were as observed under dorsal view of pronotum.

(vii) ADULT FEMALE

Dorsal view

(a) HEAD

Ground colour observed was similar at 27±1°C under isolated and crowded conditions, but two different ground colours were observed at 37±1°C under isolated and crowded conditions. Two different colours, NED COCOA-7A10 and VANDYKE BROWN, were observed at low temperature under isolated and crowded conditions, but similar and intense colour (Cameo-6F9) was observed at high temperature, both under isolated and crowded conditions. Two different colour of dots was observed at low and high temperature, both under isolated and crowded conditions. 53

(b) PRONOTUM

The ground colours observed were less prominent at low temperature under isolated condition than the high temperature. Colour of streaks observed was similar at low temperature, but at high temperature two different colours were observed under isolated and crowded conditions. Colour of dots observed was more or less similar both at low and high temperatures, under both isolated and crowded conditions.

Lateral view

(a) HEAD

Two different ground colours were observed at low and high temperatures, both under isolated and crowded conditions. Colour of streaks observed at high temperature was darker than at low temperature. Colour of dots was similar to those of dorsal side of head. Colour of compound eye observed was darker at high temperature as compared to low temperature.

(b) PRONOTUM

Similar ground colour was observed at low temperature under crowded condition and also at high temperature under both isolated and crowded conditions. Colour of streaks and dots were similar as observed under dorsal view of pronotum. 54

II. Description of colour patterns in Oedaleus abruptus

(i) 1st INSTAR HOPPERS

Dorsal View

(a) HEAD

The ground colour observed at low temperature was Italian Straw (11D2) and Mustard (11J4) under isolated and crowded conditions. The ground colour observed at high temperature was similar as observed at 27±1°C under crowded condition. The colour of post ocular patch was similar under all ecological conditions.

(b) PRONOTUM

Similar ground colour, BUFE (11K7), was observed at low temperature under both isolated and crowded conditions. Similarly, Laurel oak (7J10) was observed at high temperature under both isolated and crowded conditions. Colour of mid dorsal region observed was green OLIVE GREEN (15L4) at low temperature and Beech (15E4) at high temperature respectively.

Lateral View

(a) HEAD

The ground colour observed was uniformly Hay (1212). Streaks not appeared and dark spots (dots) were less pronounced among 1st Instar hoppers.

(b) PRONOTUM

Change in ground colour was not observed while streaks and dots were fewer and less prominent. if* i 55

(ii) Ilnd INSTAR HOPPERS

Dorsal View

(a) HEAD

The ground colour observed was uniformly Glass Gr. (18D3) under all ecological conditions except at high temperature under crowded condition. Two different colours of post ocular patch observed were Rubber (15H8) and Bison- (16A10). Colour of dots were similar as observed in 1st instar.

(b) PRONOTUM

Two different ground colours, OLIVE WOOD (15E10) and Elk Lama- (16A11), were observed at two different temperatures and under isolated and crowded conditions. Similarly two different colours of mid dorsal region were observed under said conditions. Colour of dots and its frequencies were similar as observed in 1st Instar.

Lateral View

(a) HEAD

The ground colour observed was uniformly Turtle Gr. (19G5) except at low temperature and under isolated condition. Colour of streaks was less prominent and uniformly Cub (15C1) under all ecological conditions. Colour of dots was similar as observed under dorsal view of head. The colour of compound eye observed was Oyster Gy (19A2) under all ecological conditions. 56

(b) PRONOTUM

Two different ground colours, Amber white (llCl) and English Grey (15C2), were observed at two different temperatures under both isolated and crowded conditions. Colour of streaks observed was uniformly Shadow Green (20J3) under all ecological conditions. No change was observed among dots.

(iii) Ilird INSTAR HOPPER

Dorsal view

(a) HEAD

The ground colour observed was uniformly OCEAN GR (18B5) except at low temperature under isolated condition. The colour post ocular patch observed at low temperature was Chipmunk (13L9) while at high temperature it was observed as tortoise shell (14G11), similar colour was observed at low temperature under crowded condition.

(b) PRONOTUM

Two different colours were observed as Mustard Br+ (14D10) and Cognae (14J11) at low temperature under isolated condition and at high temperature under both isolated and crowded conditions. Colour of mid dorsal region was slightly darker at 37±1°C under both isolated and crowded conditions. Colour of dots was similar as mentioned in Ilnd Instar but it was more prominent in given instars. 57

Lateral view

(a) HEAD

The ground colour observed was darker at 27±1°C under crowded condition and at 37±1°C under both isolated and crowded conditions. Colour of streaks was uniformly Cub (15C1) under all ecological conditions. Colour of dots and magnitude of intensity were similar as observed under dorsal view of head.

(b) PRONOTUM

The ground colour was almost similar under all ecological conditions except at high temperature under crowded condition, which is slightly darker (SMOKE BROWN-16A2). Colour of streaks was uniformly Corydalis Or. (19B4) under all ecological conditions. Colour of dots was similar as observed under dorsal view of pronotum.

(iv) IVth INSTAR HOPPER

Dorsal view

(a) HEAD

Influence of crowded condition was not observed but effect of temperature was prominent for ground colour. Colour of post ocular patch observed was similar under all ecological conditions. No mark differences were observed among dots as compared to the last instars.

(b) PRONOTUM

Two different ground colours were observed at two different temperatures under both isolated and crowded conditions. The 58 ground colour was darker at high temperature. Colour of mid dorsal region observed was PEA GREEN (20J6) except at low temperature under isolated condition.

Lateral view

(a) HEAD

Two different ground colours observed were, Fern Gr (21A5) and Mignon Gr (21J6), at low temperature under isolated condition and at same temperature under crowded condition. Similar colour was observed at high temperature under isolated and crowded conditions. Colour of streaks observed was more or less similar under all ecological conditions. Slightly dark colour of compound eye was observed at high temperature under both isolated and crowded conditions.

(b) PRONOTUM

The ground colour observed was Rodent (16C3) at low temperature under isolated and crowded conditions, similar colour was observed at high temperature under isolated condition, but at high temperature colour get intensified. Two different colours of streaks were observed at two different temperatures but influence of crowded condition was not observed. Mostly coarse dots were observed through out the body.

(v) Vth INSTAR HOPPER

Dorsal view

(a) HEAD

Two different colours were observed at two different temperatures, slightly darker at high temperature. Influence of crowded condition was not observed on the colour of post ocular 59

patch but effect of temperature was prominent at two different temperatures. Mostly coarse dots were observed.

(b) PRONOTUM

Two different ground colours were observed at 27±1°C, but similar colour was observed at 37±1°C under both isolated and crowded conditions. Similar observations were made in case of mid dorsal region. Size and colour of dots were similar as above.

