Table 1.
List of the species used for phylogenetic analysis based on rDNA and mtDNA available in the GenBank for Pratylenchus and Hirschmanniella.
| Pratylenchus | Hirschmanniella | ||||||||
|---|---|---|---|---|---|---|---|---|---|
| 18S rDNA | ITS rDNA | 28S rDNA | COI mtDNA | 28S rDNA | |||||
| Species | Accession number/locality | Species | Accession number/locality | Species | Accession number/locality | Species | Accession number/locality | Species | Accession number/locality |
| P. speijeri | KM245059/China | P. parafloridensis | GQ988378/USA | P. hippeastri | FN554882/USA | P. coffeae | KU198943/Japan | H. oryzae | JX291141/Myanmar |
| P. speijeri | KF974690/China | P. parafloridensis | GQ988377/USA | P. hippeastri | FM994115/USA | P. coffeae | KU198942/Japan | H. oryzae | JX291142/Myanmar |
| P. coffeae | AB905286/Japan | P. floridensis | GQ988375/USA | P. hippeastri | FN554881/USA | P. coffeae | KY424075/China | Hirschmanniella sp. | DQ328686/Vietnam |
| P. coffeae | KM245066/China | P. floridensis | GQ988376/USA | P. hippeastri | FN554879/USA | P. coffeae | KY424074/China | H. oryzae | KF201169/the Philippines |
| P. coffeae | KY424134/China | P. hippeastri | FJ712933/USA | P. hippeastri | FM994114/USA | P. speijeri | KY424088/China | H. oryzae | KF201165/the Philippines |
| P. speijeri | KF974688/ China | P. hippeastri | FJ712935/USA | P. hippeastri | KP161611/China | P. speijeri | KY424087/China | H. oryzae | KF201161/the Philippines |
| P. coffeae | KY424139/China | P. hippeastri | FJ712934/USA | P. hippeastri | KC796704/Japan | P. loosi | KY424086/Japan | H. belli | EF029860/USA |
| P. coffeae | KY424140/China | P. hippeastri | FJ712936/USA | P. hippeastri | KC796705/Japan | P. loosi | KY424085/China | Hirschmanniella sp. | EF029861/USA |
| P. coffeae | KY424142/China | P. hippeastri | KY424236/China | P. hippeastri | KC796706/Japan | P. loosi | KY424084/China | Hirschmanniella sp. | KP671713/Belgium |
| P. speijeri | KY424156/China | P. hippeastri | KR029085/China | P. hippeastri | KC796707/Japan | P. loosi | KX349422/China | H. kwazuna | South Africa |
| P. speijeri | KM245067/China | P. hippeastri | KY424237/China | P. hippeastri | KY424307/China | P. hippeastri | KY424099/China | H. kwazuna | South Africa |
| P. coffeae | KY424137/China | P. hippeastri | KC796698/Japan | P. hippeastri | KR029084/China | P. hippeastri | KY424098/China | H. loofi | EU620468/Belgium |
| P. coffeae | KY424143/China | P. hippeastri | KC796701/Japan | P. hippeastri | KY424306/China | P. scribneri | KY424092/China | H. loofi | EU620469/Belgium |
| P. loosi | AB905296/Japan | P. hippeastri | KC796702/Japan | P. hippeastri | KJ001720/China | P. scribneri | KX349425/China | Hirschmanniella sp. | JX261958/Iran |
| P. loosi | KY424153/China | P. hippeastri | KC796699/Japan | P. hippeastri | GU214112/USA | P. scribneri | KY424090/China | H. mucronata | KP179327/Cambodia |
| P. loosi | KY424154/China | P. hippeastri | KJ001718/Israel | P. hippeastri | KC796703/Japan | P. scribneri | KY424091/China | H. mucronata | KF201167/the Philippines |
| P. loosi | KY424155/China | P. hippeastri | FJ712932/USA | P. parafloridensis | GU214114/USA | P. scribneri | KY424089/China | H. mucronata | KP179333/Cambodia |
| P. loosi | AB905297/Japan | P. hippeastri | KC796700/Japan | P. parafloridensis | AF170438/USA | P. vulnus | KY828317/Belgium | H. halophila | EU620464/Germany |
| P. agilis | EU130794/USA | P. hippeastri | FN554883/USA | P. parafloridensis | GU214115/USA | P. vulnus | KY828312/Belgium | H. halophila | EU620465/Germany |
