"Bionomics of the Alder Sawfly Fenusa Dohrnii (Tischbein)"
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"BIONOMICS OF THE ALDER SAWFLY FENUSA DOHRNII (TISCHBEIN)" by Kulapu Arachchige Don Wilmot Senaratne, B.Sc.(Ceyl). Thesis submitted to the University of London for the degree of Ph.D. East Mailing Research Station Maidstone Kent October 1978. 1 ABSTRACT Field and laboratory studies during 1975-77 have shown that Fenusa dohrnii is bivoltine in south - east England with adults emerging during May - June and July - August. Average fecundity was estimated to be about 62 eggs per female, the number being correlated with both weight and tibial length of adult. At 15°C and 21°C respectively, the embryonic period was 13 and 7.5 days whilst larval development took 26.6 and 16.9 days. Selection of a cocooning site in the soil by the sixth instar larva was random; no active burrowing was observed, but the larvae gained entry via cracks and patches of loose soil. Prepupal diapause was terminated to a varying degree when cocoons were chilled for about 62 days at -1.1°C, 3.3°C, 10°C and field temperatures. The highest percentage emergence of adults was from cocoons exposed to field temperatures and progressively fewer emerged as the chilling temperature was lowered from 10°C to -1.1°C. For all treatments mean time for adults to emerge after transfer to 21°C decreased with increasing duration of chilling. In the field, distribution of eggs of the second generation was correlated more closely with the negative binomial than Poisson , Neyman type A or logarithmic distributions. Studies of the population development indicated that the generation mortality varied from 95 - 98 percent of which mortality of larvae within the mines and mortality of larva'VI due to parasitism by Lathrolestes pictilis (Ichneumonidae) and Ichneutes laevis (13raconidae) contributed most towards 2 the generation K value. Two eulophid parasites Cirrospilus vittatus (Wik.) and Chrysocharis nitetis (Wik.) were also found parasitizing the young larvae. There was also circumstancial evidence that predators were responsible for some mortality of larvae but this was not confirmed either by observations or electrophoretic techniques. 3 ACKNOWLEDGEMENTS The work for this thesis was carried out at East Mailing Research Station. For the first two years of this study I was in receipt of a Colombo-plan scholar- ship. I am grateful to the Director of East Mailing Research Station for providing me with the facilities to carry out this project, the Director of the Tea Research Institute, Sri Lanka, for granting me leave of absence and the British Council for making all necessary arrangements. I am also indebted to my supervisor Dr. G.H.L. Dicker, and my Director of studies Dr. G. Murdie for their guidance, constructive criticism and helpful discussions throughout this project. Thanks are due to Mr. G.P. Barlow for statistical advice and analysis of some of the data, Dr. R.A. Murray for electrophoresis of insect samples, Dr. K.A.D. MacKenzie for the preparation of leaf sections, Dr. M.G. Fitton, J.S. Noyes and T. Huddleston of the British Museum (Natural History) for identification of parasites, Mr. D.G. Richardson for printing of the thesis, and members of the photographic studio for preparing some of the photographs. Grateful acknowledgement is also due to Annals of Applied Biology for permission to reproduce Figure 24. I also thank Dr. B.H. Howard, Dr. J.J.N. Flegg and all other members of the Department of Zoology for their invaluable assistance. I am also grateful to Mrs. G. Squires for typing this thesis and last but not least I wish to record my thanks to my wife Shiranganie for her financial assistance which made it possible to continue this work beyond two years. 4 LIST OF FIGURES Figure Page 1 Cages used to collect laboratory reared adults. 15 2 Part of the experimental area with emergence cages and sticky boards in position. 17 3 a) A recently laid egg and one containing a well developed embryo curled within the chorion. b) Blisters on the dorsal surface of a leaf of Alnus Zlutinosa indicating the position of developing egg. c) Transverse section of a leaf showing an egg between the palisade and spongy mesophyll. 21 4 Transverse sections of leaves of Alnus glutinosa a) from plants grown in the field; b) from plants raised in the glasshouse. 25 5 Ovarioles of Fenusa dohrnii showing developing and mature oocytes. 28 6 Relationship between the number of ovarioles (OV) and the weight of the adult (W). 