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The Possibility of the Existence of Mimicry Between Two Species Of University of Tennessee, Knoxville Trace: Tennessee Research and Creative Exchange Doctoral Dissertations Graduate School 3-1962 The oP ssibility of the Existence of Mimicry Between Two Species of Southern Appalachian Salamanders Lowell Preston Orr University of Tennessee - Knoxville Recommended Citation Orr, Lowell Preston, "The osP sibility of the Existence of Mimicry Between Two Species of Southern Appalachian Salamanders. " PhD diss., University of Tennessee, 1962. https://trace.tennessee.edu/utk_graddiss/3255 This Dissertation is brought to you for free and open access by the Graduate School at Trace: Tennessee Research and Creative Exchange. It has been accepted for inclusion in Doctoral Dissertations by an authorized administrator of Trace: Tennessee Research and Creative Exchange. For more information, please contact [email protected]. To the Graduate Council: I am submitting herewith a dissertation written by Lowell Preston Orr entitled "The osP sibility of the Existence of Mimicry Between Two Species of Southern Appalachian Salamanders." I have examined the final electronic copy of this dissertation for form and content and recommend that it be accepted in partial fulfillment of the requirements for the degree of Doctor of Philosophy, with a major in . J.T. Tanner, Major Professor We have read this dissertation and recommend its acceptance: Accepted for the Council: Dixie L. Thompson Vice Provost and Dean of the Graduate School (Original signatures are on file with official student records.) March 12, 1962 To the Graduate Council: � am submitting herewith a dissertation written by Lowell Preston Orr entitled "The Possibility of_ the Existence of Mimicry Between Two Species of Southern Appalachian Salamanders.�· I recommend that it be accepted in partial fulfillment of the requirements for the degree of Doctor of Philosophy, with a major in Zoology. We have read this dissertation and recommend its acceptance :· � :?ttZr.&'H� Accepted for the Council: \� /2� Dean of the Graduate School THE POSSIBILITY OF THE EXISTENCE OF MIMICRY BETWEEN TWO SPECIE S OF SOUTHERN AP PAlACHIAN SALAMANDERS A Dissertation Presented to The Graduate Ccmncil of The University of Tennessee In Partial Fulfillment of the Requirements for the Degree Doctor of Philosophy by Lowell Preston Orr March 1962 ACKNOWLEDGMENTS I am grateful to many individuals for their assistance in this study: Merlin Tuttle for his assistance in obtaining shrews used in the feeding experiments, M. T. Orr for advice and assistance in obtaining the photographs, and A. Stupka and N. J. Reid for permission to use the Great Smoky Mountains National Park facilities for the study. Stimulating comments and criticisms from R. E. Gordon, J. T. Wood, and J. E. Huheey were very much appreciated. I am especially indebted to Dr. J. T. Tanner, of the Zoology Department, University of Tennessee1 for his invaluable guidance and criticisms in d:irecting this problem. This investigation was supported in part by grants from the Tennessee Academy of Science and the Sigma Xi - RESA. Research Fund. ii 52541.9 TABLE OF CONTEN'IS SECTION PAGE I. DJ'ffi.ODUCTION . 1 II. REVIEW OF THE LITERATURE ••. • • • • • • • • • 9 The Theory of Mimicry • • • • • • • • • • • • • • • • • 9 The Ecology and Speciation of D. o. carolinensis and f• j. jordani • • • • • • • • • • • • • • • • • • • • 13 III. THE REGION OF THE STUDY • • • • • • • • • • • • • • • • • 19 IV. THE COIDR PATTERN OF ]2. Q. CAROLJNENSIS • • • • • • • • • 20 V. THE RANGES OF f.. �· JORDANI AND ]2. £. CAROLJNENSIS • • • • 32 Methods • • • • • • • • • • • • • • • • • • • • • 32 Results • • • • • • • • • • • • • • • • • 33 VI. DETERMJNATION OF THE RELATIVE· ABUNDANCE OF THE SUPPOSED MIMICS AND MODElS • • • • . • • • • • 4o Methods . 4o Description of the Transects • • • • • • • . 41 Results • • • • • • • • • • • • • • • • • • • • • • • • 44 VII. FEEDJNG EXPER lMEN TS. • • • • • • • • • • • • • • • • • • • 51 Thamnophis sirtalis sirtalis • • • • • • • • • • • • • • 51 Blarina brevicauda • • • • • • • • • • • • • • • • • • • 58 GJ!inophilus danielsi danielsi and Desmognathus quadr·a-macula tus • • • • • • • • • • • • • • • • • • • 64 Preliminary Feeding Experiments with Other Predators • • • • • • • • • • • • • • • • • • • • • • 65 iii iv SECTION PAGE VIII. DISCUSSION • • • • • • • • • 70 IX. SUMMARY • • • • • • 77 BIBLIOGRAPHY • • • • • • • • • • • • • • • 80 LIST OF TABLES I. The Presence or Absence of �· i• jordani and �· £· carolinensis in Fourteen Areas of the Great Smoky Hountains National Park • • •• • • • . • • • 35 II. Records of Red-cheeked carolinensis Occurring Outside the Range of f• i• jordani • • • • • • • • • • • 37 III. Relative Abundance of jordani, Non-Mimetic carolinensis, and Mimetic