Food Preferences of the European Hare (Lepus Europaeus
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Food Preferences of the European Hare (Lepus europaeus Pallas) on a fescue grassland. A thesis submitted in partial fulfilment of the requirements for the degree of Master of Science in Zoology in the university of Canterbury by Garry Blay. University of Canterbury 1989 TABLE OF CONTENTS Table of contents ..................................... i List of figures ....................................... ii List of tables ....... '..•.............................. iv Abstract 0 •••• II 0 a CI • It 0 It 0 Q CI 0 0 " •• 0 0 • D • 0 • II DOD. It ....... 0 0 0 • '" 0 • V Chapters. 1.0 Introduction 1 . 1 Taxonomy ....................................... 1 1.2 Introduction and Distribution of hares in N.Z .. 4 1.3 Previous research .............................. 5 1.4 Aims of this study ............................. 7 2.0 Study area 2 . 1 Introduction ................................... 8 2.2 Locality ....................................... 8 2. 3 Physiography .........................•........ 10 2.4 Climate ....................................... 10 2.5 Vegetation .................................... 12 3.0 Methods 3 .1 Introduction ..............................•... 14 3 .2 Diet st'udy .................................... 18 3.2.1 Collection of gut samples ............... 18 3.2.2 Preparation of gut samples .............. 19 3.2.3 Analysis of gut samples ....... , ......... 19 3.2.4 Preparation of reference slides ......... 20 3.3 Calorimetry ............................. , ..... 22 3.4 Neutral detergent fibre analysis .............. 24 3.5 Secondary metabolite analysis ................. 27 3.5.1 Extraction .............................. 27 3.5.2 Analysis ................................ 27 - Folin-Denis reaction ................. 27 - Vanillin reaction .................... 28 - Leucoanthocyanidin reaction .......... 29 3.6 Vegetation sampling ........................... 30 3.6.1 Availability estimate ................... 30 3.6.2 Exclosure plots ......................... 31 4.0 Results 4.1 Diet .......................................... 33 4.2 Vegetation sampling ........................... 50 4.2.1 Availability ............................ 50 4.2.2 Exclosure plots ......................... 56 4.3 Calorimetry ................................... 58 4.4 Secondary metabolite analysis ................. 67 4.5 Neutral detergent fibre analysis .............. 71 5.0 Discussion ....................................... 76 )V' Ack... owledgements ...................................... 94 References ........................................... 95 Appendices .............•.•...•...................... 103 List of Figures Figure 2.1: Location of study site .••••..•........•....• 9 Figure 3.1: Flow chart of the Van Soest method of fibre or digestibility determination .........•••. 25 Figure 4.1: Monthly diet of the hare population in the study area •• " .•..........•..•..••........••.. 40 Figure 4.2: Seasonal diet of the hare population in the study area ..................................• 41 Figure 4.3: Monthly diet of juvenile female hares ...... 42 Figure 4.4: Monthly diet of juvenile male hares .•.•.... 43 Figure 4.5: Monthly diet of adult female hares ........• 44 Figure 4.6: Monthly diet of adult male hares ........... 45 Figure 4.7: Seasonal diet of juvenile female hares ..... 46 Figure 4.8: Seasonal diet of juvenile male hares ....... 47 Figure 4.9: Seasonal diet of adult female hares ........ 48 Figure 4.10: Seasonal diet of adult male hares ......•.. 49 Figure 4.11: Monthly availability of plants in the study area ....................................... 54 Figure 4.12: Seasonal availability of plants in the study area ....................................... 55 t'll., Figure 4.13: Monthly calorific values of major die~y items ...... "" . ., .... "" .. e ... It ....... "' .... " .......... iii" .. • 60 Figure 4.14a: Calorific value x amount eaten of R. lanuginosum D It '" Iil ... It eo. I) • I) • G co I) .... (I • *' ••• to ........ II D ..... • 61 Figure 4.14b: Calorific value x amount eaten of H. pilosella. '" .... e It " It •• iii to ... I!I .. " • co .. \l •• 0: iii " a 0 1\1 " •• II' • " '" II 111 .. 62 Figure 4.14c: Calorific value x amount eaten of the grasses . ... j) .. Ii III !It ,. .... 0 " ••• "" u • " ., .. 01> • G •• ., ..... e " ., ......... 63 Figure 4.14d: Calorific value x amount eaten of the tussocks. u .. " ~ u .. 110 D iii ... bO 11 ... 0 ••••• "" It .... ., 1:1 It eo. II "" It 0 11 ••• 64 iii Figure 4.14e: Total calorific value of the major die~y species in all age and sex classes of har~"s. Sum of figures 4.14a-4.14d ....•........... 65 F~gure 4.15: NDF content of major die~y items ...••..•. 74 tv iv List of Tables .'I'ab~4:~: Percent frequency of occurrence for species listed under 'others' .....