GEOGRAPHICALLY DISTINCT BUT NON-MONOPHYLETIC MORPHS: REEXAMINING THE EVOLUTION OF COLOR IN FLORIDA BURROWING BEETLES (COLEOPTERA: GEOTRUPIDAE: PELTOTRUPES) By Emily M. Scott A Senior Honors Signature Project Presented to the Honors College East Carolina University In partial Fulfillment of the Requirements for Graduation with Honors By Emily Scott Greenville, NC December, 2019 Approved by: Michael S. Brewer, Ph.D. East Carolina University, Department of Biology 1 Geographically distinct but non-monophyletic morphs: Reexamining the evolution of color in Florida burrowing beetles (Coleoptera: Geotrupidae: Peltotrupes) Emily M. Scott Department of Biology, East Carolina University, Greenville N.C. 27858-4353, U.S.A. ABSTRACT - The genus Peltotrupes Blanchard 1888 (Coleoptera: Geotrupidae) is restricted to scrub and sandhill habitats on peninsular Florida’s ridge system. Commonly called Florida Deep Digger Scarabs, individuals excavate burrows as deep as 2-3 m and were traditionally considered to be flightless. The recognized diversity within the genus (P. profundus profundus Howden 1952, P. p. dubius Howden 1955, and P. youngi Howden 1955) is based mostly on structural color differences corresponding to distinct geographic regions (i.e., dune ridges). Herein, we provide the first molecular phylogeny of the genus Peltotrupes and highlight the lack of monophyly of the currently recognized species (i.e., color morphs). We also analyze the color of four dorsal regions of male individuals representing the full geographic and color phenotypic breadth of the genus. Although significant color differences are recovered between the currently recognized species, given their lack of monophyly, our results suggest an ecological role may be maintaining these distinct morphs. 2 Acknowledgments I would like to thank the following people for their contributions and support to the completion of this thesis: Dr. Michael Brewer, Dr. Trip Lamb, the Department of Biology, and East Carolina University. Without the advising and guidance of Dr. Michael Brewer, my research professor and project advisor, I would have been unable to proficiently complete this project. Further, the support and guidance from Dr. Trip Lamb were beneficial to the completion of this project. Additionally, I would like to thank the NC State University Insect Museum and the University of Michigan Museum of Zoology for their generosity in sharing their Peltotrupes collections for our research. 3 Table of Contents Introduction - Background 7 - Research Questions 8 Methods - Collection 9 - Molecular Work 9 - Photography 9 - Color Analyses and Protocol 10 Results - Brief Synopsis of Molecular Work 12 - Color Analysis 12 Discussion 14 Conclusion 15 Literature Cited 16 Appendix 17 4 Appendix Peltotrupes Color Measurement Protocol 17 Table 1. Shows all of the samples collected and used for color 19 analyses from peninsular Florida. Table 2. Shows all RGB values for each specimen and dorsal 33 region measurements. 5 List of Figures Figure 1. Peltotrupes youngi in sand-hill habitat. 7 Figure 2. Peltotrupes profundus in sand hill habitat. 7 Figure 3. Peltotrupes mound habitat along sand ridges of peninsular Florida. 8 Figure 4. Peltotrupes mound and burrow entrance. 8 Figure 5. High-definition photograph of Peltotrupes youngi. The photograph used 10 for color analysis. Figure 6. High-definition photograph of Peltotrupes p. profundus. 10 The photograph used for color analysis. Figure 7. High-definition photograph of Peltotrupes p. dubius. 10 The photograph used for color analysis. Figure 8. Peltotrupes profundus illustrating locations measured for color. 11 Figure 9. SPLITSTREE phylogenetic network of mitochondrial haplotypes. 12 Tip labels are color coded as Peltotrupes p. profundus (purple), P.p. dubius (blue), and P. youngi (green). Figure 10. Color frequency distributions (as integers) for the four dorsal 13 regions and associated statistical results. Taxa were compared via ANOVAs and results were investigated using Tukey’s post-hoc test. Significant results are bolded. Figure 11. Principle component analysis and PERMANOVA results 13 for color measurements. Significant results are bolded. 6 Introduction Background: The beetle genus Peltotrupes (Blanchard, 1888) is native to peninsular Florida and is composed of two species, P. profundus, and P. youngi (Howden, 1952). They are seemingly flightless beetles and are known for their iridescent coloration. Additionally, they are known as deep digger scarab, and dig deep burrows to rear their young and inhabit during the day time. Along with these species, there is a subspecies, P.p dubius (Howden, 1955). Although P.p. dubius subspecies has been classified as a subspecies of P. profundus, this was originally dismissed and has been questioned since. These species were once classified under the genus Geotrupes, but this was changed to Peltotrupes by Henry Howden after he divided the genus into the two species: profundus and youngi (Howden, 1952). Peltotrupes profundus generally have a purple coloration along their elytra while P. youngi are typically green. The independent species within Peltotrupes with their robust, iridescent-coloration occupy separate geographic regions within central Florida (Kalisz, et al., 1984). Additionally, degree of coloration and other anatomical structures make these beetles sexually dimorphic (Howden, 1955). Figure 1. P. youngi in sandhill habitat. Figure 2. P. p. profundus in sandhill habitat. 7 These beetles are considered to be distinct species; however, through recent molecular phylogenetic analysis (Brewer lab) indicates the two species are not reciprocally monophyletic. These new data raise new questions regarding their physical characteristics that distinguish the species. Given the inconsistencies between genetic and morphological data, the purpose of this study was to examine color variation within and between the two species. Figure 3. Peltotrupes mound habitat. Figure 4. Up-close mound and burrow entrance. Research Questions: (1) Do the nominal Peltotrupes taxa comprise reciprocally monophyletic groups? (2) Does the minor coloration used to delineate species and subspecies significantly differ? 8 Methods Collection: Beetle specimens were collected from sandhill regions throughout peninsular Florida, and specimen names, species, and localities were recorded in Excel (Table 1). These beetles were collected in pitfall traps (buried coffee cans baited with sucrose/yeast mixture). Molecular Work: Beetles collected were euthanized and pinned, and the right rear leg was removed for DNA extraction using Qiagen’s DNeasy kit. After extraction, the cytochrome oxidase 1 (COX1) and cytochrome oxidase 2 (COX2) genes were amplified using Polymerase Chain Reaction (PCR). After additional clean up and gel electrophoresis, the samples were sequenced, and a SPLITSTREE phylogenetic network of mitochondrial haplotypes (Figure 8). Photography: We generated composite photographic images for 101 male Peltotrupes beetle specimens representing the wide-range distribution of the genus for color analysis. Specimens were photographed using a Canon 6D camera under controlled light settings to minimize glare and ensure all specimens had similar lighting. Eight to twelve photographs were taken of each specimen and were exported to Helicon Focus, which pieced all images together to create one high definition composite image used for color analysis. 9 Figure 5. High-definition photograph of Figure 6. High-definition photograph of Peltotrupes youngi. Image used for color Peltotrupes p. profundus. Image used for analysis. color analysis. Figure 7. High-definition photograph of Peltotrupes p. dubius. Image used for color analysis. Color Analyses: Composite images were exported to GIMP, a photograph editing software. The GIMP “color-picker” tool was used to measure RGB values for four dorsal areas of each specimen and a standardized grey background. The four dorsal areas included: lateral elytrum, edge elytrum, anterior, and head (Figure 8). RGB values standardized by dividing each value by the corresponding grey background value, then multiplied by 255, the RGB value of white (The protocol of measuring color values is described in the Appendix). RGB values were then converted to hexadecimal values and integer values using the R packages RGB and STROI to provide single 10 continuous values for analysis. Specimens, grouped by taxon, were subjected to ANOVA, followed by Tukey’s post-hoc tests in R. Lastly, principal component analysis (PCA) was used to cluster taxa using all four dorsal measurements and subjected PERMANOVA to test for differences between taxa (999 iterations). Figure 8. Peltotrupes profundus illustrating locations measured for color. 