A Key to Species in the Genus Bursaphelenchus with a ...DIAGNOSIS: Bursaphelenchus hunanensis sp. n....

Proc. Helminthol. Soc. Wash.55(1), 1988, pp. 1-11

A Key to Species in the Genus Bursaphelenchus with a Description ofBursaphelenchm hunanensis sp. n. (Nematoda: Aphelenchoididae)Found in Pine Wood in Hunan Province, China1

KAMCHEW YiN,2 YUSHENG FANG,2 AND A. C. TARjAN 3

2 Plant Nematology Research Laboratory, Department of Plant Protection,South China Agricultural University, Guangzhou, China and3 Department of Entomology and Nematology, IFAS, University of Florida, Gainesville, Florida 32611

ABSTRACT: Bursaphelenchus hunanensis sp. n. is described and illustrated. It has a distinct stylet longer thanother species in the genus and has no basal thickenings. The female lacks a vulval flap and has a short postuterinebranch. The male spicules do not have a cucullus and terminate posteriad in a blunt rounded tip. The rostrumis digitate. An alphabetical list of all 49 nominal species in the genus, a key to 44 valid species, and tables ofdiagnostic data of males and females are also included.

KEY WORDS: Nematoda, Aphelenchoididae, Bursaphelenchus hunanensis sp. n., nematode taxonomy andsystematics, Pinus massoniana, key to species.

Species within the genus BursaphelenchusFuchs, 1937, were regarded only as associates ofbark beetles (Ipidae) until the insidious pine-wood nematode B. xylophilus (Steiner and Buhr-er, 1934) Nickle, 1970, started to devastate entireforests in Japan (Mamiya, 1972). Current con-cern on chance introduction of this species intonew areas has renewed interest in the genus andits species.

A Bursaphelenchus species was first found inOctober 1984 in the wood of dead pine trees,Pinus massoniana Lamb, in Huaihua County,Hunan Province, China, during a survey for thepine wil t disease caused by Bursaphelenchus xy-lophilus. Detailed studies of the nematode re-vealed that it differed from the 43 species listedin the generic review by Tarjan and Baeza-Ara-gon (1982) and 5 additional nominal species(Kakuliya and Devdariani, 1965; Devdariani etal., 1980; Giblin and Kaya, 1983; Thong andWebster, 1983; Giblin et al., 1984).

The purpose of this paper, in addition to re-porting a new species, is to provide a key and atable of essential measurements and ratios ofnominal species within the genus.

Materials and Methods

Bursaphelenchus specimens were extracted by theBaermann funnel method from wood fragments cutfrom wilted trees. Studies were made on living as wellas fixed specimens. Measurements and drawings weremade from specimens fixed in TAF (triethanolamine,

1 Florida Agricultural Experiment Station JournalSeries No. 7922.

formaldehyde, and water) and stained with poly-chrome blue. Spicule lengths were measured along themedian line of the arc of the spicule. The specimenswere mounted in glycerin. Measurements are in mi-crometers unless stated otherwise.

Information on other Bursaphelenchus species wasobtained solely from perusal of pertinent references.Drawings of spicules of each species were made fromthe original publication, and then either enlarged orreduced to a standard size. The majority of the spiculedrawings shown in Figure 13 came from Tarjan andBaeza-Aragon (1982).

Generic Diagnosis

There have been no emendations proposed byothers to the generic diagnosis of the genus givenby Nickle (1970) which is presented below:

Bursaphelenchus Fuchs, 1937Syn. Aphelenchoides (Bursaphelenchus)

(Fuchs, 1937) Riihm, 1956

DEFINITION: Aphelenchoididae. Usually long.Lips high, offset. Stylet with small rounded basalknobs. Excretory pore usually behind medianbulb. Vulval lips sometimes protruding; post-uterine sac usually long. Female tail rounded,conoid, or sharply pointed. Spicules large, nar-row, usually with prominent rostrum. Male tailstrongly arcuate, variously pointed, with short,terminal caudal alae. Usually 2 pairs of caudalpapillae, 1 adanal, the other postanal. Guber-naculum absent.