| P. agilis | EU130793/USA | P. hippeastri | FN554887/USA | P. floridensis | GU214116/USA | P. vulnus | KY424096/China | H. pomponiensis | DQ077795/USA |
| P. scribneri | EU130812/USA | P. hippeastri | FN554884/USA | P. floridensis | AF170437/USA | P. vulnus | KY424094/China | H. santarosae | EF029859/USA |
| P. scribneri | EU130811/USA | P. hippeastri | FN554888/USA | P. floridensis | GU214117/USA | P. oleae | KJ510866/Spain | P. vulnus | EU130885/USA |
| P. scribneri | KY424158/China | P. jaehni | FJ712941/Brazil | P. araucensis | FJ463261/Colombia | Rotylenchulus macrosoma | KY992847/Greece | ||
| P. scribneri | KY424159/China | P. jaehni | FJ712940/Brazil | P. araucensis | FJ463258/Colombia | ||||
| P. scribneri | EU669927/the Netherlands | P. loosi | FJ712946/Brazil | P. araucensis | FJ463260/Colombia | ||||
| P. scribneri | KY424162/China | P. loosi | FJ712942/Brazil | P. coffeae | AF170427/USA | ||||
| P. scribneri | KY424161/China | P. pseudocoffeae | KT971367/Costa Rica | P. coffeae | AF170426/USA | ||||
| P. hippeastri | KY424166/China | P. pseudocoffeae | KT175523/South Korea | P. pseudocoffeae | KT175531/South Korea | ||||
| P. hippeastri | KJ001716/Israel | P. pseudocoffeae | LC030339/Japan | P. pseudocoffeae | KT175532/South Korea | ||||
| P. scribneri | EU669958/the Netherlands | P. pseudocoffeae | LC030338/Japan | P. pseudocoffeae | KT971360/Costa Rica | ||||
| P. araucensis | FJ154950/Colombia | P. scribneri | KY424228/China | P. pseudocoffeae | KT175533/South Korea | ||||
| P. japonicus | KF385443/Japan | P. scribneri | KY424230/China | P. scribneri | KY424300/China | ||||
| P. parazeae | KY424184/China | P. agilis | FJ712891/USA | P. agilis | EU130841/USA | ||||
| P. pratensis | KC875387/the Netherlands | P. agilis | JQ039330/China | P. scribneri | JX047002/China | ||||
| P. bolivianus | KC875390/the Netherlands | P. alleni | JX081545/Canada | P. scribneri | EU130865/USA | ||||
| Zygotylenchus guevarae | AF442189/Belgium | P. gutierrezi | FJ712929/Guatemala | P. scribneri | KX842632/USA | ||||
| P. gutierrezi | FJ712930/Guatemala | P. alleni | MF155653/Canada | ||||||
| P. gutierrezi | FJ712931/Guatemala | P. speijeri | KF974713/China | ||||||
| P. gutierrezi | FR692277/Portugal | P. speijeri | KF974715/China | ||||||
| Zygotylenchus guevarai | FJ717817/Spain | P. speijeri | KY424295/China | ||||||
| P. speijeri | KF974716/China | ||||||||
| P. speijeri | KF974703/China | ||||||||
| P. coffeae | EU130846/Japan | ||||||||
| P. coffeae | EU130850/Japan | ||||||||
| P. coffeae | EU130845/Japan | ||||||||
| P. coffeae | KC490925/China | ||||||||
| P. loosi | KY424290/China | ||||||||
| P. loosi | EF446995/Iran | ||||||||
| P. loosi | KY424291/China | ||||||||
| P. loosi | KY424293/China | ||||||||
| P. loosi | JN091970/Japan | ||||||||
| P. penetrans | JX046999/China | ||||||||
| P. penetrans | JX046998/China | ||||||||
| P. dunensis | AJ890462/the Netherlands | ||||||||
| P. dunensis | AJ890460/the Netherlands | ||||||||
| P. brachyurus | KF712474/China | ||||||||
| P. brachyurus | KF712472/China | ||||||||
| P. vulnus | HM469437/China | ||||||||
| P. vulnus | KF430799/Japan | ||||||||
| P. crenatus | KX683378/the Netherlands | ||||||||
| P. crenatus | EU130853/UK | ||||||||
| P. bhattii | JN244270/China | ||||||||
| P. bhattii | JN244269/China | ||||||||
| P. parazeae | KP903445/China | ||||||||
| P. parazeae | KP903443/China | ||||||||
| P. zeae | KT033000/Kenya | ||||||||
| P. zeae | KT032999/Kenya | ||||||||
| P. bolivianus | KU198956/Bolivia | ||||||||