11 Results Brief Synopsis of Molecular Work: The SPLITSTREE phylogenetic network revealed phylogeographic structuring among mtDNA haplotypes, with four distinct phylogroups (Figure 9). However, none of the three nominal taxa were resolved as being monophletic. Instead, all of the haplotypes representing P. youngi and P.p. dubius clustered with those of P. p. profundus from the northern portion of its geographic range . Figure 9. SPLITSTREE phylogenetic network of mitochondrial haplotypes. Tip labels are color coded as P.p. profundus (purple), P.p. dubius (blue), and P. youngi (green). Color Analysis: Color morphs (i.e. species) appear to comprise distinct entities. Further, the “lateral” and “anterior” elytral areas best differentiate these morphs based on p-values from statistical tests (Figure 10). The lateral region separated P.p. profundus and P. youngi significantly, while the anterior region separated P. youngi and P.p. dubius as well as P.p. profundus and P. youngi significantly. PCA clustered taxa based on the four dorsal regions, and PERMANOVA showed differences in taxa through statistical analysis. Based on this test, P. youngi and P.p. dubius 12 are separated significantly based on color morphs, as well as P.p. profundus and P. youngi (Figure 11). Figure 10. Color frequency distributions (as integers) for the four dorsal regions and associated statistical results. Taxa were compared via ANOVAs and results were investigated using Tukey’s post-hoc test. Significant results are bolded. Figure 11. Principle component analysis and PERMANOVA results for color measurements. Significant results are bolded. 13 Discussion Although the mitochondrial gene analysis did not support the monophyly of the three taxa of Peltotrupes, their color morphs appear to comprise distinct entities. While these taxa are not supported genetically, they are supported through their color morphs. Interestingly, these beetles are burrowers and spend their time above ground in darkness (Howden, 1955). Further, the iridescent coloration that distinguishes them is structural, meaning it is caused by light rather than pigment. With this knowledge, it is interesting to see how strongly color separates these burrowing beetles, while mitochondrial genetic analysis does not. Although these beetles inhabit geographically distinct regions, there is no phylogenetic structure defining the distinct species. These inconsistencies and the results from the color analysis raise a new question: why do discrete color morphs exist despite apparent presence of gene flow? 14 Conclusion The analysis of genetic relationships among defined species and subspecies of Peltotrupes showed that these taxa are not monophyletic, as was previously thought. However, distinct coloration does appear to distinguish them, particularly from the “lateral” and “anterior” elytral regions. The ANOVA and the PERMANOVA results support significant differences between P.p. profundus and P. youngi as well as between P.p. dubius and P. youngi. However, no significant differences were observed between the subspecies P.p. profundus and P.p.dubius. Differences in the genetic and morphological results lead us to believe that an ecological role is at play in maintaining the distinct color morphs. 15 Literature Cited 1. Blanchard, F. (1888). Some account of our species of Geotrupes. Psyche: A Journal of Entomology, 5(149-150), 103-110. 2. Howden, H. F. (1952). A new name for Geotrupes (Peltotrupes) chalybaeus LeConte, with a description of the larva and its biology (Scarabaeidae). The Coleopterists Bulletin, 6(3), 41-48. 3. Howden, H. F. (1955). Biology and taxonomy of North American beetles of the subfamily Geotrupinae, with revisions of the genera Bolbocerosoma, Eucanthus, Geotrupes, and Peltotrupes (Scarabacidae). Proceedings of the United States National Museum. 4. Kalisz, P. J., & Stone, E. L. (1984). Soil mixing by scarab beetles and pocket gophers in north-central Florida. Soil Science Society of America Journal, 48(1), 169-172. 5. Magsig, C., Lamb, A., Brewer, M. (2017). Evaluating the evolutionary history and taxonomy of the deep-digger scarab beetle genus Peltotrupes. Biology Honors Thesis, East Carolina University. 16 Appendix Peltotrupes Color Measurement Protocol **DO NOT TAKE MEASUREMENTS WHERE THERE IS A GLARE ON THE SPECIMEN** Step 1. Open GIMP. Step 2. Select a file to edit. - Select “okay” for TIFF image Message pop up and select “convert” for Convert to RGB Working Space. Step 3. Once file is open, zoom in on location being measured (*command, scroll*) and select the “Color Picker Tool”. Note: this looks like an eyedropper. - After selecting tool, mark “sample average” box of the bottom left toolbar, so the radius can be adjusted. - Mode should be on “set foreground color”. Step 4. Adjust radius to 50 and select an area of grey near specimen, but be sure the area is consistent without any difference in coloration or brightness. Note: the best place is usually right above the head, to the right of the first leg. - To get the RGB data of the area, use the method: “shift key, command key, and click” and the values will appear in a small box. - Record grey measurements for red, green, and blue in spreadsheet for lateral, edge, anterior, and head. Step 5. Adjust radius to 10 and select area of the elytrum along the edge where color is most reflective/colorful. Color is most consistent/standard in the thickest part of the lateral area. - There will be two measurements for this area-one along the lateral (inside ridge) and one along the very edge of the elytrum. Note: there is a definite “ridge” that separates the lateral and edge measurements. - Collect the RGB values for both areas of measurement and record them in the spreadsheet (one under “lateral” and the other under “edge”). - Note: the radius may need to be adjusted to a smaller size in order to get a clean measurement of the edge. **MAKE SURE TO GET A CLEAN, CONSISTENT SAMPLE** Step 6. Keep radius at 10 and select area at the top of the elytrum, right as it is joined to the thorax. - Pick a spot where the color is reflective and colorful and record the RGB values in the spreadsheet 17 - Note: count 4-6 lines left of the center of the elytra and get a measurement in this vicinity. Step 7. With the radius still at 10, select area at the base of the head (right where the head and thorax connect). - Pick a spot (without glare) between the center of the head and the left eye and record RGB values into spreadsheet. Step 8. To measure the data in Excel, type “=” and then the cell number of the color of the sample and divide that by the cell number of the same color for the grey measurement. Once doing the first box for the correction, you will be able to shift down the same formula. - EXAMPLE: To get the corrected value for each RGB value, if the R value for grey measurement is 205 (and in cell B3), and the R value for the “lateral” measurement is 132 (and in cell C3), then in the cell for the corrected value, write “=C3/B3” and “enter”. - NOTE: this should give a small decimal value? once getting this value, drag the bottom right corner of the cell across to get the rest of the Green and Blue values, and then drag down to do the same for the remaining samples of each specimen. - Do the same thing for the edge, anterior, and head recorded measurements. 