••••.••••..••.. 36 Table 4.2: Individual and averaged results from exclosure plots ......•.....•.•...........•.....•. 57 Table 4.3: Maximum, minimum and av~rage calorific value in kcal/g of major dieFry items in gut ;.. samp 1 es. II •• I) ft 0 • f> .. It \I ...... " '" ..... " • D ¢I iii> .... II • It ., " " " '" '" " .. lit 59 Table 4.4: Secondary metabolite content of major diee-ry items ..•.....••...•....................... 69 I' 0- ~T=a=b=l~e~4~.5~: Ranking of major dietry items in relation to content of secondary mttabolites and ove- r a 11 rank. '" .. D .. III ............ It 0. II ... D .. ., It 0 ........ It D ••• '" a III .. It • 69 Table 4.6: Ranking of major diet~y items when mono meric and condensed tanni~s are combined and given equal weighting with phenols .•..•••..•.•.•. 70 Table 4.7: Seasonal and averageJ?ercent NDF values and ranking for major dieEry items for all seasons .................. t . .......... , ........... 72 Table 4.~: Calorific value x amount eaten (as a percentage of the diet) x inverse of the NDF va 1 ue .. .... II It .. !O '" ... I) •• I:< • II .... a iii Q .... II • " II III ...... It " III • til ... 01 .... OJ 72 Table 4.9: Calorific value x inverse of the NDF va 1 ue .. I) " ... If " g e (0 ...... '" • " " III til ....... 0 .... D III G e 0 III lit III • iii It • " " " D • D 73 v ABSTRACT Diet of the European hare (Lepus europaeus Pallas) living on a fescue (Festuca novae-zelandiae) grassland in central high country Canterbury was examined using micro scopical gut sample analys • Availability of food plants was estimated by a monthly vegetation survey. Major dietary items were analysed for calorific value, neutral detergent fibre content (digestibility) and secondary metabolites (phenols, condensed tannins and monomeric tannins). Hares in this area relied heavily on Hieracium pilosella as the major constituent of their diet. Grasses and tussocks made up the majority of the remaining food. The diet of females, males, juveniles and adults were all significantly different. Diet also varied throughout the year, although it largely reflected the availability of the various food items, except Rhacomitrium lanuginosum which was avoided. Food plant selection did not correlate with calorific value or secondary metabolite content. However, there was some correlation between the neutral detergent fibre content or digestibility of the plants measured and their level of con sumption by the hares. It appears from the results of this study that the diet of hares in the study area is influenced predominantly by the availability of the various potential food plants but is also ihfluenced by the digestibility of those plants available. 1 Chapter 1 INTRODUCTION 1.1 Taxonomy The order Lagomorpha contains rabbits, hares and pikas and was for a long time combined with the rodents, the group being dist inguished as th~ sub-order Duplicidentata, characterised by having a small second upper incisor (Southern 1964). Barrett-Hamilton (1912) placed hares and rabbits with the rodents but acknowledged that further subdivision was inevitable. The whole group was later revised and a new order was created after 1 evidence showed that the' two divisions ~ (rodents and the hares and rabbits) were distinct by the Eocene (about 60 million ybp) (Matthews 1952; Southern 1964). The current situation Order: Lagomorpha, Family; Leporidae, and Genus; Lepus. The lagomorphs are small to medium sized quadrapedal mam mals clothed in soft fur. Many have a short tail but some are tail-less. They are digitigrade or digitigrade on the fore feet and plantigrade on the hind feet. The of all feet are fur, covered and the toes each have a claw. A wide diastema is created due to the lack of canine teeth and the incisors are chisel shaped, open rooted and grow continuously. The dental featUre distinguishing 1agomorphs from rodents is the presence of an extra pair of incisors set behind the upper incisors in the lagomorphs, whereas rodents have only one pair of incisors in each jaw (Horne 1978). other less marked skull features fur- ther distinguish the lagomorphs from rodents. 2 In lagomorphs most food passes through the digestive system twice. Refection occurs during the day (Watson and Taylor 1955), fecal material being reingested directly from the anus. It is thought that refection provides the animal with vitamin B produced by bacterial action in the caecum but not absorbed there (Matthews 1971). The Order Lagomorpha contains two distinct families; 1. Ochotonidae These are the pikas or mouse hares found in Eastern Europe, Asia and mountains of western North America. They have consid erable skull differences from hares and rabbits (Southern 1964). 2. Leporida~ The family Leporidae contains nine genera and 52 species of rabbits and