29 7 Relationship between, a) number of eggs (E) and hind tibial length (L); b) number of eggs (E) and weight of adult (W); c) weight of adult (W) and hind tibial length (L); for adults shortly after (0) and approximately ten days after (.) emergence. 32 8 Development of eggs at constant temperatures of 21°C and 15°C. 34 9 Leaves showing (a) single mines and (b) coalesced mines. 36 5 Figure Page 10 Frequency of head capsule widths of larval instars and the range for each instar. 38 11 Relationship between head capsule width (Y) and larval instars (X) plotted on a log scale. 40 12 Period of development of eggs and larvae when reared at a constant temperature of 21°C or 15°C. 43 13 Relationship between area of mine and the total leaf area. 46 14 Paths followed by mature sixth instar larvae on bare stabilized soil before either cocooning or leaving observational area of 100 x 100 cm on a) October 21st, b) October 22nd. 48 15 Paths followed by mature sixth instar larvae on bare stabilized soil before cocooning in two experiments on 27th October. 49 16 a) Single and b) aggregated cocoons of Fenusa dohrnii showing the uneven surfaces. 53 17 Numbers of F. dohrnii in emergence traps at three-day intervals, 1976. 57 18 Relationship between weight of adult (WA) and the weight of mature sixth instar larva (WL). 59 19 Relationship between number of eggs (E) and a) weight of adult (W); b) weight of mature sixth instar larva (W1). 60 20 Percent emergence at 21°C in relation to period of chilling. (a) F. dohrnii (b) Parasites. 63 6 Figure Page 21 Mean time to emergence of adults after transfer to 21°C in relation to period of chilling. (a) F. dohrnii (b) Parasites. 65 22 Variation of S.E. of mean time to emergence of adults of F. dohrnii after transfer to 21°C in relation to period of chilling. 66 23 Numbers of L. pictilis and I. laevis in emergence traps, 1977. 78 24 Electrophoretic slab gels showing A) anthocorid starved for 36 h (1); anthocorid starved and then fed on larva of F. dohrnii (2); larva of F. dohrnii (3); B) anthocorid starved for 36 h (1); anthocorid starved and then fed on alder aphid (2); alder aphid (3). 82 25 Relationship between observed and expected frequencies for leaves containing no eggs (zero variate). 88 26 Relation of k to mean for 36 samples from a population of Fenusa dohrnii eggs. 89 27 Relationship between Y1 and X1 where X1 = mean2 - varianc;a number of sampling units and Y1 = variance - mean. 90 28 Relationship between variance mean (M) (V) and mean (M) for 36 samples from a population of F. dohrnii eggs. 96 29 Relationship between mean crowding(Mx) mean (M) and mean (M) for 36 samples from a population of F. dohrnii eggs. 99 7 Figure Page 30 Relationship between variance (V) and mean (M) of the number of eggs per leaf, plotted on log scale. 102 31 Relationship between Lloyd's measure of mean crowding (Mx) and mean density (M) for eggs of Fenusa dohrnii. 103 32 Early growth of Alnus glutinosa showing individual leaf clusters. 106 33 Numbers of adults, eggs, larva I and VI per 576 clusters. (a) and (b) generation I, 1976; (c) and (d) generation II, 1976. 110 34 Numbers of eggs and each larval instar per 576 clusters. Generation II, 1977. 111 35 Totals of eggs and each larval instar per 576 clusters on each sampling occasion in generation II, 1977. 112 36 Changes in mortality of F. dohrnii expressed as k values. 118 37 The k values of various mortalities of F. dohrnii plotted against the density on which they acted. (a) ki on eggs ; (b) k3 on larva I ; (c) k2 on larva I. 120 8 LIST OF TABLES Table Page 1 Measurement of 20 eggs shortly after deposition. 20 2 Force (gm/mm2) required to penetrate leaves on shoots of Alnus glutinosa grown in the field or in the glasshouse. ~3 3 Number of eggs laid in leaves of Alnus glutinosa by adults in cages. 26 4 Relationship between number of mature oocytes, adult weight, and tibial length in females shortly after, and 10 days after, emergence. 31 5 Width of head capsule, Dyar's constant, and the percentage increase. 39 6 Variation in the area mined by single larvae of F. dohrnii in the field. 45 7 Distance travelled on the soil surface and speed of 18 larvae. 50 8 The speed (cm/mta) of larvae in relation to distance travelled on the soil surface. 51 9 The number of cocoons at various depths beneath alder trees, 54 10 Percent cocoons with prepupae and pupae, 1975-76 56 11 Percentage of cocoons occupied by F. dohrnii and by parasites. 62 12 Numbers of Ichneutes laevis and Lathrolestes pictilis emerging in relation to period of chilling. 68 13 Variation of mean time to emergence of parasites from samples chilled for 182 days. 69 14 Mean temperatures (°C) at 5 cm below soil surface, East Mailing 1975-76.