18 Grey - Lateral Sample - Lateral Corrected RGB - Lateral Standardized RGB - Lateral Species Sample R G B R G B R G B R G B P120 211 203 192 84 63 91 0.39810427 0.31034483 0.47395833 90 67.5 97.5 P. profundus P142 202 193 181 65 97 114 0.32178218 0.50259067 0.62983425 60.0543478 89.6195652 105.326087 P. youngi P002 201 190 178 35 81 104 0.17412935 0.42631579 0.58426966 40.5681818 93.8863636 120.545455 P. youngi P008 202 192 181 66 47 84 0.32673267 0.24479167 0.4640884 85.4314721 60.8375635 108.730964 P. youngi P009 179 165 144 30 58 82 0.16759777 0.35151515 0.56944444 45 87 123 P. youngi P010 194 183 169 56 86 129 0.28865979 0.46994536 0.76331361 52.6937269 80.9225092 121.383764 P. youngi P011 188 177 161 32 79 86 0.17021277 0.44632768 0.53416149 41.4213198 102.258883 111.319797 P. youngi P012 204 191 175 89 67 87 0.43627451 0.35078534 0.49714286 93.3950617 70.308642 91.2962963 P. youngi P014 200 188 174 87 116 114 0.435 0.61702128 0.65517241 69.9842271 93.3123028 91.70347 P. youngi P015 193 183 170 64 106 122 0.33160622 0.57923497 0.71764706 55.890411 92.5684932 106.541096 P. youngi P016 204 192 178 59 70 92 0.28921569 0.36458333 0.51685393 68.0769231 80.7692308 106.153846 P. youngi P017 208 197 183 79 63 96 0.37980769 0.31979695 0.52459016 84.6428571 67.5 102.857143 P. youngi P018 208 197 183 78 100 121 0.375 0.50761421 0.66120219 66.5217391 85.2842809 103.19398 P. youngi P019 206 196 183 70 110 108 0.33980583 0.56122449 0.59016393 61.9791667 97.3958333 95.625 P. youngi P020 209 198 184 102 86 62 0.48803828 0.43434343 0.33695652 104.04 87.72 63.24 P. youngi P021 199 186 169 53 62 85 0.26633166 0.33333333 0.50295858 67.575 79.05 108.375 P. youngi P022 204 192 178 67 54 110 0.32843137 0.28125 0.61797753 73.961039 59.6103896 121.428571 P. profundus P031 200 187 172 58 75 80 0.29 0.40106952 0.46511628 69.4366197 89.7887324 95.7746479 P. youngi P032 204 193 179 73 120 122 0.35784314 0.62176166 0.68156425 59.0952381 97.1428571 98.7619048 P. youngi P034 204 192 176 77 83 127 0.37745098 0.43229167 0.72159091 68.4146341 73.7456446 112.839721 P. profundus P040 202 194 184 16 108 145 0.07920792 0.55670103 0.78804348 15.1672862 102.379182 137.453532 P. youngi P049 209 200 188 77 67 102 0.36842105 0.335 0.54255319 79.8170732 69.4512195 105.731707 P. youngi P050 203 193 181 30 74 98 0.14778325 0.38341969 0.54143646 37.8712871 93.4158416 123.712871 P. youngi P051 203 193 181 19 115 131 0.09359606 0.59585492 0.72375691 18.2830189 110.660377 126.056604 P. youngi P053 202 191 179 52 76 115 0.25742574 0.39790576 0.6424581 54.5679012 79.7530864 120.679012 P. youngi P054 204 194 182 53 57 100 0.25980392 0.29381443 0.54945055 64.3571429 69.2142857 121.428571 P. youngi P055 201 195 182 85 134 145 0.42288557 0.68717949 0.7967033 59.5467033 93.8736264 101.57967 P. youngi P056 201 190 178 84 62 83 0.41791045 0.32631579 0.46629213 93.5371179 69.0393013 92.4235808 P. profundus P057 202 193 182 72 46 76 0.35643564 0.23834197 0.41758242 94.6391753 60.4639175 99.8969072 P. profundus 19 P058 208 198 186 85 60 80 0.40865385 0.3030303 0.43010753 96.3333333 68 90.6666667 P. profundus P059 210 201 190 93 59 85 0.44285714 0.29353234 0.44736842 100.063291 63.4810127 91.4556962 P. profundus P060 212 202 192 86 56 85 0.40566038 0.27722772 0.44270833 96.6079295 62.907489 95.4845815 P. profundus P061 208 202 191 60 81 123 0.28846154 0.4009901 0.64397906 57.9545455 78.2386364 118.806818 P. profundus P062 217 208 198 61 109 158 0.28110599 0.52403846 0.7979798 47.4237805 84.7408537 122.835366 P. p. dubius P064 219 210 201 109 67 96 0.49771689 0.31904762 0.47761194 102.1875 62.8125 90 P. profundus P071 215 207 197 118 93 110 0.54883721 0.44927536 0.55837563 93.7383178 73.8785047 87.3831776 P. profundus P074 211 203 192 68 61 103 0.32227488 0.30049261 0.53645833 74.7413793 67.0474138 113.211207 P. profundus P075 207 202 192 87 86 117 0.42028986 0.42574257 0.609375 76.5 75.6206897 102.87931 P. profundus P078 207 198 188 83 68 103 0.40096618 0.34343434 0.54787234 83.3267717 68.2677165 103.405512 P. profundus P079 212 204 195 74 61 81 0.3490566 0.29901961 0.41538462 87.3611111 72.0138889 95.625 P. profundus P080 210 201 192 103 65 102 0.49047619 0.32338308 0.53125 97.2777778 61.3888889 96.3333333 P. profundus P081 202 193 178 85 69 87 0.42079208 0.35751295 0.48876404 89.9377593 73.0082988 92.0539419 P. profundus P082 207 200 187 122 101 133 0.58937198 0.505 0.71122995 87.3876404 72.3455056 95.2668539 P. profundus P083 210 200 189 83 59 114 0.3952381 0.295 0.6031746 82.6757813 58.7695313 113.554688 P. profundus P086 212 202 191 108 71 90 0.50943396 0.35148515 0.47120419 102.379182 67.3048327 85.3159851 P. profundus P087 P. profundus P088 194 185 173 98 74 117 0.50515464 0.4 0.67630058 86.4705882 65.2941176 103.235294 P. profundus P089 195 179 166 82 75 115 0.42051282 0.41899441 0.69277108 76.875 70.3125 107.8125 P. profundus P091 193 184 171 93 60 99 0.48186528 0.32608696 0.57894737 94.1071429 60.7142857 100.178571 P. profundus P093 194 181 167 86 62 73 0.44329897 0.34254144 0.43712575 99.2307692 71.5384615 84.2307692 P. profundus P094 198 188 175 83 47 81 0.41919192 0.25 0.46285714 100.308057 56.8009479 97.8909953 P. profundus P095 199 185 175 56 77 91 0.28140704 0.41621622 0.52 63.75 87.65625 103.59375 P. profundus P096 190 180 167 91 65 66 0.47894737 0.36111111 0.39520958 104.527027 74.6621622 75.8108108 P. profundus P097 190 176 149 59 81 114 0.31052632 0.46022727 0.76510067 59.2322835 81.3188976 114.448819 P. profundus P098 179 168 150 43 46 91 0.24022346 0.27380952 0.60666667 60.9166667 65.1666667 128.916667 P. profundus P099 201 191 178 69 57 111 0.34328358 0.29842932 0.62359551 74.2405063 61.3291139 119.43038 P. profundus P100 175 165 154 44 66 111 0.25142857 0.4 0.72077922 50.7692308 76.1538462 128.076923 P. youngi P101 196 187 176 61 103 101 0.31122449 0.55080214 0.57386364 58.6981132 99.1132075 97.1886792 P. youngi P102 193 183 172 8 133 159 0.04145078 0.72677596 0.9244186 6.8 113.05 135.15 P. youngi P103 185 172 157 69 96 96 0.37297297 0.55813953 0.61146497 67.4137931 93.7931034 93.7931034 P. youngi 20 P104 191 180 167 34 125 149 0.17801047 0.69444444 0.89221557 28.1493506 103.49026 123.36039 P. youngi P105 191 180 168 38 76 84 0.19895288 0.42222222 0.5 48.9393939 97.8787879 108.181818 P. youngi P114 185 173 161 39 64 82 0.21081081 0.3699422 0.50931677 53.7567568 88.2162162 113.027027 P. profundus P116 185 174 162 64 45 68 0.34594595 0.25862069 0.41975309 92.2033898 64.8305085 97.9661017 P. profundus P119 183 172 156 53 72 139 0.28961749 0.41860465 0.89102564 51.1931818 69.5454545 134.261364 P. profundus P121 194 183 171 72 55 78 0.37113402 0.30054645 0.45614035 89.5609756 68.4146341 97.0243902 P. profundus P122 185 175 163 99 79 118 0.53513514 0.45142857 0.72392638 85.2871622 68.0574324 101.655405 P. profundus P123 P. profundus P125 179 171 157 64 51 78 0.3575419 0.29824561 0.49681529 84.5595855 67.3834197 103.056995 P. profundus P127 178 167 155 49 87 111 0.2752809 0.52095808 0.71612903 50.5870445 89.8178138 114.595142 P. profundus P128 180 172 158 33 61 89 0.18333333 0.35465116 0.56329114 45.9836066 85 124.016393 P. profundus P129 P. profundus P130 199 187 175 108 96 134 0.54271357 0.51336898 0.76571429 81.4792899 72.4260355 101.094675 P. profundus P134 182 171 158 95 73 122 0.52197802 0.42690058 0.7721519 83.5344828 64.1896552 107.275862 P. profundus P135 194 184 173 46 116 147 0.2371134 0.63043478 0.84971098 37.961165 95.7281553 121.31068 P. profundus P136 195 186 174 102 161 183 0.52307692 0.8655914 1.05172414 58.3183857 92.0515695 104.630045 P. profundus P137 201 192 179 9 101 160 0.04477612 0.52604167 0.89385475 8.5 95.3888889 151.111111 P. profundus P138 199 188 176 65 102 155 0.32663317 0.54255319 0.88068182 51.4751553 80.7763975 122.748447 P. profundus P139 210 200 186 77 64 97 0.36666667 0.32 0.52150538 82.5 68.5714286 103.928571 P. profundus P140 200 189 177 59 131 171 0.295 0.69312169 0.96610169 41.6759003 92.534626 120.789474 P. profundus P141 201 189 174 68 99 122 0.33830846 0.52380952 0.70114943 60 87.3529412 107.647059 P. profundus P143 214 207 198 75 69 123 0.35046729 0.33333333 0.62121212 71.6292135 65.8988764 117.47191 P. profundus P144 199 188 175 55 52 104 0.27638191 0.27659574 0.59428571 66.4691943 62.8436019 125.687204 P. profundus P145 203 196 184 77 65 113 0.37931034 0.33163265 0.61413043 77 65 113 P. profundus P147 205 194 182 72 111 116 0.35121951 0.57216495 0.63736264 61.4046823 94.6655518 98.9297659 P. youngi P149 209 203 194 74 59 107 0.35406699 0.29064039 0.55154639 78.625 62.6875 113.6875 P. profundus P152 207 201 191 86 121 155 0.41545894 0.60199005 0.81151832 60.5801105 85.2348066 109.185083 P. profundus P154 180 169 156 39 31 87 0.21666667 0.18343195 0.55769231 63.343949 50.3503185 141.305732 P. profundus P155 185 174 161 58 44 72 0.31351351 0.25287356 0.44720497 85 64.4827586 105.517241 