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

A Ke y t o Specie s in th e Genu s Bursaphelenchus with a Descriptio n of

Bursaphelenchm hunanensis sp . n . (Nematoda : Aphelenchoididae )

Found in Pin e Woo d in Huna n Province , China 1

KAMCHEW YiN, 2 YUSHEN G FANG, 2 AND A . C . TARjAN 3

2 Plan t Nematolog y Researc h Laboratory , Departmen t of Plan t Protection ,

Sout h Chin a Agricultura l University , Guangzhou , Chin a

and 3 Departmen t of Entomolog y and Nematology , IFAS, Universit y of Florida , Gainesville , Florida 32611

ABSTRACT : Bursaphelenchus hunanensis sp. n . i s describe d an d illustrated . I t ha s a distinc t style t longer than

othe r specie s

in th e genu s and ha s n o basa l thickenings . The femal e lack s a vulva l flap an d ha s a shor t postuterin e

branch .

The mal e spicule s do no t hav e a cucullu s and terminat e posteria d in a blun t rounde d tip. The rostru m

is digitate . An alphabetica l lis t of al l 4 9 nomina l specie s in th e genus , a ke y t o 4 4 vali d species , and table s of

diagnosti c dat a of male s and female s are als o included . KEY WORDS : Nematoda , Aphelenchoididae , Bursaphelenchus hunanensis sp. n., nematod e taxonom y and

systematics , Pinus massoniana, key t o species .

Species withi n the genu s Bursaphelenchus

Fuchs ,

1937 , were regarde d onl y as associate s of

bar k beetle s (Ipidae ) unti l the insidiou s pine -

wood nematod e B. xylophilus (Steine r and Buhr -

er, 1934 ) Nickle , 1970, starte d to devastat e entir e

forest s in Japa n (Mamiya , 1972) . Curren t con -

cer n

on chanc e introductio n of thi s specie s int o

new area s has renewe d interes t in th e genu s and

its species .

A Bursaphelenchus species was firs t foun d in

Octobe r 198 4 in th e woo d of dea d pin e trees ,

Pinus massoniana Lamb,

in Huaihu a County,

Huna n Province , China , durin g

a surve y for th e

pin e wil t diseas e cause d by Bursaphelenchus xy -

lophilus. Detailed studies of th e nematod e re-

veale d tha t it differe d from the 4 3 specie s liste d

in th e generi c review by Tarja n and Baeza-Ara -

gon (1982 ) and 5 additiona l nomina l specie s

(Kakuliy a and Devdariani , 1965; Devdarian i et

al. , 1980 ; Gibli n and Kaya, 1983; Thon g and

Webster , 1983 ; Gibli n et al. , 1984) .

The purpos e of thi s paper , in additio n to re -

portin g a ne w species , is t o provid e a ke y an d a

tabl e of essentia l measurement s and ratio s of

nomina l specie s withi n the genus .

Material s and Method s

Bursaphelenchus specimens wer e extracte d by th e Baerman n funnel metho d from woo d fragment s cut from wilte d trees . Studie s wer e mad e on livin g as wel l as fixe d specimens . Measurement s and drawing s wer e

mad e from specimen s fixed in TA F (triethanolamine ,

1 Florida Agricultura l Experimen t Statio n Journa l

Serie s No. 7922 .

formaldehyde , and water ) and staine d wit h poly -

chrom e blue . Spicul e length s wer e measure d alon g the media n line of th e ar c o f th e spicule . The specimen s were mounte d in glycerin . Measurement s are i n mi - crometer s unles s state d otherwise .Informatio n on othe r Bursaphelenchus species was obtaine d solel y from perusa l of pertinen t references . Drawings of spicule s of eac h specie s wer e mad e fro m the origina l publication , and the n eithe r enlarge d or reduce d to a standar d size . The majorit y of th e spicul e drawing s shown in Figur e 13 cam e from Tarja n and Baeza-Arago n (1982).

Generic Diagnosi s

Ther e hav e bee n

no emendation s proposed b y

other s to th e generi c diagnosi s of th e genu s give n

by Nickl e (1970 ) which is presente d below :

Bursaphelenchus Fuchs , 193 7

Syn. Aphelenchoides (Bursaphelenchus)

(Fuchs , 1937 ) Riihm , 1956

DEFINITION : Aphelenchoididae . Usuall y long .

Lip s high ,

offset . Stylet wit h smal l rounde d basal

knobs . Excretor y pore usuall y behind media n

bulb. Vulva l lip s sometime s protruding ; post -

uterin e sac usuall y long . Femal e tai l rounded ,

conoid ,

or sharpl y pointed . Spicules large, nar-

row, usually with prominent rostrum. Male tail

strongly arcuate, variously pointed, with short,

terminal caudal

alae. Usually 2 pairs of cauda l

papillae ,

1 adanal, the othe r postanal . Guber-

naculum absent .