BIONOMICS: Mycophagous, insect associate.TYPE SPECIES: Bursaphelenchus piniperdae

Fuchs, 1937.

1

Copyright © 2011, The Helminthological Society of Washington

PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY

50 jum1,2,3

4,5,6,7,8,9,10 10 >um

10 jam

11

10

11

Figures 1-11. Bursaphelenchus hunanensis sp. n. 1. Female. 2. Male. 3. Posterior part of male showing flexureof the anterior end of the testis. 4. Female anterior end. 5. Vulva region. 6. Female tail. 7. Lateral view of cuticlein midbody region. 8. En face view of head. 9. Lateral view of male tail. 10. Ventral view of male tail. 11. Malespicule.


OF WASHINGTON, VOLUME 55, NUMBER 1, JANUARY 1988

Results and Discussion

Bursaphelenchus hunanensis sp. n.(Figs. 1-11)

FEMALES (30) (paratypes): L = 0.67 (0.58-0.79) mm; a = 35.5 (31.1-41.5); b = 8.4 (7.3-9.6); c = 15.9 (13.6-18.6); V = 77.9 (75.7-79.5)%;stylet = 22 (20-26).

MALES (15) (paratypes): L = 0.53(0.48-0.59)mm; a = 40.9 (37.6-45.3); b = 7.1 (6.5-8.8);c = 16.1 (15-19.1); T = 26.1 (16.4-31.7)%; sty-let = 20 (19-21); spicule = 15 (13-20).

FEMALE (holotype): L = 0.67 mm; a = 36.4;b = 7.7; c = 17; V = 79.2%; stylet = 23.

MAL E (allotype): L = 0.52 mm; a = 39.4;b = 6.6; c = 16.4; T = 26.3%; stylet = 19; spic-ule = 15.

FEMALE: Body ventrally curved when killedwith heat, tapering gradually near extremities.Cuticle marked by fine annules, about 1 nm widein midbody region, interrupted by a narrow lat-eral field which is about '/10 of the maximum bodywidth and has 2 longitudinal, refractive lines oneither side of a narrow refractive band (Fig. 1).Lip flattened at oral opening, set off by a con-striction. En face view of head showing 6 equallips (Fig. 8). Anterior stylet cone less than '/2 totalstylet length, posterior shaft without basal thick-ening (Fig. 4). Procorpus cylindrical, twice as longas vulval body width. Median bulb elongate-oval,with conspicuous valve just posterior to center.Excretory pore Vi to 1 length of median bulbbehind the bulb on a level with, or just behind,the nerve ring. Esophageal glands dorsally over-lapping intestine, 9 times as long as vulval bodywidth. Reproductive system prodelphic, gonadoutstretched, occupying Vs the body length. Oo-cytes arranged in single fil e except at the anteriorpart of the ovary. Vagina an anteriorly inclinedtube, without vulval flap. Postuterine branchusually shorter than vulva body diameter (Fig.5). Vulva-anus distance equal to 2.5 times taillength. Tail conoid with slight ventral curvature,3 times longer than anal body width, with a finelyrounded terminus (Fig. 6).

MALE: Body J-shaped when killed with heat.Anterior body region similar to female. Repro-ductive system monorchic, testis usually out-stretched, extending 1A of body length. Approx-imately V& of the testis reflexed anteriorly (Fig.3). Spermatogonia arranged in several rows andspermatocytes in single file. Seminal vesicle oval.Vas deferens composed of anterior cylindricalpart and posterior slender duct. Spicules paired,

CONDYLUS

CAPITULUM

ROSTRUM

CALOMUS

LAMINA

CUCULLUS

CAUDAL ALA

(BURSA)

Figure 12. Lateral view of the Bursaphelenchusspicules (after Tarjan and Baeza-Aragon, 1982).

arcuate, with blunt prominent rostrum; distal endof spicule obtuse, without cucullus (cf. Fig. 2).Center of capitulum depressed, condylus hemi-spherically rounded. Tail ventrally arcuate, ter-minus finely rounded, with short terminallytruncated bursa (Fig. 10). Two pairs of caudalpapillae present; 1 adanal pair, 1 pair about Vitail length from tail terminus.