P. profundus P026 173 166 154 45 41 73 0.26011561 0.24698795 0.47402597 72.1698113 65.754717 117.075472 P. profundus P025 205 199 187 129 122 169 0.62926829 0.61306533 0.90374332 78.3214286 74.0714286 102.607143 P. profundus 21 P159 181 173 163 70 72 114 0.38674033 0.41618497 0.6993865 69.7265625 71.71875 113.554688 P. profundus P107 185 174 161 71 59 74 0.38378378 0.33908046 0.45962733 88.75 73.75 92.5 P. p. dubius P157 172 163 148 51 60 110 0.29651163 0.36809816 0.74324324 58.8461538 69.2307692 126.923077 P. profundus P005 175 168 155 39 54 101 0.22285714 0.32142857 0.6516129 51.2628866 70.9793814 132.757732 P. profundus P004 180 170 156 57 39 67 0.31666667 0.22941176 0.42948718 89.1717791 61.0122699 104.815951 P. profundus P166 181 171 156 55 36 66 0.3038674 0.21052632 0.42307692 89.3312102 58.4713376 107.197452 P167 183 172 159 59 50 77 0.32240437 0.29069767 0.48427673 80.8870968 68.5483871 105.564516 P027 177 169 156 73 72 130 0.41242938 0.4260355 0.83333333 67.6909091 66.7636364 120.545455 P. profundus P148 210 204 192 48 35 85 0.22857143 0.17156863 0.44270833 72.8571429 53.125 129.017857 P. profundus P158 203 195 183 100 112 119 0.49261084 0.57435897 0.65027322 77.0392749 86.2839879 91.6767372 P. youngi Grey- Edge Elytrum Sample - Edge Elytrum Corrected RGB - Edge Elytrum Standardized - RGB Sample R G B R G B R G B R G B P120 211 203 192 79 66 91 0.37440758 0.32512315 0.47395833 85.3601695 71.3135593 98.3262712 P142 202 193 181 133 175 142 0.65841584 0.90673575 0.78453039 75.3666667 99.1666667 80.4666667 P002 201 190 178 67 158 154 0.33333333 0.83157895 0.86516854 45.0791557 106.306069 103.614776 P008 202 192 181 64 86 118 0.31683168 0.44791667 0.6519337 60.8955224 81.8283582 112.276119 P009 179 165 144 69 155 152 0.38547486 0.93939394 1.05555556 46.7952128 105.119681 103.085106 P010 194 183 169 103 71 89 0.53092784 0.38797814 0.52662722 99.8669202 68.8403042 86.2927757 P011 188 177 161 68 94 88 0.36170213 0.53107345 0.54658385 69.36 95.88 89.76 P012 204 191 175 121 78 98 0.59313725 0.40837696 0.56 103.888889 66.969697 84.1414141 P014 200 188 174 133 86 71 0.665 0.45744681 0.40804598 116.948276 75.6206897 62.4310345 P015 193 183 170 119 172 165 0.61658031 0.93989071 0.97058824 66.5460526 96.1842105 92.2697368 P016 204 192 178 133 76 81 0.65196078 0.39583333 0.45505618 116.948276 66.8275862 71.2241379 P017 208 197 183 111 151 179 0.53365385 0.76649746 0.97814208 64.1836735 87.3129252 103.503401 P018 208 197 183 98 161 185 0.47115385 0.81725888 1.01092896 56.2837838 92.4662162 106.25 P019 206 196 183 147 191 164 0.71359223 0.9744898 0.89617486 74.6713147 97.0219124 83.3067729 P020 209 198 184 142 74 61 0.67942584 0.37373737 0.33152174 130.722022 68.1227437 56.1552347 P021 199 186 169 55 88 95 0.27638191 0.47311828 0.56213018 58.9285714 94.2857143 101.785714 P022 204 192 178 70 146 216 0.34313725 0.76041667 1.21348315 41.3194444 86.1805556 127.5 22 P031 200 187 172 86 116 104 0.43 0.62032086 0.60465116 71.6666667 96.6666667 86.6666667 P032 204 193 179 129 160 152 0.63235294 0.82901554 0.84916201 74.5918367 92.5170068 87.8911565 P034 204 192 176 127 106 125 0.62254902 0.55208333 0.71022727 90.4608939 75.5027933 89.0363128 P040 202 194 184 70 192 193 0.34653465 0.98969072 1.04891304 39.2307692 107.604396 108.164835 P049 209 200 188 43 110 126 0.20574163 0.55 0.67021277 39.3010753 100.537634 115.16129 P050 203 193 181 107 174 168 0.5270936 0.9015544 0.9281768 60.7683742 98.8195991 95.4120267 P051 203 193 181 96 191 178 0.4729064 0.98963731 0.98342541 52.6451613 104.741935 97.6129032 P053 202 191 179 70 149 145 0.34653465 0.78010471 0.81005587 49.0384615 104.381868 101.57967 P054 204 194 182 99 60 80 0.48529412 0.30927835 0.43956044 105.627615 64.0167364 85.3556485 P055 201 195 182 138 191 164 0.68656716 0.97948718 0.9010989 71.3793103 98.7931034 84.8275862 P056 201 190 178 125 62 89 0.62189055 0.32631579 0.5 115.48913 57.2826087 82.2282609 P057 202 193 182 78 101 132 0.38613861 0.52331606 0.72527473 63.9549839 82.8135048 108.231511 P058 208 198 186 112 68 95 0.53846154 0.34343434 0.51075269 103.854545 63.0545455 88.0909091 P059 210 201 190 120 128 170 0.57142857 0.63681592 0.89473684 73.2057416 78.0861244 103.708134 P060 212 202 192 90 67 80 0.4245283 0.33168317 0.41666667 96.835443 72.0886076 86.0759494 P061 208 202 191 136 108 124 0.65384615 0.53465347 0.64921466 94.2391304 74.8369565 85.923913 P062 217 208 198 147 99 121 0.67741935 0.47596154 0.61111111 102.138965 68.7874659 84.0735695 P064 219 210 201 108 129 193 0.49315068 0.61428571 0.960199 64.0465116 76.5 114.453488 P071 215 207 197 184 115 123 0.85581395 0.55555556 0.62436548 111.184834 69.4905213 74.3246445 P074 211 203 192 108 81 122 0.51184834 0.39901478 0.63541667 88.5530547 66.414791 100.032154 P075 207 202 192 125 71 73 0.60386473 0.35148515 0.38020833 118.494424 67.3048327 69.2007435 P078 207 198 188 92 123 185 0.44444444 0.62121212 0.98404255 58.65 78.4125 117.9375 P079 212 204 195 127 177 191 0.5990566 0.86764706 0.97948718 65.4242424 91.1818182 98.3939394 P080 210 201 192 88 143 201 0.41904762 0.71144279 1.046875 51.9444444 84.4097222 118.645833 P081 202 193 178 116 80 107 0.57425743 0.41450777 0.6011236 97.6237624 67.3267327 90.049505 P082 207 200 187 104 142 211 0.50241546 0.71 1.12834225 58.0306346 79.2341357 117.73523 P083 210 200 189 133 145 185 0.63333333 0.725 0.97883598 73.25054 79.8596112 101.889849 P086 212 202 191 177 134 176 0.83490566 0.66336634 0.92146597 92.6796715 70.164271 92.1560575 P087 P088 194 185 173 84 99 181 0.43298969 0.53513514 1.04624277 58.8461538 69.3543956 126.799451 P089 195 179 166 101 93 162 0.51794872 0.51955307 0.97590361 72.3455056 66.6151685 116.039326 23 P091 193 184 171 126 84 135 0.65284974 0.45652174 0.78947368 93.1304348 62.0869565 99.7826087 P093 194 181 167 106 69 125 0.54639175 0.38121547 0.74850299 90.1 58.65 106.25 P094 198 188 175 53 117 140 0.26767677 0.62234043 0.8 43.5967742 96.2419355 115.16129 P095 199 185 175 66 133 171 0.33165829 0.71891892 0.97714286 45.4864865 91.6621622 117.851351 P096 190 180 167 106 78 108 0.55789474 0.43333333 0.64670659 92.5684932 68.1164384 94.3150685 P097 190 176 149 59 81 114 0.31052632 0.46022727 0.76510067 59.2322835 81.3188976 114.448819 P098 179 168 150 57 84 133 0.31843575 0.5 0.88666667 53.0474453 78.1751825 123.777372 P099 201 191 178 100 118 180 0.49751244 0.61780105 1.01123596 64.0703518 75.6030151 115.326633 P100 175 165 154 18 136 165 0.10285714 0.82424242 1.07142857 14.3887147 108.714734 131.896552 P101 196 187 176 81 158 139 0.41326531 0.84491979 0.78977273 54.6428571 106.587302 93.7698413 P102 193 183 172 81 165 152 0.41968912 0.90163934 0.88372093 51.8969849 105.71608 97.3869347 P103 185 172 157 64 103 91 0.34594595 0.59883721 0.57961783 63.255814 101.802326 89.9418605 P104 191 180 167 53 100 104 0.27748691 0.55555556 0.62275449 52.5875486 99.2217899 103.190661 P105 191 180 168 57 112 124 0.29842932 0.62222222 0.73809524 49.6075085 97.4744027 107.918089 P114 185 173 161 68 84 85 0.36756757 0.48554913 0.52795031 73.164557 90.3797468 91.4556962 P116 185 174 162 80 42 57 0.43243243 0.24137931 0.35185185 113.96648 59.8324022 81.2011173 P119 183 172 156 111 78 97 0.60655738 0.45348837 0.62179487 98.9685315 69.5454545 86.486014 P121 194 183 171 111 152 153 0.57216495 0.83060109 0.89473684 68.0408654 93.1730769 93.7860577 P122 185 175 163 104 81 117 0.56216216 0.46285714 0.71779141 87.8145695 68.3940397 98.7913907 P123 P124 179 170 157 74 103 143 0.41340782 0.60588235 0.91082803 58.96875 82.078125 113.953125 P125 179 171 157 72 104 144 0.40223464 0.60818713 0.91719745 57.375 82.875 114.75 P127 178 167 155 109 156 146 0.61235955 0.93413174 0.94193548 67.6277372 96.7883212 90.5839416 P128 180 172 158 90 122 120 0.5 0.70930233 0.75949367 69.126506 93.7048193 92.1686747 P129 197 187 175 74 126 144 0.37563452 0.67379679 0.82285714 54.8546512 93.4011628 106.744186 P130 199 187 175 123 140 178 0.61809045 0.7486631 1.01714286 71.122449 80.952381 102.92517 P134 182 171 158 118 84 128 0.64835165 0.49122807 0.81012658 91.1818182 64.9090909 98.9090909 P135 194 184 173 114 167 182 0.58762887 0.9076087 1.05202312 62.7861771 91.9762419 100.237581 P136 195 186 174 112 181 195 0.57435897 0.97311828 1.12068966 58.5245902 94.579918 101.895492 P137 201 192 179 9 142 167 0.04477612 0.73958333 0.93296089 7.21698113 113.867925 133.915094 P138 199 188 176 