Selective Predation in a Polyphagous
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r Robert George Holmberg NAL~a AUTHOR/NW DE L*AUTEUR ~eleciivepredation by a generalized invertebrate predator, TITLE THESIS/TlfRE DE LA T~SF ra - Pardosa vancouveri (Atachnida Araneae) Dr. A. L. Turnbull ~iii4yP SllglVmw~~~ou oImr&x 6. Permission is granted to the NATIONAL LleRARY .CF .L'autorisatim est, psr la prlsente, .ccord/e b /a WMIOT~~&- C CANADA to micrdilm this thesis end to lend or sell copier OUE NATIONALE DU CANADA 'de micrdihw cone thdse et . of the film. k prhr w & vetdm dm examp/ainr du film. The aulha osrws dhr &liution rights, md nitkti& L'Mw se rIINW ks wru c)oits da pdlicatim; ni la themis na extrrsiw extracts from it mybe pintal or dmr- tMsew de tmgs emits k celhci nr dohrsnZ 8tre i~ly~imdr 12330 - 103 Street PsmAwenr Aooa€swrr4~~~~NH Edmonton, Alberta TK: 2K3 ' I+, National Library of Canada Bibliothkque rmtionale du Canada - Catalqying Branch i' Direction du m#Coglrge' Canaban Thesss Division . Division des thbscarfadienm . - Ottawa. 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Veuillez prendre connai~s~ncedes for- this thesis. mules d'autorisation -qui accompagnent cette thh. a. - i - B 0 8 C a i 0 I - TtUS DISSERTATION LA THESE* A'CT~ HAS BEEN MICROFILMED MICR~ILMCEEL QU~ , EXACTLY AS RECEIVED NOUS L'AVONS hE CUE" ' . ' *- SE~CTIVEPREDATION IN A PoLYPIumas . 0 I INVERTEBRATE PREDATOR; Pqrdosa vanrowerl b - (AQ~NIM, 1 I, . - b Y i Robert George Holmberg B.A.(bns.), University, of Saskatchewan, 1970 - , - -. M.Sc., University of Saskatchewan, 1971 r A THESIS SUBUITTED IN PARTIAL FULFILLPIENT OF WE REQUIREHENTS FOR THE. DEGREE OF DOCTOR OF PHILOSOPHY. i- i~ the Department 0 - .. Biological Sciences - % * 4, * ~ b s I ' .. @ Robert George Holmberg 1978 \ SIMON FRASER UNIVERSITT - July 1978 All rights reserved. This thesis may not be reproduced in whole or in part, by photocopy or'other means, without permI.ssion of the author. Frontispiece. Adult female Pardosa vancouveri feeding on a "smallt1 Tenebrio molitor larva (X 11). Robert George Holmberg Doctor of Philosophy Selective predation by a generalized invertebrate predator,. Pardosa vancouveri (Arachnids Araneae) Examining Cormittee: Chairman: Dr. Robert C. Brooke, Chairman Dr. A. L. Turnbull, Sen$or Supervisor . '. Dr. J.~.M. Hackauer Dr. R. n. F.S.Sadleir 4 -Dr. Ga. Geen, N *-he Examiner - Dr. W. G. Wellington,-xternal Examiner Institute of Animal &source Ecology University of British Col-ia PARTIAL COPYRIGKT LICENSE I hereby grant to'simon Fraser. _ ,bDi"eisirty the right to lend I . - my thesis, project or extended essay (the title of'which is shown below) ;to users of heSimon Fraser University Library, and to make partial or singlecogiesonly.for such usersor in Fesponsetoa request fromthe - - 4 any , l i brary of other university, or other educational 'institution, on r: its own behalf or for one of t.s users. I'further agree that permission ,. .* for multiple &opy.i ng of thy woik f~rscholarly purpo%es may be granted % by me ch- the Dean of Graduate Studies. It. is understood that copying . ' or pub1 ibation df this mr,k for financial gain shall not be allowed , . without my written permission; Title of~Thesis/Project/Extended Essay * Selective.predation by a generaliaed invertebrate pte&$or, Pardoma 1 flncouveri- (Arachnids Araneae) Author: (signature) Robert George Holaberg ABSTRACT " " * li 1 v Selacbive predation (i-em, no.-r&adom feeding) ms sthdied in'i- laboratory system that al-lowed individuul . , lyco44 spiders, m,to feed on prey pqpulatiods one or. tro kinds o'f insects- * Ia,rioas corrbimtions of male and female frait flies . , . ' (- ..1..oQ.OtQT), asmall@and @largeebeetle larvae . m)were ased as prey- SpidQrs of both sexes and tuo age classes Here .used as predators, In 14. expe imeats, the spiders showed strong selection in six, moderate to weak eelection in five. and very weak or no .selection' in &ee. Generally, fruit flies were salected by the spideqs over mealuorms or milkweed bugs, ;nd @small@mealuorms over 'large' ones. Very weak or no 'selection occurred between male and female flies and between