Fig. 1.
Light micrograph of Bursaphelenchus mucronatus. A: Female entire body; B: Male entire body; C: Lateral field with four incisures; D: Female anterior region; E, F: Vulva; G, H: Dauer larvae anterior part; I, J: Dauer larvae tail tip; K: Female tail tip; L: Mail tail; A, B, C, D, E, F, G, I, K, L: Georgian population; H, J: BiH population; A, B – 100 µm; C-L – 10 µm.
Table 1.
Morphometric comparison between the populations of Bursaphelenchus mucronatus from BiH and Georgia with the type population of B. mucronatus (Mamiya & Enda, 1979) from Japan. All measurements are in μm and in the form: mean ± s.d. (range).
| Dauers Georgian population | ♀♀ Georgian population | ♂♂ Georgian population | Dauers BiH population | Dauers Mamiya & Enda 1979 | ♀♀ Mamiya & Enda 1979 | ♂♂ Mamiya & Enda 1979 | |
|---|---|---|---|---|---|---|---|
| n | 5 | 25 | 12 | 5 | 30 | 40 | 35 |
| L | 597.5±19.8 (570–620) | 866.4± 77.0 (738–1018) | 675.3 ± 19.7 (652.5–723.0) | 589 ± 23.7 (558–620) | 590 (500–650) | 870 (700–980) | 790 (640–970) |
| a | 43.2±2.3 (39.5–45.6) | 40.9 ± 2.6 (35.0–45.7) | 42.6 ± 3.9 (35.8–50.4) | 41.7 ± 3.6 (35.8–45.0) | 39.8 (35.1–45.8) | 41.8 (36.5–45.9) | 44.0 (38.8–51.1) |
| b | 9.5* | 11.0 ± 1.2 (8.0–13.3) | 9.1 ± 0.4 (8.4–9.9) | 9.3* | 8.9 (7.9–9.6) | 12.6 (9.6–15.6) | 11.4 (9.0–14.7) |
| c | 20.7±1.40 (18.3–21.7) | 27.1 ± 2.8 (23.3–30.1) | 20.0 ± 1.1 (18.6–22.1) | 19.4 ± 1.3 (18.0–21.1) | 21.2 (19.7–24.4) | 26.2 (19.6–30.4) | 29.1 (25.7–35.6) |
| c′ | 3.6±0.3 (3.3–4.0) | 2.9 ± 0.2 (2.5–3.3) | 2.4 ± 0.1 (2.1–2.5) | 3.5 ± 0.2 (3.0–3.7) | |||
| V [%] | - | 74.1 ± 2.0 (67.0–76.9) | - | - | 75 (73–77) | ||
| Stylet length | 8.5* | 13.3 ± 1.2 (10.2–15.1) | 13.6 ± 0.6 (12.8–14.6) | 8.8* | 15.8 (14–16) | 15.0 (14–16) | |
| Excretory pore position1) | 54.9±4.5 (49–58.1) | 83.7 ± 3.6 (73.6–89.5) | 76.7 ± 4.2 (70.5–84.9) | 63,3±7,2 (54.0–74.4) | |||
| Pharynx length | 59.6* | 77.9 ± 4.2 (72.3–91.4) | 74.9 ± 3.0 (69.2–80.8) | 58.9* | |||
| MB2) | 59.9±4.9 (54.7–67.9) | 66.6 ± 3.7 (62.6–78.3) | 64.9± 2.5 (61.5–69.5) | 58.0 ± 5.7 (50.7–67.0) | |||
| Hemizonid position1) | - | 95.8± 5.2 (76.0–101.8) | 91.0 ± 2.5 (86.9–94.9) | - | |||
| Anal/cloacal body diameter | 8.1±0.2 (7.9–8.4) | 11.1± 0.7 (10.1–12.2) | 14.0 ± 0.7 (12.7–14.8) | 8.2 ± 0.3 (7.8–8.5) | |||
| Tail length | 29.1±2.8 (27.0–33.9) | 32.0± 2.2 (26.0–36.9) | 33.9 ± 1.9 (30.4–36.9) | 28.3 ± 2.55 (27.0–33.9) | |||
| Post-uterine sac length | - | 156.5 ± 17.9 (125.0–195.6) | - | - | |||
| Spicule length (curved median line) | - | - | 31.2 ± 1.5 (28.5–33.6) | - | 26.0 (23–29) |
Fig. 2.