44 117 158 0.22110553 0.62234043 0.89772727 35.1724138 93.5266458 126.30094 24 P139 210 200 186 88 143 173 0.41904762 0.715 0.93010753 55.5445545 90.259901 109.195545 P140 200 189 177 81 163 165 0.405 0.86243386 0.93220339 50.5012225 101.625917 102.872861 P141 201 189 174 101 87 101 0.50248756 0.46031746 0.58045977 89.1176471 76.7647059 89.1176471 P143 214 207 198 111 133 173 0.51869159 0.64251208 0.87373737 67.8776978 81.3309353 105.791367 P144 199 188 175 47 106 141 0.2361809 0.56382979 0.80571429 40.7653061 91.9387755 122.295918 P145 203 196 184 103 99 149 0.50738916 0.50510204 0.80978261 74.8290598 71.9230769 108.247863 P147 205 194 182 99 154 149 0.48292683 0.79381443 0.81868132 62.7985075 97.6865672 94.5149254 P149 209 203 194 99 108 191 0.47368421 0.5320197 0.98453608 63.4296482 69.1959799 122.374372 P152 207 201 191 114 142 170 0.55072464 0.70646766 0.89005236 68.2394366 85 101.760563 P154 180 169 156 30 68 117 0.16666667 0.40236686 0.75 35.5813953 80.6511628 138.767442 P155 185 174 161 97 55 78 0.52432432 0.31609195 0.48447205 107.543478 60.9782609 86.4782609 P026 173 166 154 75 100 147 0.43352601 0.60240964 0.95454545 59.3944099 79.1925466 116.413043 P025 205 199 187 127 150 189 0.6195122 0.75376884 1.01069519 69.4957082 82.0815451 103.422747 P159 181 173 163 112 82 109 0.61878453 0.47398844 0.66871166 94.2574257 69.009901 91.7326733 P107 185 174 161 105 84 96 0.56756757 0.48275862 0.59627329 93.9473684 75.1578947 85.8947368 P157 172 163 148 91 127 151 0.52906977 0.7791411 1.02027027 62.8861789 87.7642276 104.349593 P005 175 168 155 53 91 123 0.30285714 0.54166667 0.79354839 50.6179775 86.9101124 117.47191 P004 180 170 156 79 98 158 0.43888889 0.57647059 1.01282051 60.1343284 74.5970149 120.268657 P166 181 171 156 46 56 107 0.25414365 0.32748538 0.68589744 56.1244019 68.3253589 130.550239 P167 183 172 159 99 75 102 0.54098361 0.43604651 0.64150943 91.4673913 69.2934783 94.2391304 P027 177 169 156 117 95 127 0.66101695 0.56213018 0.81410256 88.0088496 71.460177 95.5309735 P148 210 204 192 106 94 177 0.5047619 0.46078431 0.921875 71.6976127 63.5809019 119.721485 P158 203 195 183 121 143 153 0.59605911 0.73333333 0.83606557 73.9928058 87.4460432 93.5611511 Grey - Anterior Sample - Anterior Corrected RGB - Anterior Standardized RGB - Anterior Sample R G B R G B R G B R G B P120 211 203 192 132 111 143 0.62559242 0.54679803 0.74479167 87.2020725 73.3290155 94.4689119 P142 202 193 181 142 168 160 0.7029703 0.87046632 0.8839779 77.0425532 91.1489362 86.8085106 P002 201 190 178 60 124 140 0.29850746 0.65263158 0.78651685 47.2222222 97.5925926 110.185185 P008 202 192 181 86 110 137 0.42574257 0.57291667 0.75690608 65.8558559 84.2342342 104.90991 P009 179 165 144 51 119 110 0.2849162 0.72121212 0.76388889 46.4464286 108.375 100.178571 25 P010 194 183 169 97 126 135 0.5 0.68852459 0.79881657 69.0921788 89.7486034 96.1592179 P011 188 177 161 91 123 135 0.48404255 0.69491525 0.83850932 66.4899713 89.8710602 98.6389685 P012 204 191 175 101 104 132 0.49509804 0.54450262 0.75428571 76.4243323 78.694362 99.8813056 P014 200 188 174 74 108 115 0.37 0.57446809 0.66091954 63.5353535 92.7272727 98.7373737 P015 193 183 170 103 156 160 0.53367876 0.85245902 0.94117647 62.6849642 94.9403341 97.3747017 P016 204 192 178 90 146 147 0.44117647 0.76041667 0.8258427 59.921671 97.2062663 97.8720627 P017 208 197 183 119 143 176 0.57211538 0.72588832 0.96174863 69.2808219 83.2534247 102.465753 P018 208 197 183 85 128 161 0.40865385 0.64974619 0.87978142 57.9545455 87.2727273 109.772727 P019 206 196 183 157 205 193 0.76213592 1.04591837 1.05464481 72.1351351 94.1891892 88.6756757 P020 209 198 184 100 93 98 0.4784689 0.46969697 0.5326087 87.628866 81.4948454 85.8762887 P021 199 186 169 87 134 133 0.43718593 0.72043011 0.78698225 62.6694915 96.5254237 95.8050847 P022 204 192 178 95 89 122 0.46568627 0.46354167 0.68539326 79.1666667 74.1666667 101.666667 P031 200 187 172 87 110 114 0.435 0.58823529 0.6627907 71.3344051 90.192926 93.4726688 P032 204 193 179 76 122 120 0.37254902 0.63212435 0.67039106 60.9433962 97.8301887 96.2264151 P034 204 192 176 126 105 122 0.61764706 0.546875 0.69318182 91.01983 75.8498584 88.1303116 P040 202 194 184 107 157 164 0.52970297 0.80927835 0.89130435 63.75 93.5397196 97.7102804 P049 209 200 188 130 165 157 0.62200957 0.825 0.83510638 73.340708 93.0862832 88.5730088 P050 203 193 181 58 99 117 0.28571429 0.51295337 0.64640884 53.9781022 92.1350365 108.886861 P051 203 193 181 103 181 185 0.50738916 0.93782383 1.02209945 56.0021322 98.4115139 100.586354 P053 202 191 179 117 176 183 0.57920792 0.92146597 1.02234637 62.6785714 94.2857143 98.0357143 P054 204 194 182 74 126 149 0.3627451 0.64948454 0.81868132 54.0687679 92.0630372 108.868195 P055 201 195 182 100 126 141 0.49751244 0.64615385 0.77472527 69.4822888 87.5476839 97.9700272 P056 201 190 178 95 101 132 0.47263682 0.53157895 0.74157303 73.8567073 78.5213415 102.621951 P057 202 193 182 100 89 109 0.4950495 0.4611399 0.5989011 85.5704698 76.1577181 93.2718121 P058 208 198 186 112 93 119 0.53846154 0.46969697 0.63978495 88.1481481 73.1944444 93.6574074 P059 210 201 190 121 97 128 0.57619048 0.48258706 0.67368421 89.1763006 71.4884393 94.3352601 P060 212 202 192 147 120 137 0.69339623 0.59405941 0.71354167 92.7846535 75.7425743 86.4727723 P061 208 202 191 96 99 112 0.46153846 0.49009901 0.58638743 79.7394137 82.2312704 93.029316 P062 217 208 198 110 176 183 0.50691244 0.84615385 0.92424242 59.8081023 95.6929638 99.4989339 P064 219 210 201 129 117 178 0.5890411 0.55714286 0.88557214 77.5825472 70.365566 107.051887 P071 215 207 197 120 102 118 0.55813953 0.49275362 0.59898477 90 76.5 88.5 26 P074 211 203 192 152 155 201 0.72037915 0.7635468 1.046875 76.2992126 77.8051181 100.895669 P075 207 202 192 114 107 120 0.55072464 0.52970297 0.625 85.2492669 80.0146628 89.7360704 P078 207 198 188 129 106 119 0.62318841 0.53535354 0.63297872 92.9237288 76.3559322 85.720339 P079 212 204 195 128 107 132 0.60377358 0.5245098 0.67692308 88.9373297 74.346049 91.7166213 P080 210 201 192 151 97 157 0.71904762 0.48258706 0.81770833 95.0740741 61.0740741 98.8518519 P081 202 193 178 138 117 142 0.68316832 0.60621762 0.79775281 88.6397985 75.1511335 91.209068 P082 207 200 187 144 122 149 0.69565217 0.61 0.79679144 88.4819277 74.9638554 91.5542169 P083 210 200 189 77 55 98 0.36666667 0.275 0.51851852 85.3695652 60.9782609 108.652174 P086 212 202 191 146 117 159 0.68867925 0.5792 0792 0.83246073 88.2227488 70.6990521 96.0781991 P087 P088 194 185 173 132 96 121 0.68041237 0.51891892 0.69942197 96.4469914 70.1432665 88.4097421 P089 195 179 166 88 74 111 0.45128205 0.41340782 0.6686747 82.1978022 69.1208791 103.681319 P091 193 184 171 115 85 105 0.59585492 0.46195652 0.61403509 96.147541 71.0655738 87.7868852 P093 194 181 167 139 107 125 0.71649485 0.59116022 0.74850299 95.5390836 73.5444744 85.916442 P094 198 188 175 115 94 146 0.58080808 0.5 0.83428571 82.6056338 67.5211268 104.873239 P095 199 185 175 112 100 132 0.56281407 0.54054054 0.75428571 83.0232558 74.127907 97.8488372 P096 190 180 167 135 104 117 0.71052632 0.57777778 0.7005988 96.6994382 74.494382 83.8061798 P097 190 176 149 112 97 150 0.58947368 0.55113636 1.00671141 79.5543175 68.8997214 106.545961 P098 179 168 150 163 140 182 0.91061453 0.83333333 1.21333333 85.7010309 73.6082474 95.6907216 P100 175 165 154 107 142 165 0.61142857 0.86060606 1.07142857 65.9057971 87.4637681 101.630435 P101 196 187 176 133 191 175 0.67857143 1.02139037 0.99431818 67.9659319 97.6052104 89.4288577 P102 193 183 172 26 118 125 0.13471503 0.64480874 0.72674419 24.6468401 111.858736 118.494424 P103 185 172 157 66 117 126 0.35675676 0.68023256 0.80254777 54.4660194 96.5533981 103.980583 P104 191 180 167 78 148 168 0.40837696 0.82222222 1.00598802 50.4822335 95.786802 108.730964 P105 191 180 168 90 132 104 0.47120419 0.73333333 0.61904762 70.398773 103.251534 81.3496933 P114 185 173 161 84 92 119 0.45405405 0.53179191 0.73913043 72.6101695 79.5254237 102.864407 P116 185 174 162 108 91 114 0.58378378 0.52298851 0.7037037 87.9872204 74.1373802 92.8753994 P119 183 172 156 106 132 149 0.57923497 0.76744186 0.95512821 69.8449612 86.9767442 98.1782946 P121 194 183 171 126 131 160 0.64948454 0.71584699 0.93567251 77.0503597 80.1079137 97.8417266 P122 185 175 163 118 108 130 0.63783784 0.6171 4286 0.79754601 84.5224719 77.3595506 