DIAGNOSIS: Bursaphelenchus hunanensis sp.n. differs from all other species in the genus bythe length of female stylet, 20-26 /um. Amongthose species that lack a vulval flap, B. huna-nensis appears to be closest in spicule shape toB. eggersi, from which it differs due to shorterfemale body length (580-790 vs. 990-1,122 ^m)and more posteriorly placed vulva (76-80 vs. 69-70%). The spicules of B. hunanensis also resem-ble those of B. talonus, B. tritrunculus, and B.teratospicularis. The new species differs mark-edly from the first 2 species by its greater styletlength as well as the more posteriorly positionedvulva (71-73 vs. 76-80%). Bursaphelenchushunanensis appears close to B. teratospicularisin all essential measurements except "c" ratio.



The 2 species can be further differentiated by Female tail terminus mostly broadlyvariances in the spicules. In B. hunanensis, the rounded, rarely with digitate ter-rostrum is more elongate-digitate and the lamina minus xylophilus (Steiner andis more symmetrically rounded than for B. ter- Buhrer, 1934) Nickle, 19704

atospicularis. (syn.: lignicolus Mamiya andTYPE DESIGNATIONS: HOLOTYPE: Female, Kiyohara, 1972)

collected by Mrs. Qiu Shouwei, October 1984. 3(2). Female tail tapering to a pointed ter-Specimen number: Bursaphelenchus hunanensis minus, female "c" ratio 12-17holotype 1, Nematode Collection, Department borealis Korentchenko, 1980of Plant Protection, South China Agricultural Female tail definitely mucronate, fe-University, Guangzhou, China. male "c" ratio 20-30 4

ALLOTYPE: Male, same data as holotype. 4(3). Male spicule 17-21 nm long, vulvaSpecimen number: Bursaphelenchus hunanensis 68-75%allotype 1. kolymensis Korentchenko, 1980

PARATYPES: Same data as holotype and al- Male spicule 21-29 jum long, vulvalotype. One male and female para type, slide 73-77% 5T-3794p, USDA Nematode Collection, Belts- 5(4). Female stylet 12-14 ^m long, maleville, Maryland, U.S.A.; and slide 86c/8/l, Nem- spicule 21-22 /urn longatode Collection, Rothamsted Experimental fraudulentus Riihm, 1956Station, Harpenden, Herts, England. Five female Female stylet 14-16 /^m long, maleand 4 male paratypes in Nematode Collection, spicule 23-29 /zm longDepartment of Plant Protection, South China mucronatus Mamiya andAgricultural University, Guangzhou, China. Enda, 19795

TYPE HABITAT : Dead wood of Pinus masso- 6(1). Female stylet 20-26 /urn longniana Lamb. hunanensis sp. n.

TYPE LOCALITY: Hongjiang, Huaihua Coun- Female stylet shorter than 20 pmty, Hunan Province, China. (teratospicularis female stylet 18-

22 fjan) 7Definition and Current Status of the Genus y(6) Spicular lamina with a blunt-round

Bursaphelenchus ^D g

Since the review of the genus Bursaphelenchus Tip of spicular lamina more acute toby Tarjan and Baeza-Aragon (1982), 6 new finely rounded 11species have been described and 1 species has 8(7). Spicular rostrum with acute tip; malebeen transferred into the genus, making a total "a" ratio 19-28; female lengthof 49 recognized species (Table 1). 390-830 /nm hunti (Steiner, 1935)

Kakuliya et al. (1980) referred to Bursaphe- Giblin and Kaya, 1983lenchus populneus and 5. tbilisensis and Vosilite Tip of spicular rostrum digitate to(1980) to B. sexdentai which, to the best of our almost flattened; male "a" ratioknowledge, are nomina nuda and are not in- 27-45; female length 690-1,350eluded in the key or in Tables 1 and 2. jum 9

The following key to valid Bursaphelenchus 9(8). Spicule 13 ̂ m long, spicular rostrumspp. lists synonymies which were proposed by almost flattenedTarjan and Baeza-Aragon (1982). It does not list gonzalezi Loof, 1964B. conjunctus (Fuchs, 1930) and B. conurus Spicule 18-27 jum long, spicular ros-(Steiner, 1932) which were deemed species in- trum digitate 10certae sedis, B. cryphali (Fuchs, 1930) placed inspecies inquirendae, and B. ruehmi Baker, 1962 / ,no i\ A c. A- D / :,-... . . 4 Wingfield et al. (1983) reported finding B. xylophi-which was regarded as species indeterminate. lus in balsam fir which had digitate tails and which

would mate with B. xylophilus from pine, but not withKey to Bursaphelenchus Species B mucronatus Mamiya and Enda, 1979.

