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Ecological and morphological differentiation among COI haplotype groups in the plant parasitic nematode species Mesocriconema xenoplax

Julianne N. Matczyszyn
Julianne N. Matczyszyn
, 
Timothy Harris
Timothy Harris
, 
Kirsten Powers
Kirsten Powers
, 
Sydney E. Everhart
Sydney E. Everhart
 y 
Thomas O. Powers
Thomas O. Powers
   | 11 may 2022
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Article Category: Research Paper
Publicado en línea: 11 may 2022
Páginas: -
Recibido: 08 dic 2021
DOI: https://doi.org/10.2478/jofnem-2022-0009
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© 2022 Julianne N. Matczyszyn et al., published by Sciendo
This work is licensed under the Creative Commons Attribution 4.0 International License.

Figure 1

A condensed maximum-likelihood tree of 721-bp sequences isolated from the COI gene for each individual in the morphospecies Mesocriconema xenoplax. Generated in the program MEGA, the tree displays seven partitions resulting in the HG label for further analysis. Bootstrap values from neighbor-joining and maximum-likelihood trees and posterior probability from Bayesian trees showing support for each HG are given at the terminal node, neighbor-joining, maximum-likelihood, and Bayesian, respectively. Each analysis is indicated by shading in the legend. The absence of the bar indicates the absence of support. The spanning tree network of each HG constructed using statistical parsimony is given within each HG bracket. Larger circles represent a unique haplotype. Smaller circles represent the number of mutational steps between haplotypes. Networks are disconnected when they exceed the 90% connection limit. HG, haplotype group.
A condensed maximum-likelihood tree of 721-bp sequences isolated from the COI gene for each individual in the morphospecies Mesocriconema xenoplax. Generated in the program MEGA, the tree displays seven partitions resulting in the HG label for further analysis. Bootstrap values from neighbor-joining and maximum-likelihood trees and posterior probability from Bayesian trees showing support for each HG are given at the terminal node, neighbor-joining, maximum-likelihood, and Bayesian, respectively. Each analysis is indicated by shading in the legend. The absence of the bar indicates the absence of support. The spanning tree network of each HG constructed using statistical parsimony is given within each HG bracket. Larger circles represent a unique haplotype. Smaller circles represent the number of mutational steps between haplotypes. Networks are disconnected when they exceed the 90% connection limit. HG, haplotype group.

Figure 2

Distribution of Mesocriconema xenoplax HG per collection site within the United States. The color of the circle is associated with the HG number, and the size of the circle is associated with the number of individuals of each HG collected at each site. HG, haplotype group.
Distribution of Mesocriconema xenoplax HG per collection site within the United States. The color of the circle is associated with the HG number, and the size of the circle is associated with the number of individuals of each HG collected at each site. HG, haplotype group.

Figure 3

An ordination plot using dbRDA bi-plot showing the seven most influential explanatory environmental variables (arrows) overlain on the first two eigenvectors of the analysis of Mesocriconema xenoplax HG. The length of the arrows is directly proportional to the strength of the correlation between the explanatory variable and the genetic variation calculated between individuals within each HG. Circles represent HG in ordination space, with the corresponding HG number indicated in the respective circle. Each axis indicates the amount of total variation each respective axis accounts for. dbRDA, distance-based redundancy analysis; HG, haplotype group.
An ordination plot using dbRDA bi-plot showing the seven most influential explanatory environmental variables (arrows) overlain on the first two eigenvectors of the analysis of Mesocriconema xenoplax HG. The length of the arrows is directly proportional to the strength of the correlation between the explanatory variable and the genetic variation calculated between individuals within each HG. Circles represent HG in ordination space, with the corresponding HG number indicated in the respective circle. Each axis indicates the amount of total variation each respective axis accounts for. dbRDA, distance-based redundancy analysis; HG, haplotype group.