beetle larvae and milkweed bugs- V r Sobadult and adult spiders always shored similar selective tendencies- Prior feeding experiences did not, alter feeding' selections, but changes ia the phpsiul complexity of the environment did- Thus, by: adbing tree leaves to the cages, the a~mberof flies eaten by the spiders decreased sig~ificantiy. ' . C Potential benefits to the spiders which were masured included percent maturing. per&nt qurviving. rate% weight iii gain.' size gqin. and, proddctioq of false egg%acs. In prey f acombinations . that prodoc& moderate to otrong pdection, 7 L' *. out,gf 28 benefit measurhitnts statistically favored. .the spiers. 'prey combinations that oprodded w&k iqelection - tendencies gave no benefit measurements (H' = 26) that . *I statistically favored the spiders- Spiders fed t ha less- r selected prey exhibited no benefits that were sup=rior. to , those fed the &re-selected prey. Spiders fed the .om- .% selected prey did just as well as spiders fed both prey, It is hypothesized that this particular selective predation - 0 process involves prey sampling, memory of prey attributes, *3 and selection ?S prey that possess certain attribates- r Selected prey seem to confer certain bencifits to the predator- As most beaef its were associated with idcreahd biomass yithin time limits, g- tends to be more an energy maxiaizer thanr a time, minimizer, C * I thank mr senior supervisor, Dr. A. Tarnbull, for the I opportunity Co do this project 'and for his- helpful advice. I >. dso thank the other members of the department for serving on ay supervisory-. committee: Dr. L. Dill;, Dr. S. Haien (deceased), Dr. 8. lat kauer, Dr. D. ,Ba PeCott, and Dr. B. Sadleii. I am '~r.J. Hardran, Dr'. R. Leech and ," 0 T Bakshi ' reading %hi'mantiscript; ~f. C. .Villegas- for +tatisti&l advice; P. Long for taking photographs; C, Holmberg and J. Henigman forahelping with the ueighings; I. Pain for text entry; J. Bechamp for '. L ' helping with the figures; and the people of the. Simon Praser . ~n'iversityComputing Centre and, Athabasca University b Computing Services for helping solve the computer problems. I am deeply g,rateful to my uife, Catherine, for her * patience and understanding. Financial assistance uas provided by ~imonlraser a university and by a Grant from the national nebarch COUDC~I' of Canada to Dr. TarDball. TABLE OF COlITtlTS Page APPROVAL ii 8 I ABSTRACT iii v TABLE OF COITBlTS q .. .+ t- LIST OF TABLES . 4 i~ I. 4 ..,' LISP OF FIGURES ' 'xiii - 2. Significance of Selective Predation, to Biology 3 3. Components of Selective Predation a. Availability Factors b. Catchabilit y Pactors c. Preference Factors 10 4- Past Studies of Selective Predation 13 <c 5. ~conouc~odels 17 0. a. Consumption Clodels - 18 \ b. Production odei is . - c. Producer and cons&er aodels 19 ,' 6. Utility of .the Prey Selection Process to Predators v, 2 1 r 7, ~bjwtitw,Approach and Limitations .of Present stid* 26 . # + * 1 TABLE OF CoITBrTs qoatinued) . i. / J Page 3. ~e.hfit.Criteria - . 4. Relationship Between Sel-ection and Beaefit ip *. 5. Heckanisms &hind the Selective Predation . - Process 9 3' 6. Why kt Hore fhan'0ne .Kind of 103 7. as an 106. COKLUSIOIS 109 - APPEliQIX I, Specific Informatid. on the Various Predator and Prey Classek B +' APPBIDIX 11, Details -of the Selective Predation ' . ., 1 ' .) Zx~eriments APPBIDIX III, Details of the Utility of the Prey to t) Impert&nt Terms Used :in- I , / the Text ' AP+iDIX P, List of Ion-Staqdard ~bbrAidtioes viii ,' ,* * Table I. Summary .of 14 selective +predatioo+experiments , -t . - ranked ac&=ding to the relathe disrimiF&&on . coefficient; d 6 5. C '\ - . Table XI. peading rates of p. ranso.leri ih,tro prey swems' \ * C * ' consisting of a. and Zm W afar P being pre-fd with me 03 these prey. 68 /.- r-- ' Tabla XII. ~esdi&of increasing tho phYsi&l complexity of . the cages by adding tbree alder leaves. - 70 t Table IV. Benefit criteria that- were statistically 1 signifi'cant in faGr of those spiddrs that ate tie 7. more-selected prey. 72 ' Table 7. Percentage of benefit criteria that favored the r. - .. , more-selected prey. 74 . *.- t 0 t. 0 ' Table II.