1. Spicule with cucullus (cf. Fig. 12) 2 5 Baujard (1980) considered B. mucronatus a syn-c . , . . , . , , £ onym of B. lignicolus Mamiya and Kiyohara, 1972Spicule without cucullus 6 (=R xylophilu\t de Guiran et al (1985) demon.

2(1). Female tail always with digitate or strated distinct differences on the basis of enzyme elec-pointed terminus 3 trophoresis.



Table 1. Diagnostic data on Bursaphelenchus females: essential measurements and ratios.

Bursaphelenchusspecies

baker/bestiolusborealischitwoodicorneoluscrenaticryphalidigituluseggersieidmanniel viruseremuseucarpusfraudulentusfungivorusgeorgicusgonzalezihunanensis sp. n.huntiidiusincurvuskevinikolymensisleonilignicoluslignophilusmucronatusnaujacinewmexicanusnuesslinipinasteripiniperdaepityogenipoligraphiratzeburgiisachsiscolytiseanisilvestrissteinerisutoricussychnustalonusteratospicularistritrunculusvaricaudawilfordixerokarterusxylophilus

Body length(Mm)

924-1,232830

507-1,018619

650-700825-1,125

447790-1,260990-1,122795-990890-960858-990825-1,275675-900770-1,160

656690-1,100580-790390-830:):610-885660-840810-980473-716580-860710-1,010992-1,257700-980730-1,300

1,500660-1,020550-650

551§940

605-9 14§720-825660-763

840770-1,350720-1,220456^75

930634-662

800580-675

650710-890

900600-930447-6091

"a" ratio

38-4132

26-382129

39-5419

31-4437-3925-3235 1̂

4139-46

3228 3̂

3325-3631-4121-2932-3424-2526-2922-3633^t433^t646-5536-4635-51

4022-3635 7̂

4030

41-5739-41

3440

25-3235^*722-26

4423-29

3329-34

2926-36

6026-3837-48

"c" ratio

2617

12-171618

26-3219

13-2020

70-7 3f17-21

2320-2621-2610-15

2012-1914-199-13

23-2519-2217-1917-2410-1523-3220-2520-3019-30

2824-2919-25

272645

21-2214-15

2014-1911-15

925

11-1225

22-2311

15-2527

20-3023-31

Stylet lengthGum)

18-1913-1414-16

1212*1112

11-1416-1814-18

1513

12-1412-1414-16

1213-1620-2612-1411-1415-1915-1712-1512-1714-1811-1414-1613-18

1512-1511-13

1214-15

911

12-1411

15-1912-1612-14

1312-13

1118-22

1312-17

14*12-15

13

Vulval %

73-7574

71-776973

72-7471

84-8869-7072-78

7369-7368-7674-7567-73

8271-7676-8067-8376-8072-7376-8168-7569-7467-7870-7373-7771-76

7371-7669-73

767376

71-7471-72

7276-8272-7668-73

6473-74

7375-79

7173-82

70*73-8073-77

Vulvalflap

9

+

+7+9-----79

+-----9

-

-

+

+

+

?

+

7+9

+

-

+

7-7--+9

-

9

?

9

9

+

-?

+

* Measured from illustration.t An obvious error.$ Measurements of lectotype material by Nickle, 1970.§ Measurements from Fuchs, 1937.|| Measurements of lectotype material by Nickle ct al., 1981.



Table 2. Diagnostic data on Bursaphelenchus males: essential measurements and ratios.