Phylogenetic tree resulting from alignment of 28S rRNA gene sequences.
The tree was inferred by Maximum Likelihood method under the GTR + G + I model. Bootstrap values were obtained from 100 replicate. Values higher than 50% are presented. OK523378 and OK523379 are the original sequences.
Table 2.
Virtual ITS-RFLP patterns of Bursaphelenchus mucronatus from BiH and Georgia.
| BiH population, OK523378 | Georgian population, OK523379 | ||||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|
| ITS-PCR product | RsaI | HaeIII | MspI | HinfI | AluI | RsaI | HaeIII | MspI | HinfI | AluI | |
| Fragment sizes (~bp) | 884* | 490 | 623 | 355 | 411 | 678 | 412 | 623 | 355 | 411 | 678 |
| 412 | 196 | 304 | 232 | 246 | 264 | 196 | 304 | 232 | 246 | ||
| 22 | 105 | 265 | 120 | 226 | 105 | 265 | 120 | ||||
| 87 | 22 | 87 | |||||||||
| 49 | 49 | ||||||||||
| 25 | 25 | ||||||||||
Table 3.
List of Bursaphelenchus species described from Georgia.
| No. | status | Species | Insect vector | Associated plant/note | Group* | Reference |
|---|---|---|---|---|---|---|
| 1 | valid | B. chitwoodi Rühm, 1956 | Hylastes ater (Fabricius) (Coleoptera: Scolytidae) | Pinus sp. (Pinales: Pinaceae), exact pine species is not mentioned in the text | Hofmanni group | Kakulia & Shalibashvili, 1976a |
| Pinus sosnowskyi Nakai, Pinus pityosa Steven (Pinales: Pinaceae) | Kakulia, Shalibashvili, Gorgadze, 1983 | |||||
| 2 | valid | B. crenati Rühm, 1956 | Hylesinus crenatus (Fabricius) (Coleoptera: Scolytidae) | Fraxinus excelsior L. (Oleales: Oleaceae) | Xylophilus group | Kurashvili et al., 1980 |
| 3 | valid | B. eggersi Rühm, 1956 | Hylurgops palliatus (Gyllenhal) (Coleoptera: Scolytidae) | Picea orientalis (L.) (Pinales:Pinaceae); Abies sp., Larix sp.; Picea orientalis (L.), Pinus cedrus L. (Pinales: Pinaceae) | Eggersi group | Kakulia & Maglakelidze, 1973 |
| Kurashvili et al., 1980 | ||||||
| 4 | valid | B. eidmanni Rühm, 1956 | Ips typographus L. (Coleoptera: Scolytidae) | Abies sp., Larix sp., Picea orientalis (L.), Pinus cedrus L., P. sosnowskyi Nakai (Pinales: Pinaceae) | Eidemani group | Kakulia, 1971 |
| Georgia Ips typographus L. (Coleoptera: Scolytidae) | Kakulia & Devdariani, 1975** Kurashvili et al., 1980 | |||||
| 5 | valid | B. eremus Rühm, 1956 | Scolytus intricatus (Ratzeburg) (Coleoptera: Scolytidae) | Populus gracilis Grossh., Salix sp. (Salicales: Salicaceae), Castanea vulgaris Hance, Quercus iberica Steven ex Bieb., Q. pedunculata Ehrh., Q. sessiliflora Salisb. (Fagales: Fagaceae), Ulmus foliacea Gilib. (Urticales: Ulmaceae) | Eremus group | Kurashvili et al., 1980 |
| 6 | valid | B. ernoporus Devdariani, 1975 | Ernoporicus fagi (Fabricius) (Coleoptera: Scolitidae) | Fagus sp. (Fagales: Fagaceae), Carpinus sp. (Fagales: Betulaceae) | Insufficient description: not considered for Grouping | Kakulia & Devdariani, 1975** Mikaia et al., 2010 |
| Hunt, 1993 (nomen nudum) Ryss et al., 2005 (nomen nudum) | ||||||
