Mediterranean Islands Hosting Marginal and Peripheral Forest Tree Populations: The Case of Pinus brutia Ten. in Cyprus
Abstract
:1. Introduction
2. Material and Methods
2.1. The P. brutia Forest in Cyprus
2.2. Sampling Design
2.3. Assessment of Genetic Diversity and Structure
2.4. Assessment of Morphoanatomical Diversity and Structure
3. Results
3.1. Genetic Diversity and Subpopulations Structure of P. brutia in Cyprus
3.2. The Patterns of Morphoanatomical Diversity and Structure in P. brutia
4. Discussion
4.1. P. brutia in Cyprus as a Source of Peripheral and Marginalforest Genetic Resources
4.2. Patterns of Genetic Diversity of P. brutia in Cyprus
4.3. Patterns of Morphoanatomical Traits of P. brutia in Cyprus
5. Conclusions and Perspectives of Conservation Action
5.1. Inferences from the Study of a Mediterranean Oceanic Island’s Peripheral Tree Population
5.2. Future Conservation Actions
Supplementary Materials
Author Contributions
Funding
Acknowledgments
Conflicts of Interest
References
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Location (Abbr.) | Distribution Area ǂ (ha/km) | Elevation Range (m) | Sample Size | |||||
---|---|---|---|---|---|---|---|---|
Genetic Analysis | Morphoanatomical Analysis | |||||||
No. Trees | No. Mega-Gametophytes | No. Trees | No. Cones | No. Seeds | No. Needles | |||
Macheras (Mach) | 5583 ha | 400–1200 | 60 | 50 | 60 | 180 | 540 | 180 |
Limassol (Lim) | 7896 ha | 800–1200 | 60 | 50 | 60 | 180 | 540 | 180 |
Akamas (Aka) | 5743 ha | 400 | 25 | 150 * | 25 | 75 | 225 | 180 |
Adelphi (Ade) | 12,826 ha | 400–1200 | 60 | 50 | 60 | 180 | 540 | 180 |
Troodos (Tro) | 8843 ha | 400–1200 | 60 | 50 | 60 | 180 | 540 | 180 |
Pafos North (PaN) | 37,794 ha | 400–1200 | 60 | 172 | 60 | 180 | 540 | 180 |
Pafos South (PaS) | 22,365 ha | 400–1200 | 60 | 178 | 60 | 180 | 540 | 180 |
Pafos z.400 (PaZ.400) | 320 km | 400 | 60 | 100 | 60 | 180 | 540 | 180 |
Pafos z.800 (PaZ.800) | 438 km | 800 | 25 | 150 * | 60 | 180 | 540 | 180 |
Pafos z.1200 (PaZ.1200) | 35 km | 1200 | 60 | 100 | 60 | 180 | 540 | 180 |
Total | 580 | 1050 | 565 | 1695 | 5085 | 1695 |
Subpopulation | Genetic Parameters | ||||
---|---|---|---|---|---|
PPL | Na | Ne | I | HE | |
Mach | 60% | 1.800 | 1.435 | 0.331 | 0.203 |
Lim | 60% | 1.700 | 1.300 | 0.269 | 0.170 |
Aka | 70% | 2.000 | 1.442 | 0.371 | 0.217 |
Ade | 80% | 2.100 | 1.473 | 0.393 | 0.242 |
Tro | 60% | 1.800 | 1.430 | 0.335 | 0.214 |
PaN | 90% | 2.300 | 1.415 | 0.393 | 0.228 |
PaS | 80% | 2.100 | 1.415 | 0.375 | 0.220 |
PaZ.400 | 80% | 2.000 | 1.375 | 0.350 | 0.209 |