Bursaphelenchusspecies

bakeribestiolusborealischitwoodiconjunctuscorneoluscrenaticryphalidigituluseggersieidmanniclytruseremuseucarpusfraudulentlyfungivorusgeorgicusgonzalezihunanensis sp. n.huntiidiusincurvuskevinikolymensisleonilignicoluslignophilusmucronatusnaujacinewmexicanusnuesslinipinasteripiniperdaepityogenipoligraphiratzeburgiisachsiscolytiseanisilvestrissteinerisutoricussychnustalonusteratospicularistritrunculusvaricaudawilfordixerokarterusxylophilus

Body length(Mm)

858-990670

548-813547455

570-700825-1,080885-945f530-710450-990675-855830-1,040782-840750-1,020645-685570-1,030

720580-970480-590360-740:):619-643600-645890-1,120423-706510-1,060590-820912-1,147640-970630-1,200

1,250780-900500-620

570f820

589-785f630-675662-691

800660-1,290730-895476-480

740490-547

800450-490

650540-840670-750680-825520-60 1§

"a" ratio

38-4532

25-43343135

39-5151-5433-4121-3132-3941-5240-4943^*933-3731^t5

3930-3838-4519-28

3421-2427-3624-4636-5636-4748-6439-5132-52

4743-4542-51

3831

35-6538-3926-29

3827-3337-4423-26

3424-30

4728-30

2930-41

6131-4135^t5

"c" ratio

25-2618

16-29241518

29 2̂25-3715-1919-2819-2321-2631-3633^t520-2718-25

2421-2915-1918-21

3324-2615-1918-2916-2621-3118-2326-3620-27

2628-3220-25

24223428

25-2725

18-2718-22

2114

22-2425

23-3323

15-2326

24-3421-29

Stylet length(urn)

1813-1412-161216.512*11

12-1411-1214-1614-1815

11-1212-1412-1514-15

9

12-1519-2112-1311

12-1714-1612-1513-1714-1711-1414-1613-181515

11-1312

14-15109-11

12-1411

13-1812-151214

12-1411

18-2213

15-1714*

12-1813-14

Spicule length0*m)

19-2218*

16-201614.513*

17-1814-1516*

18-2418-2128*

14-1915-1621-2218*1913*

13-2016-1714

17-1818-2217-2110-2025-3021-2323-2913-2024*15-1612-141417*

13-1411-1217-18

7*18-2728-31

172219-2315*

15-1814*

13-1710*

14-1519-23

* Measured from illustration.t Measurements from Fuchs, 1937.£ Measurements of lectotype material by Nickle, 1970.§ Measurements of lectotype material by Nickle et al., 1981.



22(18).

23(22).

24(21).

10(9). Vulva 67-73%, male "a" ratio 31-45 fungivorus Franklin and

Hooper, 1962 20(19).Vulva 76-82%, male "a" ratio 27-

33 seani Giblin and Kaya, 198311 (7). Female length 456-475 /um, "c" mea-

surement 8.6-8.8, spicule thin-elongate steineri (Riihm, 1956)

Goodey, 1960Female length 580 /um and greater, 21(20).

"c" measurement 10 or more,spicule more stout 12

12(11). Male spicule 26-31 /um long 13Male spicule 24 /um or less in length

1513(12). Female without vulva flap, "c" ra-

tio = 17-21 elytrus Massey, 1960Female with vulval flap, "c" ratio

either more than 21 or less than17 14

14(13). Condylus (cf. Fig. 12) of spicular ca-pitulum with a conspicuous pos-terior flexure, male length 730-895/um silvestris (Lieutier and

Laumond, 1978) Baujard, 1980Condylus of spicule without poste-

rior flexure, male length 1,250 /umnewmexicanus Massey, 1974

15(12). Vulva 82-88% 16Vulva 78% or less (varicauda vulva

73-82%) 1716(15). Spicule relatively slender in width,

condylus digitate, male "c" ratio11-12 digitulus Loof, 1964

Spicule relatively broader in width,condylus broadly conical, male "c"ratio 24

georgicus Devdariani, Kakuliya,and Khavtasi, 1980

17( 15). Condylus of spicular capitulum witha posterior flexure 27(25).

leoni Baujard, 1980Condylus of spicule without poste-

rior flexure 18 28(27).18(17). Spicule length (measured along arc)

11-13 /um or less 19Spicule length 14 /um or greater (var-

icauda spicule 13-17 /um) 2219(18). Female with vulva flap, length 650- 29(27).

