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Adelges Insects of Silver Firs

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Adelges Insects of Silver Firs FORESTRY COMMISS 10 N BULLETIN No. 26 ADELGES INSECTS OF SILVER FIRS By I. W. VARTY, B.Sc., Ph.D. DEPARTMENT OF FORESTRY ABERDEEN UNIVERSITY EDINBURGH: HER MAJESTY’S STATIONERY OFFICE PRICE 8s. 6d. NET Forestry Commission ARCHIVE FORESTRY COMMISSION BULLETIN No. 26 ADELGES INSECTS OF SILVER FIRS By ISAAC WILLIAM VARTY, B.Sc., Ph.D. DEPARTMENT OF FORESTRY ABERDEEN UNIVERSITY EDINBURGH: HER MAJESTY’S STATIONERY OFFICE 1956 FOREWORD Insects of the genus Adelges have long been recognised as economically important pests of the valuable timber-producing silver fir trees that comprise the genus Abies. In fact the planting of the common European silver fir, Abies alba, has had to be greatly restricted in Britain because of the depredations of these insects. This Bulletin gives the results of Dr. Varty’s intensive studies of the Adelges insects in relation to forestry, which were carried out in Scotland under the auspices of the Department of Forestry of Aberdeen University. It replaces the former Bulletin No. 7, entitled The Silver Fir Chermes, which has long been out of print. FORESTRY COMMISSION, 25 Savile Row, London, W.l March, 1956 CONTENTS PAGE PAGE INTRODUCTION AND ACKNOWLEDG­ 4. Progrediens aptera and Sexupara 25 MENTS 1 5. Sexualis .... .... 29 5. Life Cycle of Adelges piceae Ratz. 30 CHAPTER 1. TAXONOMY 1. Sistens .... 30 2. Progrediens................................................. 33 1. Discussion of Family Rank 1 6. Population levels of Adelges niisslini and A. 2. Discussion of Generic Taxonomy 2 piceae and natural control 34 1. Chronology of literature... 2 1. Population levels.......................... 34 2. Morphology at the generic level.... 3 2. The effects of climate on populations 35 3. Discussion of Specific Taxonomy 4 3. Biotic factors 37 1. Chronology of the literature .... 4 2. Morphology at the specific level. .. 5 4. Description of Adelges niisslini C.B. 6 CHAPTER 3. THE REACTION OF THE TREE 1. Techniques... 6 TO ATTACK BY ADELGES NUSSLINI C.B. 2. Fundatrix 6 AND A. PICEAE RATZ. 3. Gallicola 7 1. The general appearance of the attacked trees 39 4. Sistens .... 7 1. Attack by Adelges niisslini 39 5. Progrediens aptera 7 2. Attack by Adelges piceae.... 39 6. Sexupara 8 7. Sexualis .... ... 8 2. The Anatomical Effects of Attack ... 39 1. Method of S tu d y ........................................ 39 5. Description of Adelges piceae Ratz. 9 2. Anatomical effects of the feeding by 1. Sistens .............. 9 Adelges niisslini on Silver fir .... .... 40 2. Progrediens aptera 9 3. Anatomical effects ofAdelges niisslini on 3. Progrediens alata ....................................... 10 Oriental spruce ........................................ 40 6. The Application of Paper Chromatography 4. Anatomical effects ofAdelges piceae on to Taxonomy 10 Silver f ir ........................................ 41 3. Chemical Aspects of Leaf Attack by Pro- CHAPTER 2. LIFE HISTORIES OF ADELGES gredientes of Adelgcs niisslini 43 NUSSLINI C.B. AND ADELGES PICEAE RATZ. 1. Aims and Methods 43 2. Chlorosis of leaves ............... 43 1. Salient Features of the Life Cycle within the 3. The sugar content of attacked leaves .... 43 Subfamily Adelginae with Special Refer­ 4. The effect of attack on the nitrogen ence to Adelges niisslini and A.piceae 12 content of leaves ........................... 45 2. Review of Previous Research........................... 14 5. The effect of attack on the mineral 1. Literature on the biology of subfamily content of the leaves ............... 46 Adelginae ........................................ 14 4. Susceptibility and Resistance in Silver Firs to 2. Literature on the biology ofAdelges Attack by Adelges niisslini and A. piceae 47 niisslini and A. piceae.... 15 1. Definition of resistance in host plants ... 47 3. Theories of Evolution ........................... 16 2. Evidence of resistance to Adelgids in the 3. Research Methods and Meteorological literature .................................................... 47 Records .... 17 3. Further observations on resistance and 1. Research methods. .. 17 susceptibility .... 48 2. Field Study .............. 18 4. Experimental infections ........................... 49 3. Meteorological records .... 18 5. The nature of the resistance mechanism.... 51 4. Life Cycle of Adelges niisslini C.B 18 5. pH Studies in Relation to Resistance .... 53 1. Fundatrix 18 1. Hypothesis of a relationship between host 2. Gallicola 19 pH and resistance 53 3. Sistens 22 2. Sampling 53 iii CONTENTS PAGE PAGE 3. Measurements of pH on Silver fir species 2. Applied control measures againstAdelges 63 resistant or susceptible to Adelges 3. Proposals for experimental plots ofAbies niisslini ........................... ............... 