PaZ.800 | 80% | 2.200 | 1.464 | 0.418 | 0.244 |
PaZ.1200 | 90% | 2.200 | 1.315 | 0.327 | 0.186 |
Overall | 90% | 2.300 | 1.421 | 0.381 | 0.220 |
(a) | ||||||||
Source of Variation | All Sampled Subpopulations§ | Range-Wide Subpopulations† | ||||||
d.f. | Variance Components | Percentage Variation | Fixation Indicesǂ | d.f. | Variance Components | Percentage Variation | Fixation Indicesǂ | |
Among Groups | 5 | 0.137 | 12% | PhiRT = 0.118 *** | 3 | 0.005 | 0% | PhiRT = 0.004 n.s |
Among subpopulations within Groups | 4 | 0.013 | 1% | PhiPR = 0.012 *** | 3 | 0.011 | 1% | PhiPR = 0.010 * |
Within subpopulations within Groups | 1040 | 1.017 | 87% | PhiPT = 0.129 *** | 693 | 1.088 | 99% | PhiPT = 0.014 *** |
Total | 1049 | 1.167 | 100% | 693 | 1.104 | 100% | ||
(b) | ||||||||
Source of Variation | Altitudinal Subpopulations | Aspect Subpopulations | ||||||
d.f. | Variance Components | Percentage Variation | Fixation Indicesǂ | d.f. | Variance Components | Percentage Variation | Fixation Indicesǂ | |
Among subpopulations | 2 | 0.020 | 2% | 1 | 0.014 | 1% | ||
Within subpopulations | 347 | 1.087 | 98% | PhiPT = 0.018 ** | 348 | 1.118 | 99% | PhiPT = 0.012 ** |
Total | 349 | 1.107 | 100% | 349 | 1.132 | 100% |
(a) | ||||||||||||||||
Traits | Mach | Lim | Aka | Ade | Tro | |||||||||||
μ | SD | CV | μ | SD | CV | μ | SD | CV | μ | SD | CV | μ | SD | CV | ||
Clen (mm) | 69.47 | 8.50 | 0.12 | 68.85 | 7.80 | 0.11 | 63.27 | 7.16 | 0.11 | 68.23 | 9.61 | 0.14 | 67.39 | 8.38 | 0.12 | |
Cwid (mm) | 40.40 | 3.17 | 0.08 | 37.77 | 3.31 | 0.08 | 35.335 | 3.60 | 0.10 | 38.56 | 3.65 | 0.34 | 38.53 | 3.11 | 0.08 | |
Clen/Cwid | 1.72 | 0.18 | 0.10 | 1.77 | 0.12 | 0.06 | 1.79 | 0.13 | 0.07 | 1.77 | 0.17 | 0.09 | 1.75 | 0.17 | 0.09 | |
Slen (mm) | 8.15 | 0.61 | 0.07 | 7.76 | 0.57 | 0.07 | 7.33 | 0.55 | 0.07 | 8.09 | 0.72 | 0.09 | 7.88 | 0.57 | 0.06 | |
SWid (mm) | 5.01 | 0.45 | 0.09 | 4.63 | 0.43 | 0.09 | 4.44 | 0.35 | 0.08 | 4.88 | 0.46 | 0.09 | 4.71 | 0.39 | 0.08 | |
SWing (mm) | 18.18 | 1.90 | 0.10 | 17.02 | 2.17 | 0.12 | 15.97 | 2.52 | 0.16 | 17.69 | 2.29 | 0.13 | 16.98 | 1.86 | 0.11 | |
SLenWing (mm) | 26.33 | 4.80 | 0.08 | 24.78 | 2.45 | 0.09 | 23.30 | 2.74 | 0.12 | 25.78 | 2.68 | 0.10 | 24.86 | 2.12 | 0.08 | |
NShLen (mm) | 6.82 | 1.48 | 0.21 | 6.40 | 0.99 | 0.15 | 6.81 | 0.87 | 0.13 | 6.57 | 0.96 | 0.14 | 6.46 | 1.03 | 0.16 | |
NLen (mm) | 126.76 | 15.62 | 0.12 | 106.10 | 13.01 | 0.12 | 115.72 | 12.83 | 0.11 | 131.05 | 15.63 | 0.12 | 119.33 | 14.91 | 0.12 | |
NWid (mm) | 1.33 | 0.09 | 0.06 | 1.31 | 0.06 | 0.05 | 1.43 | 0.09 | 0.06 | 1.32 | 0.08 | 0.06 | 1.28 | 0.08 | 0.06 | |