93.1179775 P123 27 P124 P125 P127 178 167 155 86 141 161 0.48314607 0.84431138 1.03870968 56.5206186 92.6675258 105.811856 P128 180 172 158 107 144 177 0.59444444 0.837 2093 1.12025316 63.75 85.7943925 105.455607 P129 P130 199 187 175 134 117 136 0.67336683 0.62566845 0.77714286 88.2945736 77.0930233 89.6124031 P134 182 171 158 124 111 134 0.68131868 0.64912281 0.84810127 85.6910569 76.7073171 92.601626 P135 194 184 173 120 162 191 0.6185567 0.88043478 1.10404624 64.6934461 87.3361522 102.970402 P136 195 186 174 112 120 145 0.57435897 0.64516129 0.83333333 75.7559682 81.1671088 98.0769231 P137 201 192 179 134 124 192 0.66666667 0.64583333 1.0726257 75.9333333 70.2666667 108.8 P138 199 188 176 132 127 172 0.66331658 0.67553191 0.97727273 78.0974478 75.1392111 101.763341 P139 210 200 186 133 104 145 0.63333333 0.52 0.77956989 88.7827225 69.4240838 96.7931937 P140 200 189 177 114 157 203 0.57 0.83068783 1.14689266 61.3291139 84.4620253 109.208861 P141 201 189 174 128 134 176 0.63681592 0.70899471 1.01149425 74.5205479 78.0136986 102.465753 P143 214 207 198 145 143 180 0.67757009 0.69082126 0.90909091 79.0064103 77.9166667 98.0769231 P144 199 188 175 132 158 195 0.66331658 0.84042553 1.11428571 69.4020619 83.0721649 102.525773 P145 203 196 184 114 104 164 0.56157635 0.53061224 0.89130435 76.0994764 69.4240838 109.47644 P147 205 194 182 132 190 204 0.64390244 0.97938144 1.12087912 63.9923954 92.1102662 98.8973384 P149 209 203 194 161 133 187 0.77033493 0.65517241 0.96391753 85.3534304 70.5093555 99.1372141 P152 207 201 191 127 146 198 0.61352657 0.72636816 1.03664921 68.7579618 79.044586 107.197452 P154 180 169 156 80 88 144 0.44444444 0.52071006 0.92307692 65.3846154 71.9230769 117.692308 P155 185 174 161 105 70 88 0.56756757 0.40229885 0.54658385 101.806084 67.8707224 85.3231939 P026 173 166 154 102 98 148 0.58959538 0.59036145 0.96103896 74.7413793 71.8103448 108.448276 P025 205 199 187 133 127 166 0.64878049 0.63819095 0.88770053 79.6126761 76.0211268 99.3661972 P159 181 173 163 123 159 185 0.67955801 0.91907514 1.13496933 67.1627409 86.8201285 101.017131 P107 185 174 161 99 110 127 0.53513514 0.63218391 0.78881988 75.1339286 83.4821429 96.3839286 P157 172 163 148 151 145 189 0.87790698 0.88957055 1.27702703 79.3917526 76.2371134 99.371134 P005 175 168 155 104 109 165 0.59428571 0.64880952 1.06451613 70.1587302 73.531746 111.309524 P004 180 170 156 114 104 128 0.63333333 0.61176471 0.82051282 84.017341 76.6473988 94.3352601 P166 181 171 156 81 82 129 0.44751381 0.47953216 0.82692308 70.7363014 71.609589 112.65411 P167 183 172 159 127 109 143 0.69398907 0.63372093 0.89937107 85.4485488 73.3377309 96.2137203 28 P027 177 169 156 111 89 129 0.62711864 0.52662722 0.82692308 86.0334347 68.9817629 99.9848024 P148 210 204 192 128 110 142 0.60952381 0.53921569 0.73958333 85.8947368 73.8157895 95.2894737 P158 203 195 183 141 177 179 0.69458128 0.90769231 0.97814208 72.3440644 90.8148893 91.8410463 Grey - Head Sample - Head Corrected RGB - Head Standardized RGB - Head Sample R G B R G B R G B R G B P120 211 203 192 133 137 140 0.63033175 0.67487685 0.72916667 82.7195122 85.2073171 87.0731707 P142 202 193 181 129 151 128 0.63861386 0.78238342 0.70718232 80.625 94.375 80 P002 201 190 178 120 176 166 0.59701493 0.92631579 0.93258427 66.2337662 97.1428571 91.6233766 P008 202 192 181 85 130 141 0.42079208 0.67708333 0.77900552 60.8848315 93.1179775 100.997191 P009 179 165 144 74 122 104 0.41340782 0.73939394 0.72222222 62.9 103.7 88.4 P010 194 183 169 155 183 179 0.79896907 1 1.0591716 76.450677 90.2611219 88.2882012 P011 188 177 161 126 152 140 0.67021277 0.85875706 0.86956522 76.8660287 92.7272727 85.4066986 P012 204 191 175 94 123 137 0.46078431 0.64397906 0.78285714 67.7118644 88.6016949 98.6864407 P014 200 188 174 100 145 159 0.5 0.7712766 0.9137931 63.1188119 91.5222772 100.358911 P015 193 183 170 91 126 133 0.47150259 0.68852459 0.78235294 66.3 91.8 96.9 P016 204 192 178 82 102 129 0.40196078 0.53125 0.7247191 66.8051118 83.0990415 105.095847 P017 208 197 183 118 111 123 0.56730769 0.56345178 0.67213115 85.4829545 80.4119318 89.1051136 P018 208 197 183 164 194 157 0.78846154 0.98477157 0.8579235 81.2038835 96.0582524 77.7378641 P019 206 196 183 143 175 164 0.69417476 0.89285714 0.89617486 75.653527 92.5829876 86.7634855 P020 209 198 184 128 131 100 0.61244019 0.66161616 0.54347826 90.9192201 93.0501393 71.0306407 P021 199 186 169 90 109 112 0.45226131 0.58602151 0.66272189 73.7942122 89.3729904 91.8327974 P022 204 192 178 98 97 126 0.48039216 0.50520833 0.70786517 77.8504673 77.0560748 100.093458 P031 200 187 172 90 101 96 0.45 0.54010695 0.55813953 79.9651568 89.738676 85.2961672 P032 204 193 179 127 149 129 0.62254902 0.77202073 0.72067039 79.962963 93.8148148 81.2222222 P034 204 192 176 110 109 134 0.53921569 0.56770833 0.76136364 79.4617564 78.7393768 96.7988669 P040 202 194 184 124 153 134 0.61386139 0.78865979 0.72826087 76.9343066 94.9270073 83.1386861 P049 P050 203 193 181 110 134 137 0.54187192 0.69430052 0.75690608 73.6220472 89.6850394 91.6929134 P051 203 193 181 115 140 147 0.56650246 0.7253886 0.8121547 72.9477612 88.8059701 93.2462687 P053 202 191 179 152 170 149 0.75247525 0.89005236 0.83240223 82.2929936 92.0382166 80.6687898 29 P054 204 194 182 121 132 129 0.59313725 0.68041237 0.70879121 80.7722513 88.1151832 86.1125654 P055 201 195 182 118 132 131 0.58706468 0.67692308 0.71978022 78.976378 88.3464567 87.6771654 P056 201 190 178 90 99 110 0.44776119 0.52105263 0.61797753 76.7558528 84.4314381 93.812709 P057 202 193 182 91 96 109 0.45049505 0.49740933 0.5989011 78.3952703 82.7027027 93.902027 P058 208 198 186 118 119 141 0.56730769 0.6010101 0.75806452 79.6031746 80.2777778 95.1190476 P059 210 201 190 125 114 153 0.5952381 0.56716418 0.80526316 81.3137755 74.1581633 99.5280612 P060 212 202 192 169 149 164 0.79716981 0.73762376 0.85416667 89.4087137 78.8278008 86.7634855 P061 208 202 191 97 114 113 0.46634615 0.56435644 0.59162304 76.3425926 89.7222222 88.9351852 P062 217 208 198 112 136 166 0.51612903 0.65384615 0.83838384 68.9855072 83.7681159 102.246377 P064 219 210 201 149 138 187 0.6803653 0.65714286 0.93034826 80.1582278 74.2405063 100.601266 P071 215 207 197 93 102 105 0.43255814 0.49275362 0.53299492 79.05 86.7 89.25 P074 211 203 192 123 143 158 0.58293839 0.7044335 0.82291667 73.9740566 86.0023585 95.0235849 P075 207 202 192 110 124 135 0.53140097 0.61386139 0.703125 76.0162602 85.6910569 93.2926829 P078 207 198 188 130 124 158 0.62801932 0.62626263 0.84042553 80.461165 76.7475728 97.7912621 P079 212 204 195 142 127 155 0.66981132 0.62254902 0.79487179 85.4009434 76.379717 93.2193396 P080 210 201 192 162 128 153 0.77142857 0.63681592 0.796875 93.2505643 73.6794582 88.0699774 P081 202 193 178 132 125 132 0.65346535 0.64766839 0.74157303 86.529563 81.940874 86.529563 P082 207 200 187 151 140 164 0.7294686 0.7 0.87700535 84.6263736 78.4615385 91.9120879 P083 210 200 189 119 135 171 0.56666667 0.675 0.9047619 71.4 81 102.6 P086 212 202 191 153 140 162 0.72169811 0.69306931 0.84816754 85.7472527 78.4615385 90.7912088 P087 P088 194 185 173 128 118 152 0.65979381 0.63783784 0.87861272 82.0100503 75.6030151 97.3869347 P089 195 179 166 95 83 104 0.48717949 0.46368715 0.62650602 85.9042553 75.0531915 94.0425532 P091 193 184 171 86 91 123 0.44559585 0.49456522 0.71929825 73.1 77.35 104.55 P094 198 188 175 123 163 176 0.62121212 0.86702128 1.00571429 67.8896104 89.9675325 97.1428571 P095 P096 190 180 167 110 101 132 0.57894737 0.56111111 0.79041916 81.7784257 75.0874636 98.1341108 P097 190 176 149 96 82 110 0.50526316 0.46590909 0.73825503 85 72.6041667 97.3958333 P098 179 168 150 101 121 138 0.56424581 0.7202381 0.92 71.5416667 85.7083333 97.75 P099 201 191 178 134 125 163 0.66666667 0.65445026 0.91573034 80.971564 75.5331754 98.4952607 P100 175 165 154 94 113 107 0.53714286 0.68484848 0.69480519 76.3375796 91.7675159 86.8949045 30 P101 196 187 176 145 193 180 0.73979592 1.03208556 1.02272727 71.3803089 95.0096525 88.6100386 P102 193 183 172 124 165 131 0.64248705 0.90163934 0.76162791 75.2857143 100.178571 79.5357143 P103 185 172 157 98 129 120 0.52972973 0.75 0.76433121 72.0172911 94.7982709 88.184438 P104 P105 191 180 168 87 141 143 0.45549738 0.78333333 0.85119048 59.7978437 96.9137466 98.2884097 P114 185 173 161 151 166 194 0.81621622 0.95953757 1.20496894 75.3522505 82.8375734 96.8101761 