| 7 | valid | B. erosus Kurashvili et al., 1980 | Orthotomicus erosus (Woll.) (Coleoptera: Scolytidae) | Abies sp., Picea orientalis (L.), Pinus sosnowskyi Nakai (Pinales: Pinaceae) Pinus pityusa Steven | Eidemani group | Kurashvili et al., 1980 |
| Kakulia, Shalibashvili, Gorgadze, 1983 | ||||||
| 8 | valid | B. eucarpus Rühm, 1956 | Scolytus mali (Bechstein & Scharfenberg) (Coleoptera: Scolytidae) | Malus domestica Borkh., Prunus sp., Sorbus sp. (Rosales: Rosaceae), Ulmaceae gen. sp. (Urticales) | Sexdentati group | Kakulia & Devdariani, 1975** |
| Kurashvili et al., 1980 | ||||||
| 9 | valid | B. fraudulentus Rühm, 1956 | Saperda carcharias(L.) (Coleoptera: Cerambycidae) | - | Xylophilus group | Kakulia & Devdariani, 1975** |
| Kurashvili et al., 1980 | ||||||
| 10 | valid | B. georgicus Devdariani et al., 1980 | Rhopalopus macropus Germar (Coleoptera: Cerambycidae) | Quercus iberica Schur. (Fagales: Fagaceae) | Insufficient description: not considered for grouping | Devdariani et al., 1980 |
| Kakulia, 1989 | ||||||
| Hunt, 1993 (nomen nudum) | ||||||
| Ryss et al., 2005 | ||||||
| 11 | valid | B. idius Rühm, 1956 | Pityogenes chalcographus L. (Coleoptera: Scolytidae) | Pinus sp. (Pinales: Pinaceae), Carpinus caucasica Grossh. (Betulales: Betulaceae), Juglans sp. (Juglandales: Juglandaceae), Populus tremula L. (Salicales: Salicaceae), Quercus iberica Steven ex Bieb. (Fagales: Fagaceae) | Six-incisure ungrouped species | Kakulia & Devdariani, 1975** |
| Kurashvili et al., 1980 | ||||||
| 12 | valid | B. incurvus Rühm, 1956 | Dendroctonus micans (Kugel.) (Coleoptera: Scolytidae) | Abies sp., Picea orientalis (L.), Pinus sylvestris L. (Pinales: Pinaceae) | Sexdentati group | Kurashvili et al., 1980 |
| 13 | valid | B. mucronatus Mamiya & Enda, 1979 | Monochamus galloprovincialis (Coleoptera: Cerambycidae) | Pinus sylvestris (L.) (Pinales: Pinaceae) | Xylophilus group | This article |
| 14 | valid | B. nuesslini Rühm, 1956 | Pityokeines curvidens (Germar) (Coleoptera: Scolytidae) | Not mentioned in the text. | Piniperdae group | Kakulia & Shalibashvili, 1976b |
| 15 | valid | B. piniperdae Fuchs, 1937 | Blastophagus piniperda (Coleoptera: Scolytidae) | Pinus sosnowskyi Nakai, Pinus pityosa Steven (Pinales: Pinaceae) | Piniperdae group | Kakulia, Devdariani, 1965 |
| Kakulia, 1967 | ||||||
| Kurashvili et al., 1980 | ||||||
| Kakulia, Shalibashvili, Gorgadze, 1983 | ||||||
| 16 | valid | B. populneus Devdariani, 1937 | Saperda populnea L. (Coleoptera: Cerambycidae) | Populus sp. | Insufficient description: not considered for grouping | Devdariani, 1973, * |
| Mikaia et al., 2010 | ||||||
| Hunt, 1993 (nomen nudum) | ||||||
| Ryss et al., 2005 (nomen nudum) | ||||||
| 17 | valid | B. ratzerburgii Rühm, 1956 | Scolytus ratzeburgii Janson (Coleoptera: Scolytidae) | Betula sp. (Betulales: Betulaceae) | Hofmanni group | Kurashvili et al., 1980 |
| 18 | valid | B. scalari Kakulia, 1989 | Saperda scalaris L. (Coleoptera: Cerambycidae) | Alnus glutinosa L. | Insufficient description: not considered for grouping | Kakulia, 1989 |