700 /um long; "c" value of male18.5 corneolus Massey, 1966

Female without vulva flap, length 30(29).

25(22).

26(25).

720 /um or greater; "c" value ofmales 21-29 20

Female stylet 12-16 /um long, spic-ular laminae strikingly widened or"mitten-shaped" at midpoint

wilfordi Massey, 1964Female stylet 9-11 /um long, spicular

laminae not excessively widened21

Male length 800 /um, spicule 7 /umlong scolyti Massey, 1964

Male length 630-675 /um, spicule 11-12 /um long

ratzeburgii (Riihm, 1956)Goodey, 1960

Females with vulval flap 23Females without vulval flap, or

presence of flap not determined 25Male stylet 15-17 /um long, spicule

length 13-17 /umvaricauda Thong and

Webster, 1983Male stylet 13-15 /um long, spicule

length 17-18 /um 24Lateral incisures present, male length

820 /um, female length 940 /umpityogeni Massey, 1974

Lateral incisures absent, male length670 /um, female length 830 /um

bestiolus Massey, 1974Male length 450-547 /um (eggersi

male length 450-990 /um) 26Male length 547 /um or more 27Male stylet 12-14 /um long, vulva

73-74%, female "c" ratio 11-12sychnus (Riihm, 1956)

Goodey, 1960Male stylet 18-22 /um long, vulva

75-79%, female "c" ratio 22-23teratospicularis Kakuliya and

Devdariani, 1965Male stylet 18-19 /on 28Male stylet 18 /um or less, average

length 16 /um or less 29Female stylet 18-19 /um, vulva 72-

74% bakeriRuhm, 1964(syn.: naujaci Baujard, 1980)

Female stylet 16-18 /um vulva 75-76% piniperdae Fuchs, 1937

Spicule length 18-24 /um 30Spicule length no greater than 18 /um

33Stylet length of both sexes 11-14 /um,



bakeri bestiolus borealis chitwoodi corneolus crenati

fraudu/entus fungivorus georgicus gonzalezi hunanensis hunt!

mucronatus naujaci newmexicanus nuesslini pinaster! piniperdae

silvestris steineri sutoricus sychnus ta/onus teratospicularis

Figure 13. Spicules of Bursaphelenchus spp. (partially after Tarjan and Baeza-Aragon, 1982).


OF WASHINGTON, VOLUME 55, NUMBER 1, JANUARY 1988 •

digitulus eggersi eidmanni elytrus eremus eucarpus

idius incurvus kevini kolymensis ^eoni lignophilus

,m

pityogeni polygraph! ratzeburgii sachsi scolyti seani

tritrunculus varicauda wilfordi xerokarterus xylophilus

Figure 13. Continued.


10 PROCEEDINGS OF THE HELMINTHOLOGICAL SOCIETY

"a" ratio 48 or morelignophilus (Korner, 1954)

Meyl, 1960Stylet length of both sexes 14 /urn or

greater, "a" ratio 39 or less 3131(30). Female length 991-1,122 /um, vul-

va 69-70%, female "a" ratio 37-39 eggersi (Riihm, 1956)

Goodey, 1960Female length 990 /um or less, vulva

72-81%, female "a" ratio 26-32 32

32(31). Male length 675-855 /urn; spiculenarrow with thornlike pointedrostrum eidmanni (Riihm, 1956)

Goodey, 1956Male length 890-1,120 /urn, spicule

broad with narrow digitate ros-trum kevini Giblin, Swan,

andKaya, 198433(29). Male length 547 /um, female length

619 Mm chitwoodi (Riihm, 1956)Goodey, 1960

(syn.: pinasteri Baujard, 1980)Male length 570 /um and greater, fe-

male length 650 /um or greater (xe-rokarterus range 600-903 /um,idius range 610-885 /um) 34