Lateral view

(a) HEAD

The ground colour observed was Mignon Gr. (21J6) at low temperature under crowded condition and also at high temperature under isolated and crowded conditions. The ground colour was less prominent at low temperature under isolated condition. Colour of streaks was uniform under most of the conditions except at low temperature under isolated condition. Colour of compound eye observed was Hemp (14H4) at 27±1°C but two slightly darker and different colours were observed at 37±1°C under isolated and crowded conditions.

(b) PRONOTUM

Influence of two different temperatures was observed for ground colours, but affect of crowded condition was not prominent. Colour of streaks found to be darker at high temperature under crowded condition. 60

(vi) ADULT MALE

Dorsal view

(a) HEAD

Both, affect of temperature and crowded condition were observed on the ground colour, thus gradient of colours were observed at two temperatures under both isolated and crowded conditions. Colour of post ocular patch observed was Grasshopper (20F6) under all ecological conditions. Very few dots remain at adult stage on the specific areas of integument.

(b) PRONOTUM

The ground colour observed at high temperature was more prominent than at low temperature. Colour of mid dorsal region doesn't show much difference under any ecological conditions. Lateral view

(a) HEAD

Two different ground colours were observed at two temperatures, but influence of crowded condition was not observed. Similarly colour of streaks was more or less same at two temperatures with no influence of crowded condition. Colour of compound eye observed was similar at low temperature under isolated and crowded conditions. While two different colours were observed at high temperature.

(b) PRONOTUM

Effect of temperature was more pronounced for ground colour than the crowded condition, which is negligible for both 61 temperatures. Two different colour of streaks was observed at two temperatures, later was prominent and darker.

(vii) ADULT FEMALE

Dorsal view

(a) HEAD

Different ground colours were observed under different ecological conditions is indicative of the influence of both temperature and crowded condition. Colour of post ocular patch observed was similar at low temperature, but two different colours were observed at high temperature under isolated and crowded conditions, which suggest the influence of crowded condition. Dots were negligible under all conditions.

(b) PRONOTUM

The ground colour observed was almost similar under all ecological conditions except at low temperature under isolated condition. Colour of mid dorsal region observed was different under all ecological condition. Effect of both temperature and crowded condition were prominent. Number of dots and its colour was similar as observed in male.

Lateral view

(a) HEAD

Two different ground colours were observed at high and low temperatures, but effect of crowded condition was not prominent. Colour of streaks observed was similar with exception at low temperature under isolated condition. 62

(b) PRONOTUM

Two different ground colours were observed at two different temperatures, again effect of crowded condition was not observed. Colour of streaks was similar at high temperature but two different colours were observed at low temperature. 63

D. MORPHOMETRICS

When the phase theory was first proposed by Uvarov (1921) for Locusta migretoria migratorioides (L.), he distinguished phase gregaria from phase solitaria by two morphometries indices; ratio of length of hind femur to the elytron length, and ratio of maximum pronotum with its length. Zolotarevsky (1930), in his work on Locusta migretoria capito (Sauss), used the same measurement, but reverse the ratios, and introduced two new measurements- maximum widths of head and pronotum height. Among these ratios the elytro-femoral ratio (E/F) has been most commonly used in studies on the morphological phase variation in different locust species and in recording the morphometries characters of populations. Similar observation was made by Rizvi and Basit (1990) for North Indian grasshoppers and described them as occasionally swarming grasshoppers.

The present work is based on statistical treatment of the various measurable characters by which a hidden tendency of phase polymorphism was evaluated. For this purpose sixteen body parts were measured and ten ratios for above body parts were taken. Measurement of different body parts and its rate of increase in various instars of P. infumata and O. abruptus were calculated at two different temperatures under isolated and crowded conditions (Tables 29-44).

The accounts of body parts measurement are given below. The measurements of various body parts were taken with the help of Vernier dial caliper (least count, 0.05 mm). All measurements were taken in centimeter. The measurements were made on 30 males and 64

30 females at two different temperatures under isolated and crowded conditions. Schematic line diagrams of these body parts are given in Fig. 30.

1.LENGTH OF BODY (L): Distance from the anterior end of head to apex of subgenital plate in male and to apex of ovipositor in female.

2.LENGTH OF ANTENNA (A): The distance from the basal segment, the scape up to the terminal segment.

3.WIDTH OF VERTEX BETWEEN THE EYES (V): The shortest distance between the compound eyes at the vertex.

4.PERPENDICULAR DISTANCE (D): Perpendicular distance between anterior end of head and to width of vertex between the eyes.

S.VERTICAL DIAMETER OF EYE (O): The maximum diameter of eye.

6.HORIZONTAL DIAMETER OF EYE (Oh): The minimum diameter of eye.

7.MAXIMUM WIDTH OF HEAD (C): The greatest width of the head in the genal region.

8.LENGTH OF PRONOTUM (P): The median pronotal carina.

9.HEIGHT OF PRONOTUM (H): The vertical distance between the lowest point of the lateral pronotal lobe and the level of the highest point on the median pronotal carina between the second and third sulci. 65

10.LENGTH OF ELYTRON (E): The branching of costal (mediastinal) and the subcostal (anterior radial) veins to the apex of the elytron.

11.LENGTH OF ANTERIOR FEMUR (AF): The external surface of the femur, directed forwards, from the junction between the trochanter and the femur to the distal end of the latter.

12.LENGTH OF MIDDLE FEMUR (MF): The anterior surface of the femur, from the suture between the trochanter and femur to the apex of the latter.

13.LENGTH OF HIND FEMUR (F): The external surface, as the maximum length from the base to the apex.

14.MAXIMUM WIDTH OF PRONOTUM (MX): The greatest distance between the surfaces of the lateral pronotal lobes in the metazona.

15.MINIMUM WIDTH OF PRONOTUM (M): The minimum distance between the surfaces of the lateral pronotal lobes

16.LENGTH OF HIND TIBIA (T): The anterior surface of the tibia, from the suture between the femur and tibia to the base of first tarsus.

Sixteen boy parts were measured and the rate of increase in various instars was calculated at two different temperatures (27±1°C and 37±rC) under isolated and crowded conditions. Graphic representation of increase in different body parts is given in (Fig. 31-38). Two sexes were taken separately. Among sixteen body parts observed; length of pronotum, length of hind femur, length of 66 antenna and length of elytron exhibited faster rate of growth as compare to other body parts, but over all rate of growth of body parts observed was higher at 37±1°C under crowded condition, least at 27±1°C under isolated condition while intermediate rate of growth was observed at low temperature under crowded condition. More or less similar result was obtained at high temperature under isolated condition. Development rate of female population was always dominated over male population under all experimental conditions.

