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Durham E-Theses Durham E-Theses Biological studies on the meadow pipit (anthus pratensis) and moorland tipulidae; members of a food chain Coulson, John Cameron How to cite: Coulson, John Cameron (1956) Biological studies on the meadow pipit (anthus pratensis) and moorland tipulidae; members of a food chain, Durham theses, Durham University. Available at Durham E-Theses Online: http://etheses.dur.ac.uk/8840/ Use policy The full-text may be used and/or reproduced, and given to third parties in any format or medium, without prior permission or charge, for personal research or study, educational, or not-for-prot purposes provided that: • a full bibliographic reference is made to the original source • a link is made to the metadata record in Durham E-Theses • the full-text is not changed in any way The full-text must not be sold in any format or medium without the formal permission of the copyright holders. Please consult the full Durham E-Theses policy for further details. Academic Support Oce, Durham University, University Oce, Old Elvet, Durham DH1 3HP e-mail: [email protected] Tel: +44 0191 334 6107 http://etheses.dur.ac.uk 2 Biological studies on the Meadow Pipit (^thus mS^S^^^ and moorland fipuiidae (Dipt.); members of a rooa - : J^y John Cameron Coulson. Observations were made on 65 species of .!gipulidae found on the Moor itouse Nature Conservancy Reserve, Westmorland, between 1953 and 1956. The larval habitats were recorded and a stu<^ was made of the abundance and seasonal distribution of adults. Particular attention was paid to the biology of two common species of Tipulidae. namely Tipula subnodicornis Zett. which lives on peat moors and Jipula paludosa Meigan vidiich lived in mineral soils, while a comparison was made between the latter species and the closely related species, Tipula oleraeea, Meigen. A study of the egg production and mortality of the Meadow Pipit was carried out by means of ringing recoveries and nest record cards, particular attention being paid to the change in clutch size, nest mortality and time of breeding with an increase in altitude. Th9 Meadow Pipit was studied at Moor House and particular attention was paid to its food supply. A study was made of feeding areas and the food which was brought to the young. It was concluded that the predation by the Meadow Pipit had little influence on the population of 2i, subnodicornis althoug^i this species and T, paludosa fomned the major part of the food brought to the nestlings. BIOLOGICAL STUDIES ON THE MEADOW PIPIT (AWTHUS PRATENSIS) AND MOORLAHD TIPULIDAE; MEMBERS OF A FOOD CHAIN. By JOHN CAMERON COULSON ~ being a thesis presented in candidature for the Degree of Doctor of Philosophy in the University of Durham, 1956. SCIENCf SECTIOtt The author wishes to express his thanks to Professor J.B. Cragg for his. guidance and encouragement throughout the investigation, also to Dr. E.S. Page for advice, on the application of suitable statistical techniques* The author is indebted to the Durham Colleges and the Nature Conservancy for research studentships which have made this study possible. Contents. Page. I. Introduction. 1« II. Literature and taxonomy. 2. III. The study area. 4. rv. The study of Tipulidae. 7. 1. Methods. 7. Sampling for eggs and larvae. 7. Emergence traps. 7. Sticky traps.. 10. Rearing of adults. 10.. 2. List of species of lipulidae recorded from the Moor Houae Nature Reserve. 11. 3. Geographical distribution of British Tipulidae. 15. 4. Notes on the Tipulidae of Moor House Reserve. 17. 5.. Larval habitats on the Moor House Reserve. 37. 6. Duration of adults and pupation. 39. 7.. The seasonal distribution of Tipulidae. 43. 8, Egg filaments in Tipulinae. 46. 9. Development of eggs. 48. 10. Key to the larvae of some moorland Tipulidae. 50. T. An ecological and biological study of Tipula subnodicornis. 56. 1. Adults. 56. Male activity. 56. Female activity. 57. Emergence rhythm of the females. 58. Egg laying. 60. Rate of egg laying. 66. 2. Release and recapture marking experiment on adults. 69. Population estimates. 73. Sex ratio. 74. Mortality. 75. Emergence. 77. 3. The egg stage. 80. Development period. 80. Mortality under field conditions. 81. The vertical distribution of eggs in the peat. 82. 4. The larval stage. 84. Instars. 84. Population study. 86, Overwinter mortality. 87. Duration of instars. 88. Growth. 89. Sex ratio. 90. Larval populations on different vegetation types. 91. The suitability of habitats. 95. 5. The distribution of adults, eggs and larvae on a Juncus squarrosus moor. 96. 6* Causes of mortality . 100. 7. The effects of drought and high rainfall on the numbers of T. subnodicornis. 102. 8» The effects of dry weather on the larval populations in different habitats. 106. 9. The food of the larvae. 107. 'VI. The study of Tipula paludosa. 109. 1. The adult. 109. Emergence period. 109. Emergence rhythm. 