Figure 4

Scatterplots of discriminant analysis of principal components, calculated on the category of land cover that distinguishes the sequential data the most. Points represent observed individual Mesocriconema xenoplax nematodes identified at their respective land cover type, connected to the population centroids. The center of each component is represented as black grid lines. Land cover types with fewer than five individual nematodes were removed from the analysis to reduce the amount of noise.
Scatterplots of discriminant analysis of principal components, calculated on the category of land cover that distinguishes the sequential data the most. Points represent observed individual Mesocriconema xenoplax nematodes identified at their respective land cover type, connected to the population centroids. The center of each component is represented as black grid lines. Land cover types with fewer than five individual nematodes were removed from the analysis to reduce the amount of noise.

Figure 5A–E

Stacked bar plot of the probability of each Mesocriconema xenoplax nematode to be reassigned to its respective population, defined according to land cover type; A. deciduous forest, B. developed open space, C. woody wetlands, D. cultivated crops, and E. herbaceous. Only those land cover types with ≥5 individuals were included in the analysis. The vertical axis is the probability (y-axis) of each individual (x-axis) being reassigned to its respective population, and the width of the bars depends on population size.
Stacked bar plot of the probability of each Mesocriconema xenoplax nematode to be reassigned to its respective population, defined according to land cover type; A. deciduous forest, B. developed open space, C. woody wetlands, D. cultivated crops, and E. herbaceous. Only those land cover types with ≥5 individuals were included in the analysis. The vertical axis is the probability (y-axis) of each individual (x-axis) being reassigned to its respective population, and the width of the bars depends on population size.

Figure 6

Canonical plot of the seven Mesocriconema xenoplax HG are indicated by color and significant morphological features: knob width (knob), total body length (L), and stylet length (Sty). Rays are scaled in the canonical plots to 1.5. The larger ellipses denote 50% of the observations, assuming normality, respective to size, for the mean canonical variables. The smaller ellipses represent a 95% confidence ellipsoid for the means on the canonical variable. HG, haplotype group.
Canonical plot of the seven Mesocriconema xenoplax HG are indicated by color and significant morphological features: knob width (knob), total body length (L), and stylet length (Sty). Rays are scaled in the canonical plots to 1.5. The larger ellipses denote 50% of the observations, assuming normality, respective to size, for the mean canonical variables. The smaller ellipses represent a 95% confidence ellipsoid for the means on the canonical variable. HG, haplotype group.

Summary of morphological statistics by HG of Mesocriconema xenoplax specimens.

HG 08 HG 09 HG 10 HG 11 HG 12 HG 13 HG 14
N 3 13 9 14 12 16 12
R 99 ± 2.6 94 ± 4.3 94 ± 3.9 101 ± 4.7 95 ± 3.4 98 ± 7.2 100 ± 3.9
Rv 7 ± 0.6 7 ± 0.9 6 ± 1.1 7 ± 0.9 7 ± 0.8 7 ± 1.2 7 ± 0.9
Rex 29 ± 2.1 27 ± 1.6 28 ± 1.9 27 ± 1.8 26 ± 1.6 27 ± 17 27 ± 1.9
Body annule width (µm) 6.4 ± 0.7 7.0 ± 1.0 7.3 ± 0.7 6.0 ± 0.7 6.8 ± 0.7 6.1 ± 0.6 7 ± 0.7
Length (µm) 598.2 ± 43.2 634.6 ± 83.2 621.6 ± 50.5 587.2 ± 63.0 604.8 ± 65.9 579.9 ± 49.3 642 ± 44.7
Stylet length (µm) 83.7 ± 2.1 87.6 ± 5.7 89.6 ± 3.8 72.6 ± 3.0 74.9 ± 3.6 73.3 ± 4.9 71 ± 1.4
Stylet knob width (µm) 13.3 ± 0.6 13.5 ± 0.9 13.3 ± 1.1 12.0 ± 0.7 12.3 ± 1.4 11.6 ± 1.3 14 ± 0.6
VUL 554.2 ± 41.3 594.2 ± 79.4 581.8 ± 52.3 549.7 ± 59.7 565.3 ± 61.0 539.4 ± 46.5 597 ± 40.1
MBW 52.7 ± 1.5 51.8 ± 4.8 55.8 ± 9.1 45.6 ± 6.7 45.6 ± 5.1 47.2 ± 5.6 52 ± 3.7
VBW 40.3 ± 2.5 38.0 ± 3.5 40.3 ± 5.9 35.0 ± 4.2 35.3 ± 4.5 36.7 ± 4.5 40 ± 4.6
ESO 145.7 ± 1.2 159.1 ± 13.7 159.6 ± 6.0 136.2 ± 7.0 141.0 ± 6.6 142.4 ± 6.1 146 ± 8.9
V 92.6 ± 0.8 93.6 ± 1.0 93.5 ± 1.2 93.6 ± 1.2 93.5 ± 0.6 93.0 ± 0.7 93 ± 1.2

Locality and GenBank accession information for Mesocriconema xenoplax specimens in this study.