carolinensis in Eight Transects • • 1' • • • ' . • • • • • 45 v LIST OF FIGURES FIGURE PAGE 1. Red-Cheeked Phase of Desmognathus ochrophaeus carolinensis Dunn ••• . • • 2 2. Plethodon jordani jordani Blatchley . � 3 3. Normal Color Pattern of Desmognathus ochrophaeus carolin�nsis Dunn • • • • • • • • • • • • • • . • • 4 4. Phylogeny of the Family Plethodontidae • . • . • • • • • • • 17 A Color Variant of n. o. carolinensis • • • • • • • • • • • • 22 . • . • . • • • • • 23 6. A Color Variant of D. £· carolinensis • • ?. A Color Variant of D. o. carolinensis • • • • • • • • • • • • 24 B. A Color Variant of D. o. carolinensis • • • • • . • • • • 25 - • 9. A Color Varifl,nt of -D. o. carolinensis • • • • • • • • • • • • 26 10. A Color Variant of D. o. carolinensis • • • • • • • • • • • • 27 11. A Color Variant of D. £• carolinensis • • • • . • • 28 12. A Color Variant of D. £• carolinensis • • • • • • • • • • • • 29 13. A Color Variant of D. £• carolinensis • • • • • • • • • • • • 30 14. Areas Sampled fo� �· i• jordani and D. o. carolinensis in the Great Smoky Mountains • • • • • • • • • • • • • • • 34 15. Relative Aqundance of jordani and Red-Cheeked carolinensip • • • • • • • • • • • • • • • • • • • • • 47 �6. Correlation between the Number of jordani in Transects ru1d the Per Cent of Red-Cheeked carolinensis in the Total carolinensis Population • • • • • • • • • • • • • • • 50 vi rli FIGURE PAGE 17. Feeding Experiment Results of the Control Group of Thamnophis sirtalis • • • • • � • • • • • • • • • 56 18. Feeding Experiment Results of the Experimental Group of Thamnophis sirtalis • • • • • • • • • • • • • • • 57 19. Feeding Experiment Results of the Control Group of Blarina brevicauda • • • • • . "62 20. Feeding �eriment Results of the Experimental • • • • 6 Group of Bla�ina brevicauda • • • • • • • • • • • • 3 I. INTRODUCTION Investigators familiar with the amphibia of the Great Smo� Mountains have long-·been puzzled by the amazing resemblance between the red, orange, or yellow-cheeked variants of the DesmognathI us ochrophaeus carolinensis Dunn population (Fig. 1) and the red-cheeked salamander, Plethodon jordani jordani Blatchley (Fig. 2). In �is study� these two species will be referred to by their sub-specific names, carolinensis and jordani. The problem is especially intri�ing in that the frequency of the red-cheeked variant in carolinensis ·is highest where this polymorphic species is sympatric with jordani in the pmok!es, Outside the range of jordani the incidence of the red cheek coloration among normally patterned carolinensis (Fig. 3) is quite lmv-. Numerous individuals have suggested that the similari�y ean be best explained in ter;ms of mimicry. According to the theory of mimicry, the two distantly related species ar e protected by having s.imila:r;- colpr patterns which ttadvertise11 to predators an undesirable characterist�c found in one or both of the species sharing the color pattern. TWo qasi� types of m.imicry have been proposed, In Batesian m.imicry, the red chi;:Jek patch of jordani could be considered warning or aposematic coloratiop which would 11advertise11 to predators some distasteful or unde-sirable quality found ill. jordani (the model). The red-cheeked variant of carol.inensis (the m�ic ), which exhibited pseudaposematic or .false warning coloratio� would thus gain survival value from its close 1 2 FIGURE 1 #. ' - RED-CHEEKED PHASE OF DESHOGNATHUS OCHROPHAEUS CAROLJNENSIS DUNN 3 r _,. FIGURE 2 PLETHOION JORDANI JORDANI BLATCHLEY FIGURE 3 NORMAL COLOR PATTERN OF DESMOGNATHUS OCHROPHAEUS CA ROLINENSIS, DUNN resemblance to the model even though it possessed none of the undesirable qualities. In Mullerian mimicry, both j ordani and the red-cheeked carolinensis would possess undesirable qualities and thus the model and the mimic �e both protected by undesirable characteristics. Although this problem has been recognized for over thirty-five years, no field research has been conducted to actually test the hypoth­ esis of mimicry between jordani and carolinensis. The purpose of this research project is to investigate the possibility that mimicry exists bett.reen these two species by both field and laboratory approaches. Dun� (1927) first described the yellow-cheeked variety of Desmognathus as a new mountain race and referred to it as Desmognathus fuscus imitator. Later (1928) Pope concluded from his studies that it vJas merely an unstable color variety of Desmognathus fuscus carolinensis. Bishop (1947), however, regarded it as a subspecies of Desmognathus ochrophaeus and assigned it its present status of Desmognathus ochrophaeus carolinensis. Attempts to account for this color variation have been made by several investigators. Dunn (1927) concluded that his specimens eY�ibited 11a clear case of mimicry," and that "coincidence
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