NThic (mm) | 0.82 | 0.05 | 0.06 | 0.79 | 0.04 | 0.05 | 0.88 | 0.06 | 0.07 | 0.82 | 0.06 | 0.07 | 0.78 | 0.06 | 0.07 | |
NResIn | 3.73 | 0.89 | 0.23 | 3.43 | 0.98 | 0.28 | 4.23 | 0.48 | 0.11 | 3.64 | 0.79 | 0.21 | 3.34 | 0.88 | 0.26 | |
NResDo | 6.05 | 0.98 | 0.16 | 5.78 | 0.99 | 0.17 | 6.72 | 0.95 | 0.14 | 6.16 | 1.06 | 0.17 | 5.64 | 1.24 | 0.22 | |
NResTot | 9.78 | 1.48 | 0.15 | 9.21 | 1.69 | 0.18 | 10.95 | 1.10 | 0.10 | 9.80 | 1.60 | 0.16 | 8.99 | 1.85 | 0.20 | |
NStomDo | 8.98 | 1.17 | 0.13 | 9.00 | 0.83 | 0.09 | 8.92 | 0.87 | 0.10 | 8.90 | 1.07 | 0.12 | 9.06 | 1.21 | 0.13 | |
NStomIn | 3.75 | 0.76 | 0.21 | 3.87 | 0.65 | 0.16 | 3.49 | 0.74 | 0.21 | 3.66 | 0.60 | 0.16 | 3.73 | 0.68 | 0.18 | |
NStoRow | 12.74 | 1.63 | 0.12 | 12.71 | 1.64 | 0.13 | 12.41 | 1.26 | 0.10 | 12.56 | 1.39 | 0.11 | 12.78 | 1.57 | 0.12 | |
NSto/Row | 83.37 | 8.69 | 0.10 | 83.37 | 9.75 | 0.11 | 84.25 | 10.68 | 0.13 | 87.47 | 8.14 | 0.09 | 81.65 | 9.57 | 0.11 | |
Nstom | 13,471 | 2917 | 0.21 | 11,268 | 2472 | 0.21 | 12,062 | 2171.60 | 0.18 | 14411 | 2730.80 | 0.19 | 12,465 | 2654.40 | 0.21 | |
NTeh | 70.23 | 9.43 | 0.13 | 71.43 | 12.08 | 0.17 | 75.07 | 8.24 | 0.11 | 70.73 | 10.61 | 0.15 | 67.21 | 13.50 | 0.20 | |
Traits | PaN | PaS | PaZ.400 | PaZ.800 | PaZ.1200 | |||||||||||
μ | SD | CV | μ | SD | CV | μ | SD | CV | μ | SD | CV | μ | SD | CV | ||
CLen (mm) | 69.96 | 10.26 | 0.14 | 67.69 | 10.05 | 0.14 | 68.75 | 9.08 | 0.13 | 68.75 | 10.09 | 0.14 | 67.02 | 8.79 | 0.13 | |
CWid (mm) | 39.04 | 3.84 | 0.09 | 38.64 | 4.15 | 0.10 | 38.26 | 3.58 | 0.09 | 39.4 | 4.09 | 0.1 | 39.49 | 3.59 | 0.09 | |
CLen/CWid | 1.74 | 0.16 | 0.09 | 1.74 | 0.17 | 0.09 | 1.79 | 0.17 | 0.09 | 1.75 | 0.16 | 0.09 | 1.69 | 0.15 | 0.08 | |
SLen (mm) | 8.12 | 0.61 | 0.07 | 8.05 | 0.66 | 0.08 | 7.95 | 0.61 | 0.07 | 8.17 | 0.67 | 0.08 | 8.23 | 0.62 | 0.07 | |
SWid (mm) | 4.83 | 0.40 | 0.08 | 4.81 | 0.42 | 0.08 | 4.73 | 0.37 | 0.08 | 4.86 | 0.41 | 0.08 | 4.96 | 0.43 | 0.08 | |
SWing (mm) | 17.71 | 2.55 | 0.14 | 17.51 | 2.51 | 0.14 | 17.44 | 2.32 | 0.13 | 17.89 | 2.51 | 0.14 | 17.66 | 2.08 | 0.11 | |
SLenWing (mm) | 25.83 | 2.95 | 0.11 | 25.56 | 2.98 | 0.11 | 25.39 | 2.72 | 0.1 | 26.06 | 2.96 | 0.11 | 25.89 | 2.42 | 0.09 | |
NShLen (mm) | 6.54 | 1.07 | 0.16 | 6.33 | 0.89 | 0.14 | 6.11 | 0.77 | 0.12 | 6.39 | 0.91 | 0.14 | 7.04 | 1.1 | 0.16 | |
NLen (mm) | 128.47 | 19.12 | 0.15 | 114.50 | 14.14 | 0.12 | 121.7 | 21.62 | 0.17 | 127.4 | 15.86 | 0.12 | 116.49 | 15.55 | 0.13 | |
NWid (mm) | 1.27 | 0.09 | 0.07 | 1.28 | 0.07 | 0.05 | 1.29 | 0.09 | 0.07 | 1.25 | 0.08 | 0.06 | 1.29 | 0.07 | 0.05 | |