54 species ........................... 64 4. Further investigations on host pH and susceptibility to Adelges 56 SUMMARY 65 CHAPTER 4. FOREST RELATIONS APPENDICES Forest Status and Distribution of the Adelges Appendix 1. Meteorological data 66 of Silver fir ....................................... 56 Appendix 2. Estimation of chlorophyll con­ 1. The spread and present distribution of tent in leaves of Abies alba Adelges niisslini .... ........................... 56 attacked by progredients of 2. Descriptions of outbreaks of Adelges Adelges niisslini 68 niisslini in Scotland ............... 57 Appendix 3. Technique of sugar estimation.... 68 3. The distribution and forest status of Adelges piceae ............... 61 Appendix 4. Techniques of nitrogen estimat­ 4. The effect of site on the severity of attack 62 ion for leaves ofAbies alba .... 69 Silvicultural A spects ....................................... 62 Appendix 5. Measurements of pH in Abies spp. 70 1. The regeneration of Abies species in Adelges-infested regions ............... 62 REFERENCES 72 INTRODUCTION AND ACKNOWLEDGMENTS Adelges niisslini and A. piceae, which together are Throughout this work, the writer has used the variously known as the Adelges of Silver fir, and as Handbook o f Coniferae by Dallimore and Jackson the woolly aphids or Chermes of Silver fir, are (55) as the standard reference for tree names, and minute polymorphic insects belonging to the family the Check List of British Insects by Kloet and Hinck (93) for insect names. Phylloxeridae (Aphidoidea, Hemiptera). These two This investigation has been conducted under the species occur both in Europe and North America supervision of Professor H. M. Steven, Forestry where Adelges has assumed considerable silvicultural Department, University of Aberdeen, whose help­ importance because of the susceptibility of certain fulness and stimulating suggestions have been very species of Abies. In Britain, where both insects and much appreciated. The writer is indebted to Dr. E. host trees are introduced species, the persistently V. Laing and other members of the staff of the severe attack by Adelges niisslini on young trees of Forestry Department for their co-operation. Abies alba has resulted in the virtual suspension of Thanks are also due to Dr. P. C. de Kock, Macaulay the planting of this useful species during the last Institute for Soil Research, Aberdeen, without 30 years. whose help and encouragement the chemical In this investigation over part of the period 1950- aspects of this research could not have been 1954, the writer’s purpose has been to study the attempted, and to Dr. R. L. Mitchell, also of the biology of the Silver firAdelges as found in Scotland, Macaulay Institute, for undertaking certain analyses to consider certain aspects of susceptibility and re­ acknowledged in the text. Finally the kindness of sistance to attack shown by the various host species, Mr. C. Y. Ross, Head Forester of Dunecht Estate, and to assess the current silvicultural significance of in providing facilities to use the Forest Garden as these insects. a research plot, is gratefully acknowledged. Chapter 1 TAXONOMY 1. DISCUSSION OF FAMILY RANK subfamily, viz. the Phylloxerinae and the Cherme- sinae. Six years later after further study of the The two species Adelges piceae Ratz. and A. group, he (21) raised each of the subfamilies to niisslini C.B., under discussion in this work are separate family status. Within his new family members of the subfamily Adelginae in the family Chermesidae he differentiated two tribes, the Phylloxeridae of the superfamily Aphidoidea, as Pineini and the Chermesini. Yet in 1915 Cholod- recognised in Kloet and Hinck’s (93) Check List of kovsky (42) whose authority on the systematics of British Insects. In the course of arrival at this the group was fully equal to Borner’s, considered the taxonomic status, Adelgids as a whole have suffered Adelgids to be merely a genus within the subfamily the usual vicissitudes of treatment accorded to any Phylloxerinae in the family Aphidae. group with complex morphological and biological Borner’s classification came to be tacitly accepted characters. In the early years of scientific investiga­ until in 1928 Annand (2) published his important tions following the time of Linnaeus, the group though incomplete Contribution toward a monograph held only generic rank, Chermes, in an indefinite o f the Adelginae, (Phylloxeridae) of North America. position within the family Aphididae; but from 1857 In this work he adopted subfamily rank for the it has been known as a small family, the Chermidae group, which he now named the Adelginae in view Koch and from 1862 as the Chermesidae Passerini. of the fact that the old name Chermesinae had been In time its systematic proximity to thePhylloxera properly allotted
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