109. Activity. 112. Rate of egg laying. 116. Development of new eggs. 119. Oviposition. 119. 2. The egg. 121. 3. The larva. 122. Duration of instars. 122. Population study. 122. Effect of dry summer of 1955 on the population density. 124. YII The study of Tipula oleracea. 125. 1. The adult. 125. 2. The egg. 126. 3. The larva. 127. VTH.Study of the Meadow Pipit by means of nest record cards and ringing recoveries. 129. 1. Breeding. 129. Determination of start of laying.129. Determination of altitude of nests.130. Time, of breeding. 131. 2. Clutch size. 133. Effect of latitude on clutch size. 134. Relationship of clutch size to altitude. 136. Size of first and second clutches. 137. 3. Incubation period. 138. 4. Fledging period. 138. 5. Mortality. 138. Nest mortality. 138. Relation of altitude to total neat mortality. 143. Eggs which failed to hatch. 144. Loss of eggs from nests. 145. Loss of young from nest. 145. Determination of post-fledging mortality from ringing-recoveries. 145. 6. Discussion. 150. IX. Study of Meadow Pipit at Moor House. 156. 1. Methods. 156. Nest visit recorders. 156. Finding nests. 158. Determination of the number of food items brought to the young. 158. 2.. Rate of feeding visits by adult Meadow Pipits to young in the nest. 160. Diurnal rhythm. 162. Feeding frequency at nests with one, four and five young. 163. Effect of cold weather. 168. 3. Second broods. 170. 4. Time of breeding at Moor House. 171. 5. The clutch size at Moor House. 171. 6. Total nest mortality at Moor House. 172. 7. Feeding of Meadow Pipit and related species. 173. 8. Food brought to young at Moor House. 174. 9. Association of the Meadow Pipit with streams. 177. 10. Density of the Meadow Pipit at Moor House. .179. X. Predation of the Meadow Pipit on adults of Tipula subnodicornis. 181. XI. The relationship of Meadow Pipits and Tipulidae. 184. XII. Discussion. 186. XIII. Summary. 19 5. XIV. References. 203. X\r. Appendix. 215. BIOLOGICAL STUDIES ON THE MEADOW PIPIT (ANTHUS PRATENSIS L.) AND MOORLAND TIPULIDAE; MEIvBERS OF A FOOD CHAIN. I Introduction. Amongst the insects and birds inhabitiig British moorlands, Tipulidae and Meadow Pipits are often the commonest representatives and in spite of this abundance, very little detailed biological information is available. The investigations were carried out on the Moor House Nature Reserve, Westmorland between 1953 and 1956, an area where Campbell (1955) had shown the Meadow Pipit to be the commonest bird and where preliminary studies had shown that tipulids could be numerous. It was hoped that as a result of the present studies it would prove possible to assess the role of tipulids and Meadow Pipits in the general ecology of this fieserve and particularly the relative importance of Tipulidae as food of the Meadow Pipit. eClENCf SEOTIOI' II Literature and taxonomy. The classical study on the family Tipulidae is the work of Alexander (1920) and while this was confined to the taxonomy of the adults and larvae in the State of New York, it summariized much of the literature on . the biology of the family. The taxonomy of adult European Tipulidae had been well studied prior to this study and little difficuli^r arose in using the keys of Edwards (1938) and Coe (1951) while the incomplete keys of Mannheims (1940-53) ws/* of assistance since they contained drawings of the male genetalia for the majority of species found in Europe. The taxonomy of larvae of European Tipulidae has not been studied extensively and little has been published since the works of Beling (1879, 1886) (this work is not wholly satisfactory through the lack of illustrations). Oldham (1928) has described the larvae of Tipula paludosa and T. lateralis Meigen, while the larva of Phalacrocera replicata,.has been iUustcated by Miall& Shelford (1897). The ecology of the- Tipulidae has not been studied extensively. Bodenheimer (1923, 1924) has made a stuc^ of Tipula oleracea but T. paludosa was confused with this species. Rennie (1916, 1917) studied the life -2- history of T. paludosa and Barnes (1937) studied this species under laboratory conditions. White (1951) has studied T. lateralis in the laboratory and recorded that this species has four larval instars. Cuthbertson (1926) described the larval habitats of a number of tipulids in Scotland but did not describe any of the larvae. Barnes (1925) selected 16 habitats in Carnarvonshire and showed that the Tipulidae were most common in damp or wet habitats and Rogers (1933, 1942) came to similar conclusions from studies in Florida and Michigan, U.S.A. The seasonal change in species and the nocturnal activity of certain Tipulinae has been studied at Rothamsted by means of a light trap (Pinchin & Anderson 1936; Robertson 1939) and they concluded that the nocturnal activity of Tipula paludosa was controlled by the daily minimum temperature.
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