NThic (mm) | 0.78 | 0.05 | 0.06 | 0.77 | 0.05 | 0.06 | 0.79 | 0.06 | 0.07 | 0.77 | 0.05 | 0.06 | 0.78 | 0.05 | 0.06 | |
NResIn | 3.55 | 0.90 | 0.25 | 3.15 | 0.87 | 0.27 | 3.24 | 1.08 | 0.33 | 3.37 | 0.8 | 0.23 | 3.46 | 0.79 | 0.22 | |
NResDo | 5.84 | 1.20 | 0.20 | 5.21 | 1.21 | 0.23 | 5.21 | 1.55 | 0.3 | 5.58 | 1.16 | 0.2 | 5.8 | 1.11 | 0.19 | |
NResTot | 9.39 | 1.90 | 0.20 | 8.36 | 1.84 | 0.22 | 8.46 | 2.34 | 0.27 | 8.94 | 1.79 | 0.2 | 9.26 | 1.66 | 0.18 | |
NStomDo | 8.92 | 1.06 | 0.12 | 9.18 | 1.02 | 0.11 | 8.91 | 1.06 | 0.12 | 8.58 | 1.01 | 0.11 | 9.48 | 1.1 | 0.11 | |
NStomIn | 3.67 | 0.61 | 0.16 | 3.69 | 0.62 | 0.16 | 3.69 | 0.6 | 0.16 | 3.56 | 0.59 | 0.16 | 3.79 | 0.69 | 0.18 | |
NStoRow | 12.59 | 1.38 | 0.11 | 12.88 | 1.32 | 0.10 | 12.61 | 1.41 | 0.11 | 12.14 | 1.22 | 0.1 | 13.27 | 1.54 | 0.11 | |
NSto/Row | 84.44 | 7.33 | 0.87 | 86.64 | 8.31 | 0.09 | 85.81 | 8.32 | 0.09 | 85.09 | 8.07 | 0.09 | 85.88 | 8.82 | 0.1 | |
Nstom | 13,678 | 2837.13 | 0.20 | 12,777 | 2391.70 | 0.18 | 13,138 | 2854.60 | 0.21 | 13,163 | 2430.70 | 0.18 | 13,270.18 | 2676.90 | 0.2 | |
NTeh | 69.97 | 9.98 | 0.14 | 71.06 | 10.22 | 0.14 | 71.45 | 9.85 | 0.13 | 70.79 | 10.09 | 0.14 | 71.98 | 10.82 | 0.15 | |
(b) | ||||||||||||||||
CLen | CWid | CLen/CWid | SLen | SWid | SWing | SLenWing | ||||||||||
μ | 67.60 | 38.74 | 1.75 | 8.02 | 4.81 | 17.50 | 25.51 | |||||||||
SD | 0.22 | 0.09 | 0.004 | 0.01 | 0.01 | 0.05 | 0.06 | |||||||||
CV | 0.14 | 0.10 | 0.10 | 0.08 | 0.09 | 0.13 | 0.11 | |||||||||
NShLen | NLen | NWid | NThic | NResIn | NResDo | NResTot | NStomDo | NStomIn | NStoRow | NSto/Row | Nstom | NTeh | ||||
μ | 6.53 | 121.07 | 1.29 | 0.79 | 3.47 | 5.74 | 9.21 | 9.00 | 3.71 | 12.69 | 84.83 | 13026 | 70.74 | |||
SD | 1.06 | 17.83 | 0.09 | 0.06 | 0.91 | 1.23 | 1.88 | 1.08 | 0.66 | 1.48 | 8.83 | 2769.6 | 10.79 | |||
CV | 0.16 | 0.15 | 0.07 | 0.66 | 0.26 | 0.21 | 0.20 | 0.12 | 0.18 | 0.12 | 0.10 | 0.21 | 0.15 |
CLen | CWid | CLen/CWid | SLen | SWid | SWing | SLenWing | |||||||
Altitude zone | −0.79 n.s. | 0.13 ** | −0.24 ** | 0.20 ** | 0.23 ** | 0.04 n.s. | 0.09 ** | ||||||
NShLen | NLen | NWid | NThic | NResIn | NResDo | NResTot | NStomDo | NStomIn | NStoRow | NSto/Row | Nstom | NTeh | |
Altitude zone | 0.36 ** | −0.10 * | −0.03 n.s. | −0.05 n.s. | 0.09 * | 0.16 ** | 0.14 ** | 0.20 ** | 0.05 n.s. | 0.16 n.s. | −0.001 n.s. | 0.01 n.s. | 0.03 |
Variable | Axis 1 | Axis 2 | Axis 3 | Axis 4 | Axis 5 | Axis 6 | Axis 7 | Axis 8 | Axis 9 |
---|---|---|---|---|---|---|---|---|---|