NID Locality Ecoregion Agricultural site (Y or N) GenBank Accession No.
N3 Timmas Farm Ecological Forest Reserve, Cass County, NE, USA Central Tall Grasslands N KJ787901
N173 Middle Loup River, Hooker County, NE, USA Nebraska Sand Hills Mixed Grasslands N KJ787902
N583 Wakulla Springs State Park, Wakulla County, FL, USA Southeastern Conifer Forests N KJ787880
N584 Ichetucknee Springs State Park, Columbia County, FL, USA Southeastern Conifer Forests N KJ787885
N588 Ichetucknee Springs State Park, Columbia County, FL, USA Southeastern Conifer Forests N KJ787886
N607 Schluckebier Prairie State Natural Area, Sauk County, WI, USA Upper Midwest forest-savanna Transition Zone N KJ787881
N724 Chimney Creek, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N KJ787906
N728 Pickens County, SC, USA Southeastern Mixed Forests Y KJ787873
N729 Pickens County, SC, USA Southeastern Mixed Forests Y KJ787874
N730 Pickens County, SC, USA Southeastern Mixed Forests Y KJ787875
N733 Ozark National Forest, AR, USA Central US Hardwood Forests N KJ787882
N735 Ozark National Forest, AR, USA Central US Hardwood Forests N KJ787883
N736 Southeastern Fruit and Nut Research Station, Peach County, GA, USA Southeastern Mixed Forests Y KJ787887
N746 Crawford Bay, British Columbia, Canada Cascade Mountains Leeward Forests N KJ787907
N747 Crawford Bay, British Columbia, Canada Cascade Mountains Leeward Forests N KJ787908
N944 Timmas Farm Ecological Forest Reserve, Cass County, NE, USA Central Tall Grasslands N KJ787903
N945 Timmas Farm Ecological Forest Reserve, Cass County, NE, USA Central Tall Grasslands N KJ787904
N947 Timmas Farm Ecological Forest Reserve, Cass County, NE, USA Central Tall Grasslands N KJ787905
N999 Albright Grove, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N KJ787896
N1024 Albright Grove, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N KJ787897
N1025 Albright Grove, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N KJ787898
N1028 Albright Grove, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N KJ787899
N1072 Nine-Mile Prairie, Lancaster County, NE, USA Central Tall Grasslands N KJ787913
N1073 Nine-Mile Prairie, Lancaster County, NE, USA Central Tall Grasslands N KJ787914
N1215 Long Branch Stream Valley Park, Fairfax County, VA, USA Southeastern Mixed Forests N KJ787900
N1216 Long Branch Stream Valley Park, Fairfax County, VA, USA Southeastern Mixed Forests N MN711175
N1217 Spring Green Prairie Preserve, Sauk County, WI, USA Upper Midwest forest-savanna Transition Zone N KJ787884
N1267 George Washington Memorial Parkway, Fairfax County, VA, USA Southeastern Mixed Forests N KJ787909
N1276 George Washington Memorial Parkway, Fairfax County, VA, USA Southeastern Mixed Forests N KJ787910
N1294 Musser Fruit Research Farm, Oconee County, SC, USA Southeastern Mixed Forests Y KJ787876
N1297 Musser Fruit Research Farm, Oconee County, SC, USA Southeastern Mixed Forests Y KJ787877
N1298 Musser Fruit Research Farm, Oconee County, SC, USA Southeastern Mixed Forests Y KJ787878
N1327 Fresno County, CA, USA California Central Valley Y KU236636
N1346 George Washington Memorial Parkway, Fairfax County, VA, USA Southeastern Mixed Forests N KY574831
N1361 George Washington Memorial Parkway, Fairfax County, VA, USA Southeastern Mixed Forests N KY574643
N1368 Nine-Mile Prairie, Lancaster County, NE, USA Central tall grasslands N KJ787915
N1375 Nine-Mile Prairie, Lancaster County, NE, USA Central tall grasslands N KJ787916
N1397 George Washington Memorial Parkway, VA, USA Southeastern Mixed Forests N KY574644
N1451 George Washington Memorial Parkway, VA, USA Southeastern Mixed Forests N KJ788063
N2262 George Washington Memorial Parkway, VA, USA Southeastern Mixed Forests N KY574645
N2269 George Washington Memorial Parkway, VA, USA Southeastern Mixed Forests N MN711176
N2528 Nine-Mile Prairie, Lancaster County, NE, USA Central tall grasslands N KY574650
N2557 Autauga County, AL, USA Southeastern Mixed Forests N KY574633