P116 P119 P121 194 183 171 145 175 200 0.74742268 0.95628415 1.16959064 71.1057692 85.8173077 98.0769231 P122 P123 P124 179 170 157 78 100 98 0.43575419 0.58823529 0.62420382 72.0652174 92.3913043 90.5434783 P125 179 171 157 79 101 101 0.44134078 0.59064327 0.6433121 71.6903915 91.6548043 91.6548043 P127 178 167 155 132 177 182 0.74157303 1.05988024 1.17419355 68.5539715 91.9246436 94.5213849 P128 180 172 158 135 135 170 0.75 0.78488372 1.07594937 78.2386364 78.2386364 98.5227273 P129 197 187 175 135 148 130 0.68527919 0.79144385 0.74285714 83.3535109 91.3801453 80.2663438 P130 199 187 175 98 135 151 0.49246231 0.72192513 0.86285714 65.078125 89.6484375 100.273438 P134 182 171 158 123 142 165 0.67582418 0.83040936 1.0443038 72.9418605 84.2093023 97.8488372 P135 194 184 173 134 165 170 0.69072165 0.89673913 0.98265896 72.8571429 89.7121535 92.4307036 P136 195 186 174 136 160 161 0.6974359 0.86021505 0.92528736 75.8862144 89.2778993 89.8358862 P137 201 192 179 132 134 153 0.65671642 0.69791667 0.8547486 80.3341289 81.5513126 93.1145585 P138 199 188 176 121 160 160 0.6080402 0.85106383 0.90909091 69.9659864 92.5170068 92.5170068 P139 210 200 186 143 170 173 0.68095238 0.85 0.93010753 75.0308642 89.1975309 90.7716049 P140 200 189 177 142 177 177 0.71 0.93650794 1 73.0040323 90.9979839 90.9979839 P141 P143 214 207 198 128 166 175 0.59813084 0.80193237 0.88383838 69.5948827 90.2558635 95.1492537 P144 199 188 175 126 167 185 0.63316583 0.88829787 1.05714286 67.2175732 89.0899582 98.6924686 P145 203 196 184 112 121 162 0.55172414 0.61734694 0.88043478 72.3037975 78.1139241 104.582278 P147 205 194 182 132 161 161 0.64390244 0.82989691 0.88461538 74.1409692 90.4295154 90.4295154 P149 209 203 194 145 144 184 0.6937799 0.70935961 0.94845361 78.1712474 77.6321353 99.1966173 P152 207 201 191 144 165 164 0.69565217 0.82089552 0.85863874 77.6321353 88.9534884 88.4143763 31 P154 180 169 156 136 133 164 0.75555556 0.78698225 1.05128205 80.0923788 78.3256351 96.5819861 P155 P026 173 166 154 87 119 136 0.50289017 0.71686747 0.88311688 64.8684211 88.7280702 101.403509 P025 205 199 187 89 117 116 0.43414634 0.5879397 0.62032086 70.4813665 92.6552795 91.863354 P159 181 173 163 124 148 136 0.68508287 0.85549133 0.83435583 77.5 92.5 85 P107 185 174 161 112 133 133 0.60540541 0.76436782 0.82608696 75.5555556 89.7222222 89.7222222 P157 172 163 148 122 143 155 0.70930233 0.87730061 1.0472973 74.0714286 86.8214286 94.1071429 P005 175 168 155 105 106 157 0.6 0.63095238 1.01290323 72.7581522 73.451087 108.790761 P004 180 170 156 94 98 124 0.52222222 0.57647059 0.79487179 75.8544304 79.0822785 100.063291 P166 181 171 156 108 132 151 0.59668508 0.77192982 0.96794872 70.4347826 86.0869565 98.4782609 P167 P027 177 169 156 169 156 174 0.95480226 0.92307692 1.11538462 86.3627255 79.7194389 88.9178357 P148 210 204 192 124 114 155 0.59047619 0.55882353 0.80729167 80.4580153 73.9694656 100.572519 P158 203 195 183 134 141 128 0.66009852 0.72307692 0.69945355 84.7890819 89.2183623 80.9925558 Table 1. 32 ID # species county locality loc. lat long collectors ridge # P 001 F P. youngi Marion Ocala Nat. For., W of Lake 4 29.34821 -81.81608 Daniel Dye Mt.Dora Kerr P 002 M P. youngi Marion Ocala Nat. For., W of Lake 4 29.34821 -81.81608 Daniel Dye Mt. Dora Kerr P 003 F P. profundus Hernando Sunshine Grove Rd at 2 28.5808 -82.4998 Mansell Southern Sandy Dr Brooksville (S) P 004 P. profundus Citrus N Parkwood Ave, S of W 1 29.01604 -82.53441 Moler Southern Oakhill St. Brooksville (N) P 005 P. profundus Citrus N Parkwood Ave, S of W 1 29.01604 -82.53441 Moler Southern Oakhill St. Brooksville (N) P 006 P. profundus Citrus N Camae Point 3 29.00785 -82.5243 Moler Southern Brooksville (N) P 007 P. youngi Marion Ocala Nat. For., W of Lake 4 29.34821 -81.81608 Daniel Dye Mt. Dora Kerr P 008 M P. youngi Marion Ocala Nat. For FL Hwy 40, 12 29.1799 -81.7755 Moler Mt. Dora 0.5 mi W FR 11 P 009 M P. youngi Marion Ocala Nat. For FL Hwy 40, 12 29.1799 -81.7755 Moler Mt. Dora 0.5 mi W FR 11 P 010 M P. youngi Marion Ocala Nat. For. FR 11, 0.4 11 29.3213 -81.7736 Moler Mt. Dora mi S Co Rd 314 P 011 M P. youngi Marion Ocala Nat. For. FR 11, 0.4 11 29.3213 -81.7736 Moler Mt. Dora mi S Co Rd 314 P 012 M P. youngi Marion Ocala Nat. For. FR 11, 0.4 11 29.3213 -81.7736 Moler Mt. Dora mi S Co Rd 314 P 013 F P. youngi Marion Ocala Nat. For. FR 11, 0.4 11 29.3213 -81.7736 Moler Mt. Dora mi S Co Rd 314 P 014 M P. youngi Marion Ocala Nat. For. FL Hwy 12 29.1799 -81.7755 Moler Mt. Dora 40, 0.5 mi W FR 11 P 015 M P. youngi Putnam Ocala Nat. For. FR 66, 0.3 10 29.4342 -81.7325 Moler Mt. Dora mi E SR 19 P 016 M P. youngi Putnam Ocala Nat. For. FR 66, 0.3 10 29.4342 -81.7325 Moler Mt. Dora mi E SR 19 P 017 M P. youngi? Putnam Ocala Nat. For. FR 66, 0.3 10 29.4342 -81.7325 Moler Mt. Dora mi E SR 19 P 018 M P. youngi Putnam Ocala Nat. For. FR 66, 0.3 10 29.4342 -81.7325 Moler Mt. Dora mi E SR 19 P 019 M P. youngi Putnam Ocala Nat. For. FR 66, 0.3 10 29.4342 -81.7325 Moler Mt. Dora mi E SR 19 33 P 020 M P. youngi? Putnam Ocala Nat. For. FR 66, 0.3 10 29.4342 -81.7325 Moler Mt. Dora mi E SR 19 P 021 M P. youngi Putnam Ocala Nat. For. FR 66, 0.3 10 29.4342 -81.7325 Moler Mt. Dora mi E SR 19 P 022 M P. profundus Levy SR 24, 0.3 mi W Alachua 13 29.5097 -82.5616 Moler Northern Co line Brooksville P 023 M P. youngi? Putnam Lake Galilee Dr., 0.15 N 9 29.6264 -81.9514 Moler Trail FL Hwy 20 P 024 M P. youngi? Putnam Lake Galilee Dr., 0.15 N 9 29.6264 -81.9514 Moler Trail FL Hwy 20 P 025 P. Marion Hwy 25 7 29.04203 -81.97864 Moler profundus? P 026 P. profundus Marion Indian Lake State Forest 5 29.28685 -82.0661 Moler P 027 P. profundus Marion Indian Lake State Forest 5 29.28685 -82.0661 Moler P 028 P. Marion Silver Spring Shores, 6 29.09329 -82.00845 Moler profundus? Emerald Dr. P 029 P. Marion Silver Spring Shores, 6 29.09329 -82.00845 Moler profundus? Emerald Dr. P 030 P. Marion Silver Spring Shores, 6 29.09329 -82.00845 Moler profundus? Emerald Dr. P 031 M P. youngi Putnam Etoniah State Forest 14 29.7232 -81.8276 Moler Mt. Dora P 032 M P. youngi Putnam Etoniah State Forest 14 29.7232 -81.8276 Moler Mt. Dora P 033 F P. profundus Levy 6.6 mi SW Williston; E 16 29.3313 -82.5361 Moler Northern Levy St & SE141Cr Brooksville P 034 M P. profundus Levy 4.5 mi WSW Williston; NE 15 29.3597 -82.5139 Moler Northern 20 St, 0.6 mi W SR121 Brooksville P 035 F P. profundus Levy SR 24, 0.3 mi W Alachua 13 29.5097 -82.5616 Moler Northern Co line Brooksville P 036 P. profundus Levy SR 24, 0.3 mi W Alachua 13 29.5097 -82.5616 Moler Northern Co line Brooksville P 037 P. profundus Levy SR 24, 0.3 mi W Alachua 13 29.5097 -82.5616 Moler Northern Co line Brooksville P 038 P. profundus Levy SR 24, 0.3 mi W Alachua 13 29.5097 -82.5616 Moler Northern Co line Brooksville P 039 P. profundus Levy SR 24, 0.3 mi W Alachua 13 29.5097 -82.5616 Moler Northern Co line Brooksville P 040 M P. youngi Marion SR 42, 2 mi E SE 182 Ave 19 28.9791 -81.7704 Moler Mt. Dora Rd P 041 P. youngi? Lake SR 44 jct SR 46A 17 28.8509 -81.5303 Moler Mt. Dora? 34 P 042 P. youngi? Lake W Huff Rd, 2 mi N SR 44 18 28.855 -81.5267 Moler Mt. Dora? P 043 P. youngi? Lake W Huff Rd, 2 mi N SR 44 18 28.855 -81.5267 Moler Mt. Dora? P 044 P. youngi? Lake W Huff Rd, 2 mi N SR 44 18 28.855 -81.5267 Moler Mt. Dora? P 045 P. youngi Putnam Ocala Nat. For.(vial 1) 20 29.4906 -81.8102 Daniel Dye Mt. Dora P 046 P. youngi Marion Ocala Nat. For. (vial 2) 21 29.4879 -81.8606 Daniel Dye Mt. Dora P 047 P. youngi Putnam Ocala Nat. For. (vial 4) 23 29.4041 -81.7413 Daniel Dye Mt. Dora P 048 F P. youngi? Alachua Holden Park Rd., 0.2 mi 24 29.546 -82.058 Moler Mt. Dora NW Putnam Co line P 049 M P. youngi? Alachua Holden Park Rd., 0.2 mi 24 29.546 -82.058 Moler Mt. Dora NW Putnam Co line P 050 M P. youngi Putnam Baden Powell Rd at SR 20 25 29.6154 -81.978 Moler Mt. Dora P 051 M P. youngi Putnam Baden Powell Rd at SR 20 25 29.6154 -81.978 Moler Mt. Dora P 052 P. youngi Clay SR 21, 2.2 mi NE SR100 26 29.8099 -82.0091 Moler Trail P 053 M P. youngi Clay SR 21, 2.2 mi NE SR100 26 29.8099 -82.0091 Moler Trail P 054 M P. youngi Clay SR 21, 2.2 mi NE SR100 26 29.8099 -82.0091 Moler Trail P 055 M P. youngi Clay SR 21, 2.2 mi NE SR100 26 29.8099 -82.0091 Moler Trail P 056 M P. profundus Dixie US 19, 5.3 mi W Old Town 32 29.60855 -83.07015 Moler P 057 F P. profundus Dixie Co Rd 361, 2.1 mi S Co Rd 33 29.63539 -83.35489 Moler 358 (S. Jena) P 058 M P. profundus Dixie Co Rd 361, 2.1 mi S Co Rd 33 29.63539 -83.35489 Moler 358 (S. Jena) P 059 M P. profundus Dixie Co Rd 361, 2.1 mi S Co Rd 33 29.63539 -83.35489 Moler 358 (S. Jena) P 060 M P. profundus Dixie Co Rd 361, 2.1 mi S Co Rd 33 29.63539 -83.35489 Moler 358 (S. Jena) P 061 M P. Alachua Holden Park Rd, 0.25 mi 24 29.5462 -82.058 Moler Mt. Dora profundus? NW Putnam Co line P 062 M P. p. dubius Volusia De Leon Springs 30 29.1363 -81.3565 Moler Deland P 063 F P. p. dubius Volusia De Leon Springs 30 29.1363 -81.3565 Moler Deland P 064 M P. profundus Gilchrist SR 26, 3.0 mi W Alachua 34 29.6314 -82.7048 Moler Northern Co line Brooksville P 065 F P. profundus Gilchrist SR 26, 3.0 mi W Alachua 34 29.6314 -82.7048 Moler Northern Co line Brooksville P 066 M P. Clay Hwy 21 at Gas Line Rd 29.80734 -82.01186 Moler Trail profundus? 