Differences between means of body parts measurement of crowded and isolated adults of P. infumata and O. abruptus were calculated at two different temperatures under isolated and crowded conditions and found appreciable difference at high temperature under crowded condition. In most of the cases differences of total body length, length of hind femur, length of antennae and length of pronotum were found to be significant. Similarly ten differences between means of ratios under isolated and crowded population for above body parts were calculated, and a number of differences between means of ratios were found to be significant but only at high temperature in female population. At low temperature only one ratio was found to be significant, which is an indicative of hidden swarming tendency of these two grasshoppers. Means, standard errors, standard deviations and mean differences of body parts of adults of both P. infumata and O. abruptus under isolated and crowded conditions are given Tables 45-56.

Differences between means ratios of body parts measurement of crowded and isolated adults of P. infumata and O. abruptus were 67 calculated at two different temperatures (27±1°C and 37±1X) and critically examined the values of student t-test and significance (P)

At low temperature values of 't' oscillated from 3-5 for male but the values for females were 4-6. On the other hand only one or two indices exceeds the value of 3 (significant) in both the sexes (Tables 57-60).

Similar observations were made in case of Oedaleus abruptus. Significant values were more positive in case of female individuals. Very less significant values were obtained at low temperatures for both sexes (Tables 61-64).

Further differences between means of ratios in crowded and isolated adults of P. infumata were calculated and 7 out of 10 ratios were found to be significant both in female and male population at 37±1°C. At 27±1°C, no significant value was obtained.

In case of O. abruptus only 1 out of 10 indices were significant for each sex. At high temperature 2 out of 10 indices were significant for male population but in case of female population 6 out of 10 indices were positive. chapteR'5 DISCUSSION

Very few references of these two economically important grasshoppers are indicative of poor attention of acridologists especially to Phlaeoba infumata, which is a polyphgous pest in North India. The adults of P. infumata inhibit long grasses, dense bushes and cultivated crops; migrate into sugarcane, rice, maize, and vegetable crops for shelter and egg laying. Recently it was found to be a serious pest of a number of medicinal plants in and around Aligarh. Gupta (1980) observed P. infumata Brunn., common in tropical grasslands causing extensive damage to grasses of economic importance during his studies on food consumption, assimilation and tissue growth in Phlaeoba infumata .

The adults of Oedaleus abruptus inhabit short and dry grasses, though it was found periodically as a major pest of millet but also found damaging to most of the cultivated crops with 2-3 overlapping generations a year. Life cycle is completed from 44-82 days depending upon the experimental conditions provided (Tables 2, 3). Khan and Aziz (1973) observed O. abruptus as an important polyphagous pest with two generations a year, and being found throughout the year except during very hot and cold periods. Frequent mating was observed from the end of June to the end of September. Egg- pods of the first generation were laid in June and those the second generation during September-October. Population of 69 the nymphs and adults were largest during August-October, and smallest during December-February.

The population studies of P. infumata and O. abruptus with reference to seasonal variations are based on numerical assessment of all stages through random sampling. Seasonal variations in North India are of prime importance because of open Savanna in Aligarh. The present observations are on the similar pattern, used excellently by Richards and Waloff (1954) in British grasshoppers and by Haider (1986) on seasonal abundance and sex ratios in three species of acridids.

The life cycle of P. infumata and O. abruptus are quite similar in bio-ethological profile except delayed moulting behavior in P. infumata as compared to O. abruptus, resulting there by one more overlapping generations was observed. Xu and Lu (1989) during their study on Phlaeoba angustdoris found it as a pest of bamboo leaves with one generation per year in Zhejiang. Variations in period and time may be attributed to the abiotic regimes experienced by all stages of life cycle. Similar observations were made by Descamps (1953); Uvarov (1966); Rizvi (1985); Lecoq (1991); Marta Cigliano (2000). Seasonal variations in the field population of P. infumata and O. abruptus in different months of the years; 2002, 2003 and 2004 have been recorded on similar pattern, as observed on European grasshoppers, Italian locusts, red locusts and desert locust have been recorded by Richards 85 Waloff (1954) and Uvarov (1966). Adults of these two species are well adapted to all existing ecological changes and were 70 recorded even in December and January but without reproductive activity.

The distribution of acridoids under study may be correlated with the food plants of the area where they were collected. Similar correlation has been established by Anderson 85 Wright (1952) and Iqbal Ss Aziz (1975). Early stages of P. infumata and O. abruptus prefer weeds and grasses and invade crop fields while entering into advance stages of life cycle, may be attributed to different type of nutritional preference in food plants. This has got support from Basit (1990) who has concluded with the same contention in G. africanus and O. abruptus.

Development from hatching to the end of the adult stage take 75.97 days in males and 95 days in females, at high temperature under crowded condition, and the total food intake was found to be higher for female than male. In Phlaeoba infumata 2-3 overlapping generations was observed a year, depending upon experimental conditions. Combined effect of temperature and crowded condition seems to be a threat to its host plants, especially to medicinal plants.

Shorter copulation period, pre-oviposition period, oviposition period and post oviposition period were found at high temperature under crowded condition than at low temperature and isolated condition, indicated clearly the effect of above experimental conditions in the laboratory. Prolonged life processes may be due to lack of optimal conditions necessary for faster growth and development. Similar observations have been made by Srivastava (1957) in 71

Atractomorpha crenulata; Gregory (1961) in Locusta migratoria migratorioides; Hafez and Ibrahim (1958) in Acrida pellucida and Basil (1990) in Gastrimargus africanus and Oedaleus abruptus.

High temperature and crowded condition shifted more to P. infumata than O. abruptus towards the phase gregeria on the scale of phase polymorphism, as they exhibit close behavioral attributes of established locusts all over the world, and appeared to be more significant at high temperature under crowded condition. Therefore, these two species can be designated as swarming grasshoppers. The present observations also revealed that the isolated and crowded conditions affected the average fecundity of female of P. infumata and O. abruptus. Similar observations have been made by Norris (1950, 1952) in Locusta and in Schistocerca, Albrecht (1959) in Nomadacris and Basit (1990) in G. africanus and O. abruptus.

In P. infumata the average number of eggs per egg-pod per female was found to be 37.75 at 27±1°C and 65.95 at 37±1°C. Size of the egg-pod laid earlier was longer than laid latter. Norris (1950) and Ewer (1977) have recorded gradual decrease in size of the egg-pods, if laid many times by a single female in Schistocerca gregaria and Locusta migretoria. In O. abruptus the average number of eggs per egg-pod per female was found to be 26.03 at 27±1°C and 28.76 at 37±1°C. The data suggest that the magnitude of effect of temperature is higher in case of P. infumata as compared to O. abruptus. Almost 100% increment was found in the number of eggs per 72 egg-pod per female in case of P. infumata but slight increment was observed in case of O. abruptus at 37±1°C (Table 5).