N2558 Autauga County, AL, USA Southeastern Mixed Forests Y MN711177
N2577 Chilton County, AL, USA Southeastern Mixed Forests Y KY574624
N2604 Chilton County, AL, USA Southeastern Mixed Forests Y MN711178
N2610 Chilton County, AL, USA Southeastern Mixed Forests Y MN711179
N2611 Chilton County, AL, USA Southeastern Mixed Forests Y MN711180
N2615 Chilton County, AL, USA Southeastern Mixed Forests Y KY574634
N2618 Chilton County, AL, USA Southeastern Mixed Forests Y KY574635
N2619 Chilton County, AL, USA Southeastern Mixed Forests Y MN711181
N2622 Chilton County, AL, USA Southeastern Mixed Forests Y KY574623
N2694 Albright Grove, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MF770909
N2719 Albright Grove, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711182
N2720 Albright Grove, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711183
N2727 Purchase Knob, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711184
N2728 Purchase Knob, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711185
N2842 Double Springs Gap, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711186
N2844 Double Springs Gap, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711187
N2849 Konza Prairie Biological Station, Riley County, KS, USA Flint Hills Tall Grasslands N KY574651
N2850 Konza Prairie Biological Station, Riley County, KS, USA Flint Hills Tall Grasslands N KY574652
N2851 Konza Prairie Biological Station, Riley County, KS, USA Flint Hills Tall Grasslands N MN711188
N2853 Konza Prairie Biological Station, Riley County, KS, USA Flint Hills Tall Grasslands N MN711189
N2855 Double Springs Gap, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711190
N2857 Double Springs Gap, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711191
N2858 Double Springs Gap, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711192
N2863 Konza Prairie Biological Station, Riley County, KS, USA Flint Hills Tall Grasslands N KY574653
N2864 Konza Prairie Biological Station, Riley County, KS, USA Flint Hills Tall Grasslands N MN711193
N2869 Double Springs Gap, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N KY574646
N2873 Double Springs Gap, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711194
N2874 Double Springs Gap, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711195
N2891 Konza Prairie Biological Station, Riley County, KS, USA Flint Hills Tall Grasslands N KY574654
N2892 Konza Prairie Biological Station, Riley County, KS, USA Flint Hills Tall Grasslands N KY574655
N2893 Konza Prairie Biological Station, Riley County, KS, USA Flint Hills Tall Grasslands N MN711196
N2896 Konza Prairie Biological Station, Riley County, KS, USA Flint Hills Tall Grasslands N MN711197
N2933 West Point, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N KY574647
N2945 West Point, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711198
N2998 Cades Cove, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711199
N3008 Goshen Prong, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N KY574648
N3013 Trillium Gap, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711200
N3078 Gifford Woods State Park, VT, USA New England Acadian Forests N KY574639
N3084 Hayden Prairie State Preserve, Howard County, IA, USA Central tall grasslands N MN711201
N3112 Hayden Prairie State Preserve, Howard County, IA, USA Central tall grasslands N MN711202
N3124 Hayden Prairie State Preserve, Howard County, IA, USA Central tall grasslands N KY574656
N3153 George Washington Memorial Parkway, VA, USA Southeastern Mixed Forests N KY574649
N3154 George Washington Memorial Parkway, VA, USA Southeastern Mixed Forests N MN711203
N3208 Ichetucknee Springs State Park, FL, USA Southeastern Conifer forests N KY574625
N3246 Oconaluftee, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711204
N3253 Oconaluftee, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711205