| P. bolivianus | KU198955/Bolivia | ||||||||
| P. neglectus | HM469438/China | ||||||||
| P. neglectus | MG205581/China | ||||||||
| P. brzeskii | AM231928/France | ||||||||
| P. brzeskii | AM231927/France | ||||||||
| P. thornei | KX258736/Iran | ||||||||
| P. thornei | KX258737/Iran | ||||||||
| P. thornei | EU130881/Moldova | ||||||||
| Zygotylenchus guevarai | JQ917439/Iran | ||||||||
Table 2.
Nematode species and GenBank accession numbers used for the present study.
| Species | Gene | GenBank accession number | Origin | Sample codes |
|---|---|---|---|---|
| P. hippeastri | 18S rDNA | MH324470 | Potchefstroom, South Africa | ESW 1 |
| P. hippeastri | ITS rDNA | MH324471 | Potchefstroom, South Africa | ESW 2 |
| P. hippeastri | 28S rDNA | MH324472 | Potchefstroom, South Africa | ESW 3 |
| P. hippeastri | 28S rDNA | MH324473 | Potchefstroom, South Africa | ESW 4 |
| P. hippeastri | COI of mtDNA | MH324474 | Potchefstroom, South Africa | ESW 5 |
| H. anchoryzae | 28S rDNA | MK571451 | Royan, Iran | IR Royan |
| H. anchoryzae | COI of mtDNA | MK583962 | Royan, Iran | IR Royan |
Table 3.
Morphometrics of H. anchoryzae from Iran and P. hippeastri from South Africa. All measurements are in μm and in the form: mean ± s.d. (range).
| Species | H. anchoryzae | P. hippeastri | |
|---|---|---|---|
| Locality | Royan | Potchefstroom | |
| Province | Mazandaran Province | North-West Province | |
| Country | Iran | South Africa | |
| Habitat | Mentha aquatica | Willow tree | |
| n | 4 ♀♀ | 2 ♂♂ | 7 ♀♀ |
| L | 1,796 ± 71 (1,740–1,895) | 1,273, 1,625 | 522.6 ± 69.5 (424–614) |
| a | 65.0 ± 2.0 (63.6–68.0) | 62.1, 62.5 | 28.5 ± 3.1 (18.6–31.8) |
| b | 10.3 ± 1.4 (9.0–11.6) | 6.6, 8.3 | 3.8 ± 0.7 (2.7–4.9) |
| c | 18.6 ± 2.3 (16.4–21.9) | 15.7, 18.8 | 18.7 ± 3.1 (11.7–23.6) |
| c′ | 5.5 ± 1.6 (4.2–7.2) | 4.2, 5.4 | 2.2 ± 0.2 (2.0–2.3) |
| V | 55.4 ± 1.9 (53–57) | – | 77.1 ± 3.5 (73–82) |
| Lip region height | 4.1 ± 0.2 (4–5) | 3, 4 | 2.6 ± 0.7 (1.9–3.7) |
| Lip region diameter | 10.3 ± 0.5 (10–11) | 9, 11 | 8.5 ± 1.7 (7–11) |
| Stylet length | 20.2 ± 1 (19–21) | 18, 19 | 15.4 ± 1.6 (13–18) |
| Stylet conus length | 9.8 ± 0.7 (9–11) | 9, ? | 7.6 ± 0.7 (6.4–8.2) |
| Stylet shaft length | 8.4 ± 0.5 (8–9) | 8, ? | 4.9 ± 1.2 (4.2–5.8) |
| Stylet knob height | 1.7 ± 0.2 (1.5–1.8) | 1.5, ? | 2.2 ± 0.4 (1.7–2.5) |
| Stylet knob width | 3.6 ± 0.3 (3.3–3.8) | 3.4, ? | 3.4 ± 0.7 (2.7–4.1) |
| DGO from stylet base | 3.2 ± 0.4 (3–4) | 3.3, 3.7 | 2.7 ± 0.9 (1.7–3.6) |
| Anterior end to centre of median bulb | 87 ± 3 (84–91) | 77, 93 | 53.8 ± 3.7 (50–60) |
| End of pharyngeal glands | 393 ± 13 (380–410) | 223, 441 | 132.9 ± 15.9 (116–149) |
| Median bulb length | 12.7 ± 0.2 (12–13) | 13 | 13.8 ± 2.7 (11–16) |
| Median bulb width | 16.6 ± 0.5 (16–17) | 18 | 9.1 ± 1.8 (7–10) |
| Excretory pore – anterior end | 150 ± 2 (148–152) | 119, 143 | 95.7 ± 15.1 (80–112) |
| Maximum body diameter | 28 ± 2 (26–29) | 21, 26 | 20.9 ± 3.4 (17–27) |
| Anal body diameter | 18 ± 4 (13–20) | 15, 20 | 12.9 ± 1.0 (12–14) |
| Anterior genital tract length | ? | – | 160.5 ± 36.5 (102–210) |
| Tail length | 101 ± 4 (96–106) | 81, 87 | 31.7 ± 6.4 (26–43) |
| Number of tail annuli | 73 ± 9 (63–80) | ? | 21.3 ± 2.8 (18–23) |
| Vulva to anus distance | – | – | 77.5 ± 7.7 (72–83) |
| Post-vulval uterine sac length | – | – | 20.6 ± 3.9 (16–26) |
| Lateral field width | 7.5 | 7 | 5.5 ± 0.7 (5–6) |
| Phasmid-anus distance | 57.8 ± 7.6 (46–64) | 49, 61 | 17.2 ± 4.1 (15-22) |
| Spicules | – | 27, 30 | – |
| Gubernaculum | – | 8.6-8.7 | – |