34(33). Spicule length 14-15 yum (exc. poli-graphi spicule 13-18 /um) 35

Spicule length 15 /um or more (ere-mus range 14-19 /um) 38

35(34). Male length 650 /um, vulva 71%tritrunculus Massey, 1974

Male length greater than 680 /um orless than 643 /um, vulva 73% ormore 36

36(35). Condylus of spicule a reduced pro-tuberance, male length 619-643/urn

idius (Riihm, 1956) Goodey, 1960Condylus of spicule prominent, male

length 680-825 /um 3737(36). Vulva 73-80%, stylet length both

sexes 12-18 /^mxerokarterus (Riihm, 1956)

Goodey, 1960(syn.: sutoricus Devdariani, 1974)

Vulva 73%, stylet length both sexes11 /um talonus (Thorne, 1935)

Goodey, 196038(34). Rostrum of capitulum sharply

pointed 3 9Rostrum of capitulum digitate 42

39(38). Condylus rectangular to rectangu-lar-elongate 40

Condylus digitate to broadly digitate41

40(39). Female stylet 12-14 /urn long, malelength 750-1,020 /um, male "c"value 3 3-4 5

eucarpus (Riihm, 1956)Goodey, 1960

Female stylet 15-19 /um long, malelength 600-645 /um, male "c" val-ue 24-26 incurvus (Riihrn, 1956)

Goodey, 196041(39). Male length 780-900 /um, "c" value

of female 24-29, spicular laminaewith symmetrically curved dorsalsurface nuesslini (Riihm, 1956)

Goodey, 1960Male length 662-691 um, "c" value

of female 14-15, spicular laminaewith dorsal surface forming an 80°angle sachsi (Riihm, 1956)

Goodey, 196042(38). Female stylet 9-11 /um long, female

"c" value at least 26, spicular ca-pitulum slightly depressed 43

Female stylet 13 /um long, female "c"value 23, spicular capitulummoderately depressed

eremus (Riihm, 1956)Goodey, 1960

43. Female stylet 9 /um long, male length589-785/um, spicule length 13-14/urn poligraphi Fuchs, 1937

Female stylet 11 /um long, malelength 825-1,080 yum, spiculelength 17-18 /um

crenati (Riihm, 1956)Goodey, 1960

The essential measurements and ratios (Tables1, 2) and spicule shapes found within the species(Fig. 13) which follow are an amplification ofthose presented in Tarjan and Baeza-Aragon(1982), which are brought up to date with theinclusion of the 6 recently described species andadditional data on the older species. It is incum-bent upon the authors to caution potential usersthat several of the species are poorly describedand lacking essential data. Almost no mentionof nematode variation is made, hence the keyand diagnostic data must be relied upon onlywith adequate reservation. Spicule shape has beenassumed to exhibit little, if any, variation. Be-


OF WASHINGTON, VOLUME 55, NUMBER 1, JANUARY 1988 11

cause of the paucity of differentiating morpho-logical characters between species, the shape ofthe spicule probably is the most visible featurecharacterizing species. It is unfortunate that vari-ation in spicule shape and other morphologicalfeatures have not yet been extensively investi-gated, except for some of the studies conductedby Giblin and Kaya (1983) and Giblin et al.(1984). When adult variation is intensively stud-ied, the genus may possibly be reduced to halfits present species.

Acknowledgments

We are grateful to Professor Jiang Jingbo ofthe Biology Department of Zhongshan Univer-sity (Sun Yat-Sen University) for his review ofan earlier version of this manuscript and Drs. A.M. Golden and Zafar Handoo of the U.S. De-partment of Agriculture, Belts ville, Maryland andDr. R. M. Giblin-Davis, Fort Lauderdale REC,University of Florida for supplying photocopiesof references.

Literature Cited

Baujard, P. 1980. Three new species of Bursaphelen-chus (Nematoda: Tylenchida) and observations onthe genus. Revue de Nematologie 3:167-177.

de Guiran, D., M. J. Lee, A. Daltnasso, and M. Bon-giovanni. 1985. Preliminary attempt to differ-entiate pinewood nematodes (Bursaphelenchusspp.) by enzyme electrophoresis. Revue de Ne-matologie 8:88-90.