N3316 Twin Creeks, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N KY574640
N3320 Twin Creeks, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N KY574832
N3342 Torreya State Park, FL, USA Southeastern Conifer forests N KY574636
N3361 Raspberry Island, Apostle Islands National Lakeshore, WI, USA Western Great Lakes Forest N KY574641
N3371 Raspberry Island, Apostle Islands National Lakeshore, WI, USA Western Great Lakes Forest N MN711206
N3372 Raspberry Island, Apostle Islands National Lakeshore, WI, USA Western Great Lakes Forest N MN711207
N3373 Raspberry Island, Apostle Islands National Lakeshore, WI, USA Western Great Lakes Forest N MN711208
N3374 Raspberry Island, Apostle Islands National Lakeshore, WI, USA Western Great Lakes Forest N MN711209
N3481 Wakulla Springs State Park, FL, USA Southeastern Conifer forests N MN711210
N3491 West Point, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MF770951
N3492 West Point, Great Smoky Mountains National Park, USA Appalachian Blue Ridge Forests N MN711211
N5508 Oconaluftee, Great Smoky Mountains National Park, USA Appalachian-Blue Ridge Forests N MN711216
N5587 Tuskegee National Forest, Macon County, AL, USA Southeastern Mixed Forests N KY574626
N5592 Canyonlands-South, Big Thicket National Preserve, TX, USA Piney Woods forests N KY574627
N5593 Canyonlands-South, Big Thicket National Preserve, TX, USA Piney Woods forests N KY574628
N5603 Canyonlands-South, Big Thicket National Preserve, TX, USA Piney Woods forests N MF770959
N5638 Tuskegee National Forest, Macon County, AL, USA Southeastern Mixed Forests N KY574629
N5643 Big Sandy Creek, Big Thicket National Preserve, TX, USA Piney Woods forests N KY574630
N5645 Big Sandy Creek, Big Thicket National Preserve, TX, USA Piney Woods forests N KY574631
N5712 Ichetucknee Springs State Park, FL, USA Southeastern Conifer forests N MN711217
N5713 Ichetucknee Springs State Park, FL, USA Southeastern Conifer forests N MN711218
N5714 Ichetucknee Springs State Park, FL, USA Southeastern Conifer forests N MN711219
N5715 Ichetucknee Springs State Park, FL, USA Southeastern Conifer forests N MN711220
N5726 Black Hills National Forest, Lawrence County, SD, USA South Central Rockies forests N KY574642
N5727 Black Hills National Forest, Lawrence County, SD, USA South Central Rockies forests N MN711221
N5728 Black Hills National Forest, Lawrence County, SD, USA South Central Rockies forests N MN711222
N5731 Black Hills National Forest, Lawrence County, SD, USA South Central Rockies forests N MN711223
N5813 Cuming County, NE, USA Central Tall Grasslands Y KY574637
N5814 Cuming County, NE, USA Central Tall Grasslands Y MN711224
N5815 Cuming County, NE, USA Central Tall Grasslands Y MN711225
N5816 Cuming County, NE, USA Central Tall Grasslands Y KY574632
N5943 Tulare County, CA, USA Central California Valley Y KY574638
P74053 Fresno County, CA, USA Central California Valley Y KJ787911
P194033 Konza Prairie Biological Station, Riley County, KS, USA Flint Hills Tall Grasslands Y KJ787912
P231026 Southeastern Fruit and Nut Research Station, Peach County, GA, USA Southeastern Plains Y KJ787888
P231028 Southeastern Fruit and Nut Research Station, Peach County, GA, USA Southeastern Plains Y KJ787889
P231030 Southeastern Fruit and Nut Research Station, Peach County, GA, USA Southeastern Plains Y KJ787879
P231031 Southeastern Fruit and Nut Research Station, Peach County, GA, USA Southeastern Plains Y KJ787890
P231032 Southeastern Fruit and Nut Research Station, Peach County, GA, USA Southeastern Plains Y KJ787891
P231034 Southeastern Fruit and Nut Research Station, Peach County, GA, USA Southeastern Plains Y KJ787892
P231035 Southeastern Fruit and Nut Research Station, Peach County, GA, USA Southeastern Plains Y KJ787893
P231036 Southeastern Fruit and Nut Research Station, Peach County, GA, USA Southeastern Plains Y KJ787894
P231037 Southeastern Fruit and Nut Research Station, Peach County, GA, USA Southeastern Plains Y KJ787895