Figure 1:
Line drawings of Hirschmanniella anchoryzae. (A) anterior portion of the female; (B) cephalic region of the female; (C–E) female posterior end; (F) male posterior end; (G) female reproductive system; (H, I) status of females after relaxation; (J, K) status of male after relaxation.

Figure 2:
PCA analysis of the different population of H. anchoryzae.
Table 4.
Genetic pairwise distance estimation of 28S rDNA of Hirschmanniella species using Mega 7.
| Species | Locality | 1 | 2 | 3 | 4 | 5 | 6 | 7 | 8 | 9 | 10 | 11 | 12 | 13 | |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| 1 | H. anchoryzae | Iran | 0.035 | 0.049 | 0.051 | 0.032 | 0.048 | 0.055 | 0.052 | 0.053 | 0.056 | 0.024 | 0.058 | 0.053 | |
| 2 | H. pomponiensis | USA | 0.109 | 0.040 | 0.039 | 0.007 | 0.038 | 0.045 | 0.044 | 0.044 | 0.047 | 0.024 | 0.047 | 0.041 | |
| 3 | Hirschmanneilla sp. | Iran | 0.147 | 0.126 | 0.031 | 0.038 | 0.027 | 0.030 | 0.030 | 0.016 | 0.014 | 0.046 | 0.022 | 0.030 | |
| 4 | Hirschmanneilla sp. | USA | 0.146 | 0.122 | 0.102 | 0.035 | 0.012 | 0.014 | 0.014 | 0.031 | 0.030 | 0.040 | 0.031 | 0.004 | |
| 5 | H. santarosae | USA | 0.101 | 0.019 | 0.123 | 0.110 | 0.033 | 0.039 | 0.038 | 0.039 | 0.042 | 0.022 | 0.042 | 0.036 | |
| 6 | H. belli | USA | 0.137 | 0.116 | 0.088 | 0.036 | 0.103 | 0.007 | 0.006 | 0.026 | 0.026 | 0.038 | 0.031 | 0.011 | |
| 7 | H. oryzae | Myanmar and the Philippines | 0.152 | 0.131 | 0.097 | 0.043 | 0.119 | 0.016 | 0.000 | 0.029 | 0.028 | 0.042 | 0.031 | 0.012 | |
| 8 | Hirschmanneilla sp. | Vietnam | 0.146 | 0.129 | 0.098 | 0.044 | 0.117 | 0.015 | 0.000 | 0.030 | 0.029 | 0.044 | 0.031 | 0.012 | |
| 9 | H. kwazuna | South Africa | 0.149 | 0.131 | 0.050 | 0.097 | 0.119 | 0.082 | 0.090 | 0.092 | 0.010 | 0.039 | 0.018 | 0.028 | |
| 10 | H. loofi | Belgium | 0.157 | 0.139 | 0.044 | 0.097 | 0.128 | 0.083 | 0.089 | 0.092 | 0.026 | 0.041 | 0.017 | 0.028 | |
| 11 | H. halophila | Germany | 0.073 | 0.075 | 0.140 | 0.121 | 0.069 | 0.113 | 0.124 | 0.128 | 0.118 | 0.124 | 0.043 | 0.040 | |
| 12 | H. mucronata | Belgium | 0.166 | 0.145 | 0.074 | 0.103 | 0.132 | 0.100 | 0.100 | 0.100 | 0.058 | 0.055 | 0.132 | 0.031 | |
| 13 | Hirschmanneilla sp. | Belgium | 0.149 | 0.124 | 0.098 | 0.009 | 0.111 | 0.030 | 0.036 | 0.036 | 0.089 | 0.089 | 0.120 | 0.101 |
1 Notes: Accession numbers: 1 = MK571451; 2 = DQ077795; 3 = JX261958; 4 = EF029861; 5 = EF029859; 6 = EF029860; 7 = JX291141, JX291142, KF201161, KF201165, KF201169; 8 = DQ328686; 9 = EU620466, EU620467; 10 = EU620468, EU620469; 11 = EU620464, EU620465; 12 = KP179327, KP179333, KF201167; 13 = KP671713.