| 19 | valid | B. sexdentati Rühm, 1960 | Ips sexdentatus (Boerner) (Coleoptera: Scolytidae) | Picea orientalis (L.), Pinus sosnowskyi Nakai, Pinus pityosa Steven (Pinales: Pinaceae) | Sexdentati gropu | Kakulia & Devdariani, 1975** |
| Kurashvili et al., 1980 | ||||||
| Kakulia, Shalibashvili, Gorgadze, 1983 | ||||||
| 20 | valid | B. sutoricus Devdariani, 1974 | Monochamus sutor (L.) (Coleoptera: Cerambycidae) | Pinus sp. (Pinales: Pinaceae) | Sexdentati group | Devdariani, 1974 |
| 21 | valid | B. tbilisiensis Kakulia, 1989 | Saperda carcharias L. (Coleoptera: Cerambycidae) | Not mentioned in the text. | Insufficient description: not considered for grouping | Kakulia, 1989 |
| Hunt, 1993 (nomen nudum) | ||||||
| Ryss et al., 2005 (nomen nudum) | ||||||
| 22 | valid | B. wekuae Kakulia et al., 1978 | Trypophloeus sp. (erroneously named as Trypodendron sygnatum) (Coleoptera: Scolytidae) | Carpinus caucasica Grossh. (Betulales: Betulaceae), Fagus orientalis Lipsky. (Fagales: Fagaceae) | Insufficient description: not considered for Grouping | Kakulia et al., 1978** |
| Kurashvili et al., 1980 | ||||||
| Kakulia, 1989 | ||||||
| Hunt, 1993 (nomen nudum) | ||||||
| Ryss et al., 2005 | ||||||
| 23 | valid | B. xerocarterus Rühm, 1956 | Scolytus scolytus (Fabricius) (Coleoptera: Scolytidae) Scolytus scolytus (Fabricius), S. multistriatus (Marsh.) (Coleoptera: Scolytidae) | Ulmus foliacea Gilib., Zelkova sp. (Urticales: Ulmaceae), Carpinus caucasica Grossh. (Betulales: Betulaceae), Juglans sp. (Juglandales: Juglandaceae), Populus nigra L. (Salicales: Salicaceae) | Hofmanni/Abietinus group | Kakulia & Devdariani, 1967 |
| Kakulia & Devdariani, 1975** | ||||||
| Kurashvili et al., 1980 | ||||||
| Ryss et al., 2005 | ||||||
| 24 | species inquirendae vel incertae sedis | B. conurus (Steiner, 1932) | - | - | - | Kakulia, 1989 |
| Hunt, 1993 | ||||||
| Ryss et al., 2005 | ||||||
| 25 | species inquirendae vel incertae sedis | B. conjuctus | - | - | - | Kakulia, 1989 |
| Hunt, 1993 | ||||||
| Ryss et al., 2005 | ||||||
| 26 | species inquirendae vel incertae sedis | B. rüehmi | - | - | - | Kakulia, 1989 |
| Hunt, 1993 | ||||||
| Ryss et al., 2005 | ||||||
| 27 | nomen nudum | B. hylesini | - | - | - | Devdariani, 1975** |
| Kurashvili et al., 1980 | ||||||
| This article (Nomina Nuda) | ||||||
| 28 | Transfer to other genus | B. lignophilus (Körner, 1954) | - | Laimaphelenchus lignophilus (Körner, 1954) | - | Kakulia, 1989 |
| Meyl, 1961 | Goodey, 1960 | Ryss et al., 2005 | ||||
| 29 | Transfer to other genus | B. teratospicularis Kakulia & Devdariani, 1965 | - | Devibursaphelenchus teratospicularis | - | Kakulia & Devdariani, 1965 |
| Kakulia, 1989 | ||||||
| Ryss et al., 2005 | ||||||
| Braasch, 2009 | ||||||
| 30 | Transfer to other genus | B. typographi Kakulia, 1967 | - | Devibursaphelenchus typographi | - | Kakulia, 1967 |
| Kakulia, 1989 | ||||||
| Ryss et al., 2005 | ||||||
| Braasch, 2009 |
Legend:
* = grouping is based on Braasch et al. (2009);