Effect of temperature on the development of eggs/day was notable enough to affect even the hatching percentage, which was higher at high temperature. These observations clearly revealed that the number of egg-pods and their size are determined by number of egg-pods laid by these grasshoppers under different ecological conditions. Observation made by Majeed (1978) in G. transverses and by Basit (1990) in O. abruptus and G. africanus are on the same contention and agreement. Attempts made by Ackonor (1988) regarding effect of soil moisture and temperature on hatching and survival in Locusta migratoria was significant and Amatobi (1996) on egg development and nymphal emergence of Kraussaria angulifer in relation rainfall has been supportive to the ecological behavior.

The development of hoppers is of great interest as the acridologists have been reported variations in the number of hopper instars, which normally varried from 4-6 in male and 5-7 in female acridids. The number of hopper instars and the variations therein have been tabulated by Uvarov (1966), in his manual of Locust and Grasshoppers, Vol. 1, and p.286 and attributed to the food of the grasshoppers. There is no variations in the number of hopper instars, exclusively five hopper instars were observed under all experimental conditions both in P. infumata and O. abruptus (Table 10), which is most common figure among acridoids. Similarly reversal of wing observed after third hopper instar in both 73 these grasshoppers, during which the wing rudiments turn on their axes, so that the outer surfaces become the inner and the rudiments of tegmina become covered by those of wings, both lying dorsally and directed obliquely upwards. Any variation in the number of hoppers and occurrence of wing reversal strongly recommend their stability to the different environmental conditions. Basit (1990) and recently Rizvi et al. (2003) have reported similar observations in Acrida exaltata. Roonwal (1952b) stated that most species of acridids pass through 5-nymphal instars in tropical areas especially in North India.

Dyar's law has been successfully applied to hopper instars of P. infumata and O. abruptus as applied by Basit (1990) in Gastrimargus africanus. Calculated width of head of hoppers was found to be different at two different temperatures under isolated and crowded conditions, which is always dominated for female population than the male population. Calculated width of head was observed higher for crowded females at 37±1°C, again indicated a direct impact of crowded condition on the postembryonic development (Table 11-14).

The most important aspects of the present studies have been chromoecology and morphometries in relation to suspected swarming behaviour, and may also help acridologists to evaluate the behavioural changes very easily under different environmental scenerio. Use of light microscope to study chromological changes not only magnified the specimen under study, but also magnified the 74 concept of colour changes throughout the hopper instars and at adult stage. Slight and gradual changes in ground colour, number of dark spots and colour of streaks on the head and pronotum can easily distinguished, thus effect of above ecological conditions can be studied without any ambiguity. Chromoecological studies may not only be used as a tool to investigate phase polymorphic behaviour in acridoids, but also to differentiate hopper instars, which is an important aspect for acridologists, as miss evaluation of instar hoppers will certainly render the results, whether working on any aspect of biological and ethological profiles on acridoids. Similarly chromo-morphological studies like formation of eye stripes, changes in streaks on the head and pronotum, numerical variations in dark spots (fine and coarse dots) and ground colours may prove worth as responses under different ecological stress.

Appreciable changes in colour patterns were observed during chromoecological studies and found fully in agreement with Sword 86 Simpson (2000), Colvin 86 Cooter (1995), Guerrucci 86 Voisin (1988), Eterovick et al, (1997), Islam (1998), Konno (1998), Sobolev (1990), Stebaev (1986), Badruddin et al, (2003) and Khan et al, (2003) while experimentally working on colours as bioindicators and polychromatic behaviour. A variation in colour and its pattern has been recorded from first instar to the adult stage. Observations made by Hunter-Jones and Ward (1959), Rowell (1970), Richard 86 Waloff (1954), Basit (1990) in Gastrimargus africanus, in various British grasshoppers and O.abruptus respectively are of immense value in acridid ethology. Rowell 75

(1970) suggested that the environment determines the coloration in Gastrimargus africanus. Rizvi (1985) had created a concept of chromoecology as a visual gadget in phase polymorphism in acridoids. Some useful observations made by Guerruci and Viosin (1988) on the influence of environmental factors on the colour morphs, by Sobolev (1990) on the cryptic behaviour of locusts, by Garlinge et ah, (1991) on sex-related morphs frequencies in Acrida conica, by Islam et ah, (1994) on parental effects on the colouration, by Suresh and Muralirangan (1995) a colour morphs of Acrida exaltata in the agroecosystem of Tamil Nadu, by Briddle et al, (2001) speciation in grasshoppers with colour pattern and by Schmidt and Albutz (2002) on sex-linked coloration in desert locust have contributed in establishing a new aspect of chromoecology in acridids.

The morphometrical parameters used for the determination of swarming behaviour, have been enhanced to 16 indices and 10 body parts ratios. Most of them have been found significant, mainly at high temperature and under crowded condition, while few parameters do not favored. Differences between means of body part measurements of crowded and isolated adults of Phlaeoba infumata and O. abruptus were calculated at two different temperatures under isolated and crowded conditions and found appreciable changes at high temperature under crowded condition. Out of 16 body parts; differences in pronotum height, length of hind femur, length of tibia, length of antenna and length of pronotum were found to be significant under crowded condition in female population. Ten differences between 76 means of ratios of different body parts calculated and a number of differences between means of ratios were found significant but only at high temperature in female population under isolated and crowded population. At low temperature only one ratio was found significant, which indicated greater susceptibility of female population toward high temperature and crowded condition.

Substantial differences between measurements of body parts under two experimental conditions (isolated and crowded) analyzed critically for each character by calculating the difference between the two means. The standard errors of the mean values of 16 body part measurements were examined and found a mark difference in the said paramertes when compared with their respective means. The standard errors of the means are somewhat higher under isolated condition than the crowded condition for both low and high temperatures. Hence, these morphometrical treatments can be used as reliable estimates for determining the phase polymorphism among acridids.

Occasional gregarization and aggregation in P. infumata and O. abruptus are of great ethological significance. Similar excellent observations have been made by Joyce (1952b), Ghouri 86 Ahmad (1960), Basit (1990) in G. africanus and O. abruptus. Rizvi (1985) made similar observations on few North Indian acridids, and is supported by chromatic changes exhibited by the locusts, explained excellently by Stower (1959). Morphometrical and chromoecology studies, indicated that the two species, though exhibit occasional swarming -'Jlly I. behaviour, but definitely inclined toward s^''^~pffase polymorphism. The fundamental observations made by Tatsute et al, (2000), Schmidt (2001), Rehman et al, (2002) and Colombo (2003) while working on polymorphism in acridid with reference to morphometries has been found fully justified.

Though morphometrical attributes are not very close to the established locusts like Locusta migratoia migretorioides (E/F- 1.79, for phases solitaria and 2.09, for phases gregaia] but other attributes like colour changes, marching activity of hoppers and higher reproductive potential are close to locusts, therefore such grasshoppers may be treated as predecessor to the locust, a phenomenon in action and warning to the acridolgists to take necessary steps for discouraging such acridids to avoid locust plague in future.