Summary of genetic statistics for the 721-bp sequence isolated from the COI gene for each individual HG among the seven HG identified within the morphospecies Mesocriconema xenoplax. HG are organized in the table below according to their phylogenetic relationships shown.

HG na Sb PIc Hdd ke hf Pig
13 28 11 5 0.738 1.354 10 0.002
12 16 15 9 0.858 4.683 8 0.006
11 16 8 2 0.342 1.308 3 0.002
14 18 10 6 0.863 1.941 8 0.003
8 8 1 1 0.429 0.428 2 <0.001
10 23 7 3 0.711 1.628 6 0.002
9 20 60 50 0.932 19.53 13 0.027
Pooled 132 214 182 0.963 56.15 53 0.078

Population comparison defined by four categories of land use associated with the morphospecies Mesocriconema xenoplax in a pairwise AMOVA.

Number of Individuals Deciduous forest Developed open space Woody wetlands Cultivated crops Herbaceous
Deciduous forest 25 – 0.010 <0.001 0.011 <0.001
Developed open space 5 0.263 – 0.92 0.168 0.001
Woody wetlands 10 0.375 –0.114 – 0.027 <0.001
Cultivated crops 10 0.118 0.087 0.217 – 0.009
Herbaceous 9 0.654 0.725 0.757 0.330 –

Calculations of Birky’s 4× test on the morphospecies Mesocriconema xenoplax HG interclade divergence (K), intraclade variation (θ), and estimate of reciprocal monophyly (K/θ).

HG Ka θb K/θc
13 0.029 0.006 4.833*
12 0.029 0.031 0.935
11 0.060 0.006 10.000*
14 0.102 0.009 11.333*
8 0.079 0.003 26.333*
10 0.086 0.006 14.333*
9 0.089 0.096 0.927

Species delimitation statistics corresponding to the neighbor-joining tree HG built from the COI gene sequence for the morphospecies Mesocriconema xenoplax. Statistics include the intra-distance of each HG, inter-distance of each HG compared to its sister clade, ratio of intra- and inter-distance, P ID(strict), P ID(liberal), Av(MRCA-tips), Rodrigo’s P(AD), and Rosenberg’s P(AB). HG are organized in the table below according to their phylogenetic relationships shown.

HG Closest HG Intra-distance Inter-distance - closest Intra/inter P ID (Strict) P ID (Liberal) Av(MRCA-tips) P(AD) R(AB)
13 12 0.004 0.03 0.13 0.95 0.98 0.002 0.24 7.00E−14
12 13 0.007 0.03 0.23 0.92 0.97 0.005 1.00 7.00E−14
11 13 0.002 0.063 0.02 0.99 1.00 0.004 0.05 3.60E−18
14 8 0.008 0.118 0.07 0.97 0.99 0.004 0.05 1.40E−24
8 13 <0.001 0.095 0.01 0.94 1.00 <0.001 0.05 6.70E−15
10 8 0.005 0.098 0.05 0.98 1.00 0.004 0.41 3.62E−03
9 8 0.03 0.104 0.29 0.90 0.97 0.020 0.56 3.90E−26

Isolation by distance metric calculated by Mantel test statistic (Mantel R), and associated P-value for each of the seven HG within the morphospecies Mesocriconema xenoplax. HG are organized in the table below according to their phylogenetic relationships shown.

HG Mantel R P-value
13 0.048 0.731
12 0.206 0.118
11 0.989 <0.001*
14 –0.108 0.443
8 0.874 0.021*
10 0.627 <0.001*
9 0.284 0.005*
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