Figure 3:
The Bayesian inference tree of Hirschmanniella anchoryzae (Ebsary and Anderson, 1982) from Iran and other related species based on the sequences from 28S rDNA under GTR+I+G model (−lnL = 3,374.3581; AIC = 6,856.7162; freqA = 0.2269; freqC = 0.2193; freqG = 0.3068; freqT = 0.2471; R(a) [AC] = 0.6057; R(b) [AG] = 2.7984; R(c) [AT] = 0.9170; R(d) [CG] = 0.2612; R(e) [CT] = 3.5814; R(f) [GT] = 1; p-inv = 0.1960; shape = 0.5340).

Figure 4:
Line drawings of Pratylenchus hippeastri. (A) female anterior end; (B, C) stoma; (D) female reproductive system; (E) entire female; (F) lateral field; (G) post uterine sac; (H, I) female posterior end (arrow indicates phasmid).

Figure 5:
Light photomicrographs of Pratylenchus hippeastri. (A, B) anterior end (arrows indicate hemizonid); (C) reproductive system (arrow indicates vulva); (D) entire body (black arrow indicates vulva, white arrows indicate phasmids); (E) posterior end (arrow indicate anus).

Figure 6:
Scanning electron microscope photographs of Pratylenchus hippeastri. (A) entire body (black arrow indicates vulva); (B, C, E) lip region in lateral, frontal and ventral views, respectively); (D) female anterior region; (F) excretory pore (arrow); (G) lateral field (arrows indicate longitudinal incisures); (H) vulval region; (I, J) female posterior end in ventral and lateral views, respectively (arrow indicates phasmid); (K) anus.
Table 5.
Correlation of morphometric data of P. hippeastri from South Africa.
| Variables | L | a | b | c | ć | V | Stylet | DGO | Tail | PUS | Phasmid | MB | Pharynx |
|---|---|---|---|---|---|---|---|---|---|---|---|---|---|
| L | 1 | −0.236 | 0.786 | 0.336 | −0.340 | −0.453 | 0.277 | −0.404 | 0.768 | 0.672 | 0.626 | 0.931 | −0.841 |
| a | −0.236 | 1 | −0.516 | 0.589 | −0.210 | −0.285 | −0.601 | −0.166 | −0.605 | −0.846 | 0.001 | −0.449 | −0.205 |
| b | 0.786 | −0.516 | 1 | −0.187 | 0.198 | −0.503 | 0.792 | −0.287 | 0.904 | 0.806 | 0.797 | 0.709 | −0.679 |
| c | 0.336 | 0.589 | −0.187 | 1 | −0.882 | 0.082 | −0.488 | −0.692 | −0.344 | −0.191 | −0.134 | 0.140 | −0.359 |
| ć | −0.340 | −0.210 | 0.198 | −0.882 | 1 | −0.427 | 0.471 | 0.607 | 0.275 | −0.079 | 0.407 | −0.280 | 0.103 |
| V | −0.453 | −0.285 | −0.503 | 0.082 | −0.427 | 1 | −0.181 | −0.141 | −0.543 | 0.059 | −0.913 | −0.329 | 0.811 |
| Stylet | 0.277 | −0.601 | 0.792 | −0.488 | 0.471 | −0.181 | 1 | −0.257 | 0.588 | 0.679 | 0.531 | 0.192 | −0.197 |
| DGO | −0.404 | −0.166 | −0.287 | −0.692 | 0.607 | −0.141 | −0.257 | 1 | 0.089 | −0.268 | −0.094 | −0.120 | 0.347 |
| Tail | 0.768 | −0.605 | 0.904 | −0.344 | 0.275 | −0.543 | 0.588 | 0.089 | 1 | 0.771 | 0.736 | 0.831 | −0.614 |
| PUS | 0.672 | −0.846 | 0.806 | −0.191 | −0.079 | 0.059 | 0.679 | −0.268 | 0.771 | 1 | 0.296 | 0.738 | −0.261 |
| Phasmid | 0.626 | 0.001 | 0.797 | −0.134 | 0.407 | −0.913 | 0.531 | −0.094 | 0.736 | 0.296 | 1 | 0.472 | −0.856 |
| MB | 0.931 | −0.449 | 0.709 | 0.140 | −0.280 | −0.329 | 0.192 | −0.120 | 0.831 | 0.738 | 0.472 | 1 | −0.655 |
| Pharynx | −0.841 | −0.205 | −0.679 | −0.359 | 0.103 | 0.811 | −0.197 | 0.347 | −0.614 | −0.261 | −0.856 | −0.655 | 1 |