The differences in morphometries and colour patterns under isolated and crowded conditions suggested that the species under study have the tendency of unstable gregariousness and may shift eco-ethological niche to tall the number of locust species. Present observations will certainly fill the gaps of the biological information and other aspects, during postembryonic development of these two economically important grasshoppers. chapteR'6 78

SUMMARY

Phlaeoba infumata is one of the most abundant grasshoppers in northern India. Adults are brown to carbon black, medium size, head elongated, face oblique and frontal ridge raised between median ocellus. They are distributed in India (Bihar, Orissa and Uttar Pradesh), Nepal, Myanmar, Thailand, Malaysia and China.

Both adult Phlaeoba infumata and their hoppers are a serious pest of a number of agricultural crops like wheat, sugarcane, millet, maize, rice, jute and economically important grasses, which make them as polyphagous in nature. Apart from these it was found to be a serious pest of bamboo in China and recently reported as a pest of a number of medicinal plants. Adults were found generally in long grasses and immature stages in short grasses. Both adults and nymphs have tendency to aggregate during the hottest part of the day in bushes and crop fields.

The adults of Oedaleus abruptus inhibit short and dry grasses, general body colour green to brown with a prominent white streaks on the dorsum of pronotum, distributed all over the world, and it was found to be a periodically major pest of millet with 3 overlapped generations a year. Under laboratory conditions 3-4 generation was observed depending upon the experimental conditions provided as life cycle is completed from 44-82 days, cause damage to a number of cultivated crops in North India and also reported from different geographical areas throughout the globe especially from African countries, therefore has polyphagous status. 79

Khan and Aziz (1973) observed Oedaleus abruptus as an important polyphagous pest with two generations a year, and being found throughout the year except during very hot and cold periods. Mating occurred from the end of June to the end of September. Egg-pods of the first generation were laid in June and those the second generation during September-October. Population of the nymphs and adults were largest during August - October, and smallest during December-February.

The present work is a preliminary attempt to evaluate the biology, ecolgy, behaviour and population studies especially with reference to North India. Chromoecoloygy and morphometrical characteristics were chosen as tools to evaluate the magnitude of behavioral changes and extent to which polymorphic behaviour by these two grasshoppers by providing ecological conditions gradient. Two different temperatures (27±rC and 37±1°C) and number of hoppers per jar (isolated and crowded conditions), combined to each other, thus four different conditions were obtained. Newly emerged hoppers of both grasshoppers reared at two different temperatures (27±1°C and 37±1°C) under isolated and crowded conditions. During each instar morphometrical measurement, colour changes and their bionomics were recorded and analyzed using suitable statistical treatments.

The population size of these grasshoppers has been estimated with reference to seasonal variations on monthly basis of all stages through random sampling. Both these acridids pests form a common population in North Indian grasslands and cultivated crops, their life-cycles are of similar nature, except flight potential and vigorous activities of hoppers in P. infumata a unique feature as 80 compared to O. abruptus which was found mostly active when crowded otherwise show least activities. It was found that the populations were very high during March 85 April, but most abundant with reproductive activities during rainy season in August 86 September. Similar observation was made by Aziz (1978) for seasonal variation in the population of the hoppers of Oedaleus abruptus and Gastrimargus transuersus, respectively.

The population of hoppers and adult were found higher where grasses like Cyperous rotundas, Seteria glauca, Paspalum distichum, Andropogon adoratus and Cynodon dactylon were dominating the vegetation. The early nymphal instars have preferred Cynodon dactylon while the advance stages migrate to the crop fields. It was found that the distribution of these species is directly related to the preferential value of the food plants available. Adult of these acridids are seen active even in extreme cold of December and January but without reproductive activities

Different processes of life cycle were observed under different ecological conditions and it was found that the least time was taken at high temperature under crowded condition, while at low temperature prolonged biological processes were observed which suggest preference of temperature and higher density for faster rate of development.

Copulation in these two species preceded by elaborate courtship with especial reference to long copulating time and the mechanism involved in oviposition is of the acridian pattern. The egg laying period was shorter at high temperature under crowded condition. Similarly the size of egg-pods, length and breadth of egg 81 and average number of eggs per egg-pod per female was higher at high temperature under crowded condition the viability of egg-pods, eggs and incubation period was found affected by high temperature but influence of crowded condition was not observed. Diapause doesn't observe at any stage of life cycle under different laboratory conditions.

The development of hoppers of these two species is of great interest with reference to number of hopper instars and it was found that under all above ecological conditions five hopper instars were observed in both sexes. The process of alar rudiments reversal was similar in both grasshoppers, which was observed after third instars in both sexes. Moist soil was found to be most preferred breeding ground. Dyar's law was successfully applied in both grasshoppers. The measurements of head width of the successive instars were made separately in both sexes and within the same sex. The head width in successive instars increases in a geometrical progression.

Chromoecology of these grasshoppers were studied under different ecological conditions to know the magnitude of swarming behavior. Only head and thorax were taken into account as they exhibit prominent colour patterns. Out of the three components of colour patterns (ground colour, streaks and dark spots) change in streaks and dark spots were mainly considered as the variations in ground colours may be attributed to its type.

Colour changes were observed visually, using hand lens and under light microscope which not only magnified the object (Dark spots on the head and pronotum) but also magnified the concept of colour and changes during post embryonic development. Only head and pronotum were selected to observe colour changes, as they 82 exhibit prominent colour patterns especially dark spots and streaks. Colour variations were recorded in the form of colour plates under all ecological conditions and deciphered the colour changes using colour dictionary by Maerz and Paul (1950), London. Bold patterns mainly dark brown and black colours were observed at high temperatures under crowded condition. Biologically, the most important difference between the crowded and isolated population was higher activities and tendency of aggregation characterised by higher feeding potential and band formation.

For morphometrical analysis sixteen body parts were taken into account to calculate rate of increase of different body parts, ratios between different body parts, differences between means of body parts measurement, differences between mean ratios of different indices. A number of body parts found to be significant especially length of hind femur, pronotum length and height, length of hind tibia and length of elytron etc. at 37oC under crowded condition in both sexes but female is always dominated over male.

Chandra (1983) observed a small concentration of Oedaleus senegalensis in Rajasthan desert, which is of great importance in desert locust populations in relation to habitats. Rafeeq 86 Rizvi (1989) and Razak 86 Rizvi (1989) have made some observations on the suspected gregariousness in Oedaleus senegalensis and Acrida exaltata, respectively.