35 P 067 F P. profundus Madison CR 53, 0.25 mi N 38 30.2635 -83.2833 Moler Lafayette Co line P 068 F P. profundus Lafayette CR 53, 1.2 mi N US 27 37 30.1462 -83.2932 Moler P 069 F P. profundus Dixie CR 349, 0.15 mi SCR 351A 43 29.7064 -82.9846 Moler (NE 512 Ave) P 070 F P. profundus Dixie US 19, 5.3 mi W Old Town 29.60855 -83.07015 Moler P 071 M P. profundus Taylor Beach Rd. (CR361) 8.9 mi 40 29.7607 -83.4975 Moler N Steinhatchee P 072 F P. profundus Taylor Hampton Spring Rd, 0.4 42 30.0409 -83.7266 Moler mi NE Coney Island P 073 F P. profundus Taylor Hampton Spring Rd, 0.4 42 30.0409 -83.7266 Moler mi NE Coney Island P 074 M P. profundus Taylor FL Hwy 51, 4.9 mi S US 19 39 29.7186 -83.3607 Moler P 075 M P. profundus Taylor FL Hwy 51, 4.9 mi S US 19 39 29.7186 -83.3607 Moler P 076 F P. profundus Gilchrist CR 340, 2.15 mi E US 129 43 29.7955 -82.8311 Moler P 077 F P. profundus Gilchrist CR 340, 2.15 mi E US 129 43 29.7955 -82.8311 Moler P 078 M P. profundus Hamilton SR 6, ~1 mi W I-75 30.50637 -83.06778 P 079 M P. profundus Hamilton CR 141, ~4.8 mi S SR6 30.42931 -83.17017 P 080 M P. profundus Hamilton CR 141. ~4.8 mi S SR6 30.42931 -81.17017 P 081 M P. profundus Marion CR 326, ENE 58th Ave. 29.23983 -82.05167 P 082 M P. profundus Hamilton CR 141, ~ 4.8 mi S SR6 30.42931 -83.17017 P 083 M P. profundus Marion CR 326 ENE 58th ave. 29.23 983 -82.05167 P 084 Mycotrupes Marion CR 326 ENE 58th ave. Cotton Plant gaigei P 085 Geotrupes Alachua Fl. 7818 SW CR 346 egeriei P 086 M P. profundus Marion Fl. Co. Rd. 326 E of NE 29.2403 -82.051 P. Moler 58th Ave. P 087 M P. profundus Hamilton Fl. SR 6, 0.08 mi E SW CR 30.48575 -83.19907 P. Moler 143 P 088 M P. profundus Hamilton Fl. SR 6, 0.08 mi E SW CR 30.48575 -83.19907 P. Moler 144 P 089 M P. profundus Hamilton Fl. SR 6, 0.08 mi E SW CR 30.48575 -83.19907 P. Moler 145 P 090 F P. profundus Marion Fl. US 27, 1.3 mi N SR 42. 28.9987 -81.99824 P. Moler P 091 M P. profundus Marion Fl. US 41 0.5 mi S CR 464 29.23511 -82.44105 P. Moler 36 P 092 F P. profundus Marion Fl. US 41 0.5 mi S CR 464 29.08914 -82.45154 P. Moler at Rainbow Springs P 093 M P. profundus Madison I-10 E-bound rest area 30.36751 -83.24831 P. Moler P 094 M P. profundus Madison I-10 E-bound rest area 30.36751 -83.24831 P. Moler P 095 M P. profundus Madison I-10 E-bound rest area 30.36751 -83.24831 P. Moler P 096 M P. profundus Madison Twin Rivers State Forest 30.59969 -83.26375 P. Moler P 097 M P. profundus Madison Twin Rivers State Forest 30.59969 -83.26375 P. Moler P 098 M P. profundus Madison Twin Rivers State Forest 30.59969 -83.26375 P. Moler P 099 M P. profundus Madison Twin Rivers State Forest 30.59969 -83.26375 P. Moler P 100 M P. youngi Marion SE 112th Place 29.06292 -81.90894 P. Moler P 101 M P. youngi Marion SE 112th Place 29.06292 -81.90894 P. Moler P 102 M P. youngi Marion SE 112th Place 29.06292 -81.90894 P. Moler P 103 M P. youngi Marion SE 112th Place 29.06292 -81.90894 P. Moler P 104 M P. youngi Marion Ocala Nat. Forest, S 28.9766 -81.772 P. Moler border P 105 M P. youngi Marion Ocala Nat. Forest, S 28.9766 -81.772 P. Moler border P 106 M P. p. dubius Volusia US 17 0.13 mi S Spring 29.13355 -81.35167 P. Moler S Spring Garden Ranch Rd. Garden Ranch P 107 P. p. dubius Volusia US 17 0.13 mi S Spring 29.13355 -81.35167 P. Moler S Spring Garden Ranch Rd. Garden Ranch P 108 P. p. dubius Putnam Lake Cond Dr., 0.25 mi S 29.4885 -81.5867 P. Moler Cresent City US 17 P 109 F P. profundus Marion Fl. US 27 1.2 mi N jct SR 42.99697 -81.99745 P. Moler P 110 F P. profundus Marion Fl. US 27 1.2 mi N jct SR 42.99697 -81.99745 P. Moler P 111 F P. profundus Osceola US 192 at Co Rd. 500 A 28.22411 -81.18109 P. Moler P 112 F P. profundus Osceola US 192 at Co Rd. 500 A 28.22411 -81.18109 P. Moler P 113 F P. profundus Osceola US 192 at Co Rd. 500 A 28.22411 -81.18109 P. Moler P 114 M P. profundus Polk Tiger Creek Preserve 27.809 -81.4896 P. Moler Lake Wales P 115 F P. profundus Polk Tiger Creek Preserve 27.809 -81.4896 P. Moler Lake Wales P 116 M P. profundus Indian Roseland 27.83766 -80.48475 P. Moler Atlantic Coastal River P 117 F P. profundus Indian Roseland 27.83766 -80.48475 P. Moler Atlantic Coastal River 37 P 118 F P. profundus Indian Roseland 27.83766 -80.48475 P. Moler Atlantic Coastal River P 119 F P. profundus Indian Roseland 27.83766 -80.48475 P. Moler Atlantic Coastal River P 120 M P. profundus Indian Roseland 27.83766 -80.48475 P. Moler Atlantic Coastal River P 121 M P. profundus Indian Roseland 27.83766 -80.48475 P. Moler Atlantic Coastal River P 122 M P. profundus Indian Roseland 27.83766 -80.48475 P. Moler Atlantic Coastal River P 123 M P. profundus Highlands US 27, S side of Josephine 27.3719 -81.40122 P. Moler Lake Wales Ck. P 124 F P. profundus Highlands US 27, S side of Josephine 27.3719 -81.40122 P. Moler Lake Wales Ck. P 125 M P. profundus Polk Hwy 630 E of Frostproof 27.75978 -81.46325 P. Moler Lake Wales P 126 F P. profundus Polk Hwy 580 E of Haines City 28.12093 -81.55084 P. Moler Lake Wales P 127 M P. profundus Polk Hwy 580 E of Haines City 28.12093 -81.55084 P. Moler Lake Wales P 128 M P. profundus Polk Hwy 580 E of Haines City 28.12093 -81.55084 P. Moler Lake Wales P 129 M P. profundus Polk Hwy 580 E of Haines City 28.12093 -81.55084 P. Moler Lake Wales P 130 M P. profundus Polk Hwy 580 E of Haines City 28.12093 -81.55084 P. Moler Lake Wales P 131 F P. profundus Marion Fl. US 41 0.5 mi S CR 464 29.08914 -82.45154 P. Moler a t Rainbow Springs P 132 F P. profundus Marion 29.19319 -82.40241 P. Moler Cotton Plant P 133 F P. profundus Marion 29.19319 -82.40241 P. Moler Cotton Plant P 134 M P. profundus Marion 29.19319 -82.40241 P. Moler C otton Plant P 135 M P. profundus Lake SE of SR 44 jct SR 46A 17 28.8509 -81.5303 P. Moler P 136 M P. profundus Lake SE of SR 44 jct SR 46A 17 28.8509 -81.5303 P. Moler P 137 M P. profundus Lake NE jct SR44 and La Place 28.8516 -81.5656 P. Moler Ct. P 138 M P. profundus Lake NE jct SR44 and La Place 28.8516 -81.5656 P. Moler Ct. P 139 M P. profundus Lake NE jct SR44 and La Place 28.8516 -81.5656 P. Moler Ct. P 140 M P. profundus Lake NE jct SR44 and La Place 28.8516 -81.5656 P. Moler Ct. P 141 M P. profundus Lake NE jct SR44 and La Place 28.8516 -81.5656 P. Moler Ct. 38 P 142 M P. youngi Lake Powerline Rd. E of SR 19 29.051 -81.6341 P. Moler P143 M P. profundus Marion CR 326 ENE 58th Ave. 29.2406 -82.0516 P. Moler P144 M P. profundus Marion CR 326 ENE 58th Ave. 29.2406 -82.0516 P. Moler P145 M P. profundus Marion CR 326 ENE 58th Ave. 29.2406 -82.0516 P. Moler P146 M P. profundus Marion CR 326 ENE 58th Ave. 29.2406 -82.0516 P. Moler Not Tissued P146 P. youngi Marion Ocala NF SR40, 0.4 Mi W 29.1783 -81.777 P. Moler FR 11 P147 P. youngi Marion Ocala Nat. For. FR 11, 0.4 29.3219 -81.7727 P. Moler m i S Co Rd 314 148 F profundeus M arioN 149 M dubius 150 F p rofunus Marion 151 M 152 M 153 M 154 M 155 M 156 M 157 M 158 M 159 M P148 M P. profundus Citrus W. Basket Way at N Travis 28.9889 -82.4482 P. Moler Not tissued Dr. P149 M P. profundus Citrus W. Basket Way at N Travis 28.9889 -82.4482 P. Moler Dr. P150 F P. profundus Levy US Hwy 41, 1.4 mi N 29.2349 -82.441 P. Moler Marion Co. line P151 F P. profundus Marion FL Hwy 4 at SE 172 Terr. 29.1727 -81.866 P. Moler P152 M P. profundus Marion FL Hwy 4 at SE 172 Terr. 29.1727 -81.866 P. Moler P153 F P. profundus Highlands US 27, S side of Josephine 27.3719 -81.40122 P. Moler Lake Wales Ck. Table 2. 39 40