Devdariani, T. G., G. A. Kakuliya, and D. D. Khavtasi.1980. A new species of nematode from the smallmaple Capricorn beetle, Rhopalopus macropus.Soobshcheniya Akademii Nauk Gruzinskoi SSR98(2):457-459.

Fuchs, A. G. 1937. Neue parasitische und halbpar-asitische Nematoden bei Borkenkafern und einigeandere Nematoden. I. Teil die Parasiten der Wald-gartner Myelophilus piniperda L. und minor Har-tig und die Genera Rhabditis Dujardin, 1845 undAphelenchus Bastian, 1865. Zoologische Jahr-biicher Abteilung fur Systematik Oekologie undGeographic der Tiere. Jena 70(5-6):291-380.

Giblin, R. M., and H. K. Kaya. 1983. Bursaphelen-chus seani n. sp. (Nematoda: Aphelenchoididae),

a phoretic associate of Anthophora bomboidesstanfordiana Cockerell, 1904 (Hymenoptera: An-thophoridae). Revue de Nematologie 6(1):39-50.

-, J. L. Swan, and H. K. Kaya. 1984. Bursa-phelenchus kevini n. sp. (Aphelenchida: Aphelen-choididae), an associate of bees in the genus Ha-lictus (Hymenoptera: Halictidae). Revue deNematologie 7(2): 177-187.

Kakuliya, G. A., and T. G. Devdariani. 1965. Anewspecies of nematode Bursaphelenchus teratospic-ularis Kakuliya and Devdariani, sp. nov. (Nem-atoda: Aphelenchoidea). (Georgian text). Bulletinof the Academy of Sciences of the Georgian SSR38(1):187-191.

, , and L. K. Maglakelidze. 1980.Nematodes of Cerambycidae, parasites of trees inEastern Georgia. (Georgian text.) Pages 109-110in Konferentsiya Ukrainskogo Parazitdogiches-kogo Obshchestra Tezisy doklador Chast 2. "Nau-kova Dunka," Kiev, USSR.

Mamiya, Y. 1972. Pine wood nematode, Bursaphe-lenchus lignicolus Mamiya and Kiyohara, as acausal agent of pine wilting disease. Review ofPlant Protection Research 5:46-60.

Nickle, W. R. 1970. A taxonomic review of the gen-era of the Aphelenchoidea (Fuchs, 1937) Thorne,1949 (Nematoda: Tylenchida). Journal of Nema-tology 2:375-392.

, A. M. Golden, W. Mamiya, and W. P. Wergin.1981. On the taxonomy and morphology of thepine wood nematode, Bursaphelenchus xylophilus(Steiner and Burher 1934) Nickle, 1970. JournalofNematology 13:385-392.

Riihm, W. 1956. Die Nematoden der Ipiden. Par-asitologische Schriftenreihe. Jena 6:1-435.

Tarjan, A. C., and C. Baeza-Aragon. 1982. An anal-ysis of the genus Bursaphelenchus Fuchs, 1937.Nematropica 12:121-144.

Thong, C. H. S., and J. M. Webster. 1983. Nematodeparasites and associates of Dendroctonus spp. andTrypodendron lineatum (Coleoptera: Scolytidae),with a description of Bursaphelenchus varicaudan. sp. Journal ofNematology 15:312-318.

Vosilite, B. S. (Vosylyte, B.). 1980. A new species ofnematode, Diplogasteroides sexdentati n. sp. andnotes on the biology of ecto-nematodes of Ipssexdentatus. (Russian text.) Pages 21-28 inGel'minty nasekomykh. "Nauka," Moscow,USSR.

Wingfield, M. J., A. Blanchette, and E. Kondo. 1983.Comparison of the pine wood nematode, Bursa-phelenchus xylophilus from pine and balsam fir .European Journal of Forest Pathology 13:360-373.


A Key to Species in the Genus Bursaphelenchus with a ...DIAGNOSIS: Bursaphelenchus hunanensis sp. n....

Documents

Transcript of A Key to Species in the Genus Bursaphelenchus with a ...DIAGNOSIS: Bursaphelenchus hunanensis sp. n....