% of variance explained | 40.6 | 27.9 | 14.0 | 7.2 | 4.4 | 3.0 | 2.0 | 0.6 | 0.2 |
NWid | −0.628 * | 0.277 | 0.216 | 0.191 | 0.199 | −0.126 | −0.185 | −0.273 | 0.136 |
NThic | −0.578 * | 0.557 | 0.159 | 0.103 | 0.296 | −0.074 | −0.268 | −0.126 | 0.086 |
SLen | 0.446 * | 0.145 | 0.281 | 0.149 | 0.185 | 0.166 | −0.055 | 0.033 | 0.188 |
NLen | 0.328 | 0.742 * | −0.273 | 0.023 | −0.052 | −0.181 | −0.033 | −0.214 | 0.258 |
NStoRow | 0.250 | 0.439 * | 0.011 | 0.409 | 0.060 | −0.154 | −0.408 | −0.269 | 0.224 |
NShLen | −0.047 | 0.171 | 0.579 * | 0.009 | −0.288 | −0.218 | 0.046 | −0.063 | 0.293 |
SWid | 0.348 | 0.259 | 0.475 * | 0.109 | 0.419 | 0.108 | −0.039 | 0.113 | −0.019 |
NStom | 0.068 | 0.060 | −0.028 | 0.639 * | 0.012 | 0.334 | −0.193 | 0.262 | 0.018 |
NResIn | −0.263 | 0.375 | 0.282 | −0.182 | −0.553 * | 0.128 | −0.287 | 0.011 | 0.001 |
NResTot | −0.296 | 0.415 | 0.278 | −0.168 | −0.530 * | 0.118 | −0.131 | −0.216 | −0.078 |
CWid | 0.337 | 0.136 | 0.325 | −0.218 | 0.432 * | 0.073 | 0.156 | −0.019 | −0.023 |
CLen | 0.153 | 0.076 | −0.040 | −0.093 | 0.363 * | 0.166 | −0.018 | −0.060 | 0.231 |
NStomDo | −0.019 | −0.159 | 0.394 | 0.246 | 0.087 | −0.452 * | −0.388 | −0.424 | −0.002 |
NTeh | −0.110 | 0.002 | 0.063 | 0.341 | −0.102 | 0.361 * | 0.354 | −0.277 | 0.263 |
SWing | 0.253 | 0.154 | 0.148 | 0.008 | 0.329 | 0.344 * | 0.079 | −0.122 | −0.167 |
SLenWing | 0.324 | 0.167 | 0.195 | 0.042 | 0.328 | 0.337 * | 0.055 | −0.098 | −0.099 |
NStomIn | −0.005 | −0.136 | 0.190 | −0.154 | 0.213 | 0.139 | −0.454 * | −0.411 | 0.376 |
NStomDo | 0.004 | −0.141 | 0.364 | 0.158 | 0.168 | −0.387 | −0.428 | −0.472 * | 0.134 |
NResDo | −0.250 | 0.343 | 0.189 | −0.097 | −0.339 | 0.069 | 0.117 | −0.454 * | −0.160 |
CLen/CWid | −0.128 | −0.022 | −0.380 | 0.083 | 0.105 | 0.159 | −0.136 | −0.007 | 0.424 * |
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Eliades, N.-G.H.; Aravanopoulos, F.A.; Christou, A.K. Mediterranean Islands Hosting Marginal and Peripheral Forest Tree Populations: The Case of Pinus brutia Ten. in Cyprus. Forests 2018, 9, 514. https://doi.org/10.3390/f9090514
Eliades N-GH, Aravanopoulos FA, Christou AK. Mediterranean Islands Hosting Marginal and Peripheral Forest Tree Populations: The Case of Pinus brutia Ten. in Cyprus. Forests. 2018; 9(9):514. https://doi.org/10.3390/f9090514
Chicago/Turabian StyleEliades, Nicolas-George H., Filippos (Phil) A. Aravanopoulos, and Andreas K. Christou. 2018. "Mediterranean Islands Hosting Marginal and Peripheral Forest Tree Populations: The Case of Pinus brutia Ten. in Cyprus" Forests 9, no. 9: 514. https://doi.org/10.3390/f9090514