These observations certainly leading to accept the fact that both the species are having distinct behaviour of gregarization, swarm forming and having ability of mass active behaviour with local migratory instincts. These observations are the substantial addition to the knowledge of polymorphism in acridoids. 83

On the basis of the present studies it can convincingly be concluded that the species may become gregarious and migratory under suitable ecological parameters. The life-cycle of these species hitherto unknown, have been made known. A complete chromoecological profile, under all considerations, has been completed and the entire research work will be in addition to Entomologist in general and very especial to acridologists the world over. chapteR'7 REFERENCES

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CS 9S !Z5 E CONDITIONS Body Parts 27± 1° C 37± r C Isolated Crowded Isolated Crowded

Dorsol View Head

Ground colour 11 D2 11 D2 10H2 10H2 Italian Straw Italian Straw Cockatoo Cockatoo

Streak 7C7 7C7 7A 10 7A 10 Bonito Fuscous + Bonito Fuscous + Natal Brown Natal Brown

Dots 7H9 7H9 7H9 7H9 Liver Brown + Liver Brown + Liver Brown + Liver Brown + Pronotum

Ground colour 11 D2 11 D2 11 D2 10J3 Italian straw Italian straw Italian straw Amber Y

Streak

Dots 12F12 12F12 I2F12 12F12 Sunstone Sunstone Sunstone Sunstone

Lateral View Head

Ground colour 11 E5 11 J2 11 J2 11 J2 Raffia Pineapple + Pineapple + Pineapple +

Streak

Dots

Compound Eye I4B2 14B2 14H4 14 B 2 Sirocco + Sirocco + Hemp Sirocco + Pronotum

Ground colour 13D6 11 J2 11 J2 11 J2 CRACKER Pineapple + Pineapple + Pineapple +

Streak

Dots

Table. 15. Phlaeoba infumata: 1st Instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. CONDITIONS Body Parts 27± 1° C 37± 1° C Isolated Crowded Isolated Crowded

Dorsol View Head

Ground colour 14F6 I4F6 14E10 14E10 Airedale Airedale Seminole Seminole

Streak 15C 1 15C 1 15C 1 15C2 Cub Cub Cub English Grey

Dots 47 C 6 47 C 6 48 C 5 48 C 5 SLATE SLATE Sooty Black Sooty Black Pronotum

Ground colour 14E7 15E7 15E7 15E7 Stag ACORN ACORN ACORN

Streak 48 A 4 48 A 4 48 A 4 15H11 Slag Slag Slag Brown Sugar

Dots 48 C 5 48 C 5 48 C 5 14C6 Sooty Black Sooty Black Sooty Black Slate Black +

Lateral View Head

Ground colour 12K3 II C3 11C3 II C3 Cloudy Amber Sheepskin Moth + Sheepskin Moth+ Sheepskin Moth +

Streak

Dots

Compound Eye I3C3 I3C3 I3C3 13E4 Lido Lido Lido Sudan Pronotum

Ground colour 13D2 13D2 13D2 I4H4 Bronze Clair Bronze Clair Bronze Clair Hemp

Streak

Dots

Table. 16. Phlaeoba infumata: Ilnd Instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. CONDITIONS Body Parts 27± r C 37± V C Isolated Crowded Isolated Crowded

Dorsol View Head

Ground colour 15L4 15L4 15L6 15L6 OLIVE GREEN OLIVE GREEN Metallic Green^ Metallic Green^

Streak 15E4 15E4 15E4 15E4 Beech Beech Beech Beech

Dots 48 C 7 48 C 7 48 C 7 48 C 7 GRAPHITE + GRAPHITE + GRAPHITE + GRAPHITE + Pronotum

Ground colour 15L5 15L5 I4K3 14K3 Roman Green Roman Green Serpentine Serpentine

Streak 15A4 15A4 15A4 Limestone Limestone Limestone

Dots 48 C 4 48 C 4 48 C 4 48 C 4 Cobblestone Cobblestone Cobblestone Cobblestone

Lateral View Head

Ground colour 12 E 2 13C2 12E2 12E2 Sallow Lint Sallow Sallow

Streak 1212 15E7 12 12 15E7 Hay ACORN Hay ACORN

Dots

Compound Eye 14B4 14B4 14B4 14B2 Antilope DUST + Antilope DUST + Antilope DUST + Sirocco Pronotum

Ground colour 13K3 12K3 12J2 12J2 Olivesheen Cloudy Amber Absinthe Y Absinthe Y

Streak

Dots

Table. 17. Phlaeoba infumatai Ilird Instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. CONDITIONS Body Parts 27± r C 37± r C Isolated Crowded Isolated Crowded

Dorsol View Head

Ground colour 10K3 10K3 11 12 11 12 LEMON Y LEMON Y Malmaison Malmaison

Streak 8E7 8E7 7C7 7C7 Lava- Lava- Bonito Fuscous Bonito Fuscous

Dots 8C8 8C8 8C9 8C9 Chaetura Black Chaetura Black London Smoke + London Smoke + Pronotum

Ground colour 11 Kl 11 K 1 11 K 1 13G9 Acacia Acacia Acacia Sayal Br

Streak 7A8 7A8 7A8 6A 10 ROSE GREY ROSE GREY ROSE GREY Army Br

Dots 48 C 5 48 C 5 7E11 7E11 Sooty Black Sooty Black Trotteur Tan Trotteur Tan

Lateral View Head

Ground colour 14C6 14B3 14J2 13E6 Caucasia + Twine Dune + Silver Fern Arizona

Streak 15 A2 8E7 15E4 15E4 Traprock + Lava Beech Beech

Dots 10A2 14A4 10A2 12B5 Buff + Omiond + Buff + FALLOW

Compound Eye 6A9 6A9 6A9 7H7 Manon Manon Manon Cedar + Pronotum

Ground colour 11 C3 11 C3 12K3 12K3 Sheepskin Moth + Sheepskin Moth + Cloudy Amber Cloudy Amber

Streak 10A2 10A2 10A2 12B5 Buff+ Bufr+ Buff + FALLOW

Dots

Table. 18. Phlaeoba infumata: IVth Instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. CONDITIONS Body Parts 27± 1" C 37± r C Isolated Crowded Isolated Crowded

Dorsol View Head

Ground colour 15J5 14H4 141 10 141 10 OLIVE DRAB Hemp Cinnamon Br Cinnamon Br

Streak 8E7 8C 10 8C 10 8C 10 Lava- Moose Moose Moose

Dots 48 A 4 48 A 4 48 A 4 48 A 4 Slag Slag Slag Slag Pronotum

Ground colour 15L5 14L6 14D7 14D7 Roman Green CITRINE Adobe Adobe

Streak 7A 10 7A 10 8Jg gJ8 NEW COCOA NEW COCOA Java Brown Java Brown

Dots 16H12 16H12 16H12 16H12 Bronzesheen Bronzesheen Bronzesheen Bronzesheen

Lateral View Head

Ground colour 13B4 13F6 13F6 13F6 Tea Time + Praine + Prairie + Praine +

Streak 15 A3 15C4 15C4 15A3 Beaverpelt Pelt + Pelt + Beaverpelt

Dots 9D2 I0C4 IOC 4 9D2 CREAM SUNSET SUNSET CREAM

Compound Eye 12J5 12J5 12 J 5 15E4 LIGHT STONE LIGHT STONE LIGHT STONE Beech Pronotum