Figure 7:
PCA analysis of the different population of P. hippeastri.

Figure 8:
Cluster dendrogram for different populations of P. hippeastri using morphometric data. Red values represent AU (approximated unbiased) values. Green values on the right branch indicate BP (bootstrap probability). Florida 1 (Inserra et al. 2007) and Florida 2 (De Luca et al. 2010).

Figure 9:
The Bayesian inference tree of Pratylenchus hippeastri from South Africa and other related taxa based on the sequences from 18S rDNA under GTR+I+G model (−lnL = 5,036.0855; AIC = 10,236.171; freqA = 0.2586; freqC = 0.2234; freqG = 0.2663; freqT = 0.2517; R(a) [AC] = 1.29106; R(b) [AG] = 2.99041; R(c) [AT] = 1.68788; R(d) [CG] = 0.89263; R(e) [CT] = 6.4881; R(f) [GT] = 1; p-inv = 0.5010; Shape = 0.4870).

Figure 10:
The Bayesian inference tree of Pratylenchus hippeastri from South Africa and other related taxa based on the sequences from ITS rDNA under GTR+I+G model (−lnL = 7,745.2851; AIC = 15,674.5702; freqA = 0.2437; freqC = 0.2123; freqG = 0.255; freqT = 0.2889; R(a) [AC] = 1.07478; R(b) [AG] = 2.56737; R(c) [AT] = 1.63147; R(d) [CG] = 0.53909; R(e) [CT] = 2.91622; R(f) [GT] = 1; p-inv = 0.2300; Shape = 1.3540).

Figure 12:
The Bayesian inference tree of Pratylenchus hippeastri from South Africa and other related taxa based on the sequences from COI of mtDNA under GTR+I+G model (−lnL = 2,667.1378; AIC = 5,446.2756; freqA = 0.2552; freqC = 0.0926; freqG = 0.1917; freqT = 0.4604; R(a) [AC] = 0.01; R(b) [AG] = 9.49586; R(c) [AT] = 2.67386; R(d) [CG] = 3.1383; R(e) [CT] = 8.02121; R(f) [GT] = 1; p-inv = 0.2150; Shape = 0.4730).

Figure 11:
The Bayesian inference tree of Pratylenchus hippeastri from South Africa and other related taxa based on the sequences from 28S rDNA under GTR+I+G model (−lnL = 7,451.6325; AIC = 15,235.265; freqA = 0.2081; freqC = 0.2296; freqG = 0.3327; freqT = 0.2296; R(a) [AC] = 0.83418; R(b) [AG] = 2.50021; R(c) [AT] = 1.25212; R(d) [CG] = 0.34218; R(e) [CT] = 4.6954; R(f) [GT] = 1; p-inv = 0.2510; Shape = 0.6830).