Ground colour 13E5 13E5 I4A6 14A6 Mastick Mastick Buckskin Buckskin

Streak 9D2 I0C4 10C4 9D2 CREAM SUNSET SUNSET CREAM

Dots

Table. 19. Phlaeoba infumata: Vth Instar: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. CONDITIONS Body Parts 27± 1° C 37± 1° C Isolated Crowded Isolated Crowded

Dorsol View Head

Ground colour 16E12 16E12 16J6 16A7 Bronze Lustre^ Bronze Lustre^ Beetle Eagle Clove Brown

Streak 15 A5 15 A4 15A4 15A4 Log Cabin + Hair Brown Hair Brown Hair Brown

Dots 15H 11 15H 11 15H 11 15H11 Brown Sugar Brown Sugar Brown Sugar Brown Sugar Pronotum

Ground colour 15H 11 16C9 I6A 11 16A 11 Brown Sugar Bronze Brown Elk Lama - Elk Lama -

Streak

Dots 15 A12 I5A12 15C 11 I5C 11 BURNT UMBER'' BURNT UMBER'' COCOA BROWN + COCOA BROWN +

Lateral View Head

Ground colour 15 A6 15C 1 15A6 14K5 BEAVER Cub BEAVER OLD GOLD

Streak 8E10 8E10 8E10 8J8 SEAL SEAL SEAL Java Brown

Dots 11 B2 I5A2 8J8 14L8 PUTTY Traprock + Java Brown Syrup

Compound Eye 14A4 13D2 13D2 14A4 Ormond + Bronze Clair Bronze Clair Ormond + Pronotum

Ground colour 14H4 13E4 I5A2 15E4 Hemp Sudan Traprock Beach

Streak 11 B2 15 A2 15A7 15A2 PUTTY Traprock Soapstone Traprock +

Dots

Table. 20. Phlaeoba infumata: Adult Male: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. CONDI TIONS Body Parts 27± r C 37± r C Isolated Crowded Isolated Crowded Dorsol View Head

Ground colour I5J5 15J5 7E9 7A 11 OLIVE DRAB OLIVE DRAB Kaffa VANDYKE BROWN ''

Streak 7A 10 7A 11 6F9 6F9 NEW COCOA VANDYKE BROWN'' Cameo Br Cameo Br

Dots 8C12 8C 12 8C 10 8C 10 Bracken Bracken Moose Moose Pronotum

Ground colour 15C6 15A7 16H12 16A 12 GREY 31 Pony Brown Bronzesheen Biskra Date -

Streak 14 B 4 14B4 15L11 I6C5 Antilope DUST+ Antilope DUST+ BufTalo Rat

Dots I6C9 16C9 16C9 16C9 Bronze Brown Bronze Brown Bronze Brown Bronze Brown

Lateral View Head

Ground colour 14 K 5 14K5 14J9 14J9 OLD GOLD OLD GOLD MUMMY MUMMY

Streak 8E5 I5H7 15C5 8E8 AFRICAN + Olive Brown Rabbit + AFRICAN

Dots II A3 II A3 11 A4 11 A4 BISQUE BISQUE NUDE NUDE

Compound Eye I4A4 11 J2 15J5 15J5 Ormond + Silver Fern OLIVE DRAB OLIVE DRAB Pronotum

Ground colour 13E6 I3K7 13K7 I3K7 Anzona Isabella Isabella Isabella

Streak 11 A3 11 A3 11 A4 8A 10 BISQUE BISQUE NUDE SEPL\ +

Dots

Table. 21. Phlaeoba infumata: Adult Female: Colour Names and Codes as observed using Colour Dictionary by Maerz & Paul, London. Body Parts CONDITIONS 27± 1" C 37± r C Isolated Crowded Isolated Crowded Dorsol View Head

Ground colour 11D2 11J4 11J4 I1J4 Italian straw Mustard Mustard Mustard

Post ocular patch 19D2 19D2 19D2 19D2 Reed Gr Reed Gr Reed Gr Reed Gr

Dots

Pronotum

11 K7 11 K7 7J 10 7JI0 Ground colour BUFE BUFE Laurel oak Laurel oak

11 B3 11 B3 11 B3 9B2 Mid Dorsal Region CHAMPAGNE CHAMPAGNE CHAMPAGNE POLAR BEAR

Dots

Lateral View Head

Ground colour 1212 1212 1212 1212 Hay Hay Hay Hay

Streak 1212 1212 1212 12L2 Hay Hay Hay 0/ive Y Dots

14H4 14H4 19C2 19C2 Compound Eye Hemp Hemp Water Gr Water Gr Pronotum

11 I 1 11 I 1 Ground colour 11 1 1 11 1 1 Chartreuse Y Chartreuse Y Chartreuse Y Chartreuse Y

Streak

Dots

Table. 22. Oedaleus abruptus : 1st instar: Colour names and codes as observed using Color Dictionary by Maerz & Paul, London. Body Parts CONDITIONS 27± 1° C 37± r C Isolated Crowded Isolated Crowded Dorsol View Head

Ground colour 18D3 18D3 18D3 19K3 Glass Gr Glass Gr Glass Gr Chrysolite Gr

Post ocular patch I5H8 15H8 16A10 16AI0 Rubber + Rubber + Bison - Bison -

Dots

Pronotum

Ground colour 15E10 15E10 I6A11 I6A11 OLIVE WOOD OLIVE WOOD Elk Lama- Elk Lama-

Mid Dorsal Region 15L4 15L4 I5E4 15E4 OLIVE GREEEN OLIVE GREEEN Beech Beech

Dots

Lateral View Head

I9G5 I9G5 Ground colour 19C2 19G5 Turtle Gr Turtle Gr Water Gr Turtle Gr 15C1 15C1 Streak I5CI 15CI Cub Cub Cub Cub

Dots

19A2 19A2 Compound Eye 19A2 19A2 Oyster Gy Oyster Gy Oyster Gy Oyster Gy Pronotum

I5C2 15C2 llCl UCl Ground colour English Grey English Grey Amber white Amber white 20J3 20J3 20J3 20J3 Streak Shadow Green Shadow Green Shadow Green Shadow Green

Dots

Table. 23. Oedaleus abruptus : Ilnd linstar: Colour names and codes as observed using Color Dictionary by Maerz & Paul, London. Body Parts CONDITIONS 27± 1° C 37± 1° C Isolated Crowded Isolated Crowded Dorsol View Head

Ground colour 1815 18B5 18B5 18B5 SkyGr OCEAN GR OCEAN GR OCEAN GR

Post ocular patch I3L9 14G11 14G11 14G11 Chipmunk Tortoise Shell Tortoise Shell Tortoise Shell

Dots

Pronotum

Ground colour 14DI0 14J11 I4J11 14J11 Mustard Br + Cognae Cognae Cognae

Mid Dorsal Region 1716 1716 18F3 18F3 Viridine Gr Viridine Gr Glass Gr Glass Gr

Dots

Lateral View Head

22B6 Ground colour 20D2 22B6 22B6 PALMETTO Green Stone PALMETTO PALMETTO 15C1 Streak I5C1 15C1 15C1 Cub Cub Cub Cub

Dots

I9K3 Compound Eye 19G5 I9G5 I9K3 Chrysolite Gr Turtle Gr Turtle Gr Chrysolite Gr Pronotum

16A2 I5C2 I5C2 15C2 Ground colour SMOKE BROWN English Grey English Grey English Grey 19B4 I9B4 19B4 19B4 Streak Corydalis Gr Corydalis Gr Corydalis Gr Corydalis Gr

Dots

Table. 24. Oedaleus abruptus : IlIrd Instar: Colour names and codes as observed using Color Dictionary by Maerz & Paul, London. Body Parts CONDITIONS 27± r C 37± V C Isolated Crowded Isolated Crowded Dorsol View Head

Ground colour 19K6 I9K6 19K6 19K6 SEAGR SEAGR SEAGR SEAGR

Post ocular patch 2033 2033 2033 2033 Shadow Green Shadow Green Shadow Green Shadow Green

Dots

Pronotum

Ground colour 16A12 I6A12 16A6 16A6 Biskra Date Biskra Date TAUPE TAUPE

Mid Dorsal Region 20K5 20J6 20J6 20J6 Verdant Gr PEA GREEN PEA GREEN PEA GREEN

Dots Lateral View Head

Ground colour 21A5 2IJ6 2U6 21J6 Fern Gr Mignon Gr Mignon Gr Mignon Gr Streak 8E7 8E7 8E7 7C7 Lova- Lova- Lova- Bom to Dots

Compound Eye 14A4 I4A4 I4J2 I4J2 Ormond+ Omiond+ Silver Fern Silver Fern Pronotum

Ground colour 16C3 16C3 16C3 22G1 Rodent Rodent Rodent Vetiver Gr

Streak 22L2 22L2 22L4 22L4 Palm Palm Caller Gr Caller Gr Dots

Table. 25. Oedaleus abruptus : IVth linstar: Colour names and codes as observed using Color Dictionary by Maerz & Paul, London. Body Parts CONDITIONS 27± 1" C 37± 1" C Isolated Crowded Isolated Crowded Dorsol View Head

Ground colour 20J5 20J5 20C9 20C9 Absinthe Gr Absinthe Gr Peacock Gr Peacock Gr

Post ocular patch 21L3 21L3 21L5 21L5 Coyrge Gr Coyrge Gr GLASS GR GLASS GR

Dots

Pronotum

Ground colour 16A9 16A12 16A12 I6A12 Owl Biskra Date- Biskra Date- Biskra Date-

Mid Dorsal Region 20J3 20J3 I6J6 16J6 Shadow Green Shadow Green Beetle Beetle

Dots Lateral View Head

21F3 21J6 21J6 2IJ6 Ground colour Silver Gr Mignon Gr Mignon Gr Mignon Gr

I4B4 15A2 15A2 15A2 Streak Antelope Traprock+ Traprock+ Traprock+

Dots

14 H 4 I4H4 19C2 21 B 1 Compound Eye Hemp Hemp Water Gr Quaker Gy Pronotum

Ground colour 7H7 7H7 7H7 7H7 Cedra+ Cedra+ Bonito Bonito Streak 20A2 20A2 20A2 2215 Mineral grey Mineral grey Mineral grey SAGE GR Dots

Table. 26. Oedaleus abruptus : Vth linstar: Colour names and codes as observed using Color Dictionary by Maerz & Paul, London. Body Parts CONDI TIONS 27± V C 37± 1" C Isolated Crowded Isolated Crowded Dorsol View Head

Ground colour 2217 22GI 22E8 22L4 ArtGr Wetiver Gr Civette Gr Callar Gr

Post ocular patch 20F6 20F6 20F6 20F6 Grasshopper Grasshopper Grasshopper Grasshopper

Dots

Pronotum

Ground colour I5A5 16A12 16A12 16AI2 Log Cabrn+ Biskra Date Biskra Date Biskra Date

Mid Dorsal Region 20J5 20K6 20K6 20K6 Absinthe Gr Piquant Gr Piquant Gr Piquant Gr

Dots Lateral View Head

Ground colour 21L5 21L5 2IL5 21L5 GRASS GR GRASS GR GRASS GR GRASS GR

Streak 7C7 7C7 7C8 7C8 Bonito Bonito MAUVETAUPE MAUVE TAUPE Dots

Compound Eye I4H4 14 H 4 19C2 21 Bl Hemp Hemp Water Gr Quaker Gy Pronotum

Ground colour 7A8 7A8 7A2 7A2 ROSE GREY ROSE GREY Elephant Skin Elephant Skin

Streak 23J5 23J5 22C7 22C7 Elm Green Elm Green Cactus Cactus

Dots

Table. 27. Oedaleus abruptus : Adult Male: Colour names and codes as observed using Color Dictionary by Maerz & Paul, London. Body Parts CONDITIONS 27± V C 37± 1» C Isolated Crowded Isolated Crowded Dorsol View Head

Ground colour 20C9 20A9 22K7 20C9 Peacock Gr Kllemey Gr Spinach Gr Peacock Gr

Post ocular patch 19K2 19K2 20CI 20D2 Citron Gr Citron Gr Eucalyptus Gr Green Stone

Dots

Pronotum

Ground colour 16C9 16A12 16A12 16A12 Bronze Brown Biskra Date- Biskra Date- Biskra Date-

Mid Dorsal Region 21J6 21F3 2117 2U6 Mignon Gr Sliver Gr Fern Mignon Gr

Dots Lateral View Head

22K7 22K7 Ground colour 23L4 23U Spinach Gr Quaker Green Quaker Green Spinach Gr 15C1 15C1 Streak 15A3 15C1 Cub Nutria Cub Cub

Dots

21 B 1 14 H 4 19C2 Compound Eye 14H4 Quaker Gy Hemp Water Gr Hemp Pronotum

13C3 13A3 13C3 Ground colour 13 A3 Lido Cuban Sand- Cuban Sand- Lido 22B5 23J5 22B5 Streak 23L5 Mistletoe Cedar Green Elm Green Mistletoe

Dots

Table. 28. Oedaleus abruptus : Adult Female: Colour names and codes as observed using Color Dictionary by Maerz & Paul, London. II — — ti-i

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