BIONOMICS : Mycophagous , insec t associate .

TYPE SPECIES : Bursaphelenchus piniperdae

Fuchs, 1937 .

1

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50 ju m

1,2, 3

4,5,6,7,8,9,1 0 10 >u m

10 jam

11

1 0

11

Figure s 1-11. Bursaphelenchus hunanensis sp. n . 1. Female . 2. Male . 3. Posterio r par t of mal e showin g flexure of the anterio r end of the testis . 4. Femal e anterio r end . 5. Vulv a region . 6. Female tail. 7. Latera l view of cuticl e in midbod y region. 8. E n fac e vie w of head . 9. Latera l view of mal e tail. 10. Ventra l view of mal e tail. 11. Mal e spicule .

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OF WASHINGTON , VOLUM E 55, NUMBE R 1, JANUAR Y 1988

Result s and Discussio n

Bursaphelenchus hunanensis sp. n .

(Figs . 1-11 )

FEMALE S (30) (paratypes) : L = 0.6 7 (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) .

FEMAL E (holotype) : L = 0.6 7 mm ; a = 36.4 ;

b = 7.7 ; c = 17 ; V = 79.2% ; style t = 23 .

MAL E (allotype) : L = 0.5 2 mm ; a = 39.4 ;

b = 6.6; c = 16.4; T = 26.3% ; style t = 19 ; spic -

ule = 15 .

FEMALE : Body ventrall y curve d whe n kille d

wit h heat , taperin g graduall y nea r extremities .

Cuticl e marke d

by fin e annules , abou t 1 nm wide

in midbod y region, interrupte d by a narro w lat -

era l

fiel d whic h is abou t '/10 o f th e maximu m bod y

widt h

and ha s 2 longitudinal , refractiv e line s on

eithe r sid e of a narro w refractiv e ban d (Fig . 1).

Lip flattene d a t ora l opening , set of f b y a con -

striction .

En fac e view of hea d showin g 6 equal

lip s (Fig .

8). Anterio r style t con e les s tha n '/2 tota l

style t length , posterio r

shaft withou t basa l thick -

enin g (Fig .

4). Procorpus cylindrical , twic e as lon g

as vulva l bod y width . Media n bul b elongate-oval ,

wit h conspicuou s valv e jus t posterio r

to center .

Excretor y

pore Vi to 1 lengt h of media n bulb

behin d the bul b on a leve l with , or jus t behind ,

the nerv e ring . Esophageal glands dorsall y over -

lappin g intestine ,

9 times as lon g as vulva l bod y

width . Reproductiv e syste m prodelphic , gona d

outstretched , occupyin g

Vs the bod y length . Oo-

cyte s arrange d in singl e file except at th e anterio r

par t

of th e ovary . Vagin a an anteriorl y incline d

tube , withou t vulva l

flap. Postuterin e branc h

usually shorte r tha n vulv a bod y diamete r (Fig .

5). Vulva-anu s distanc e equa l to 2. 5 time s tai l

length .

Tail conoi d with sligh t ventra l curvature ,

3 times longe r tha n ana l bod y width , wit h a finel y

rounde d terminu s (Fig . 6).

MALE : Body J-shape d whe n kille d wit h heat .

Anterio r bod y regio n simila r

to female . Repro-

ductiv e syste m

monorchic , testis usuall y out -

stretched , extendin g

1A of bod y length . Approx -

imatel y

V& of th e testi s reflexe d anteriorl y (Fig .

3). Spermatogoni a arranged in severa l row s and

spermatocyte s in singl e file. Semina l vesicle oval.

Vas deferen s compose d of anterio r cylindrica l

part and posterio r slende r duct . Spicule s paired ,

CONDYLU S

CAPITULUM

ROSTRUM

CALOMU S

LAMINA

CUCULLU S

CAUDAL ALA

(BURSA )

Figure 12. Latera l view of th e Bursaphelenchus spicule s (after Tarja n and Baeza-Aragon , 1982).

arcuate , wit h blun t prominen t rostrum ; dista l end

of spicul e obtuse , withou t cucullu s (cf . Fig . 2).

Cente r of capitulu m depressed , condylu s hemi-

sphericall y rounded. Tail ventrall y arcuate , ter -

minu s

finel y rounded , wit h shor t terminall y

truncate d burs a (Fig . 10) .

Two pair s of cauda l

papilla e present ;

1 adanal pair , 1 pair abou t Vi

tail lengt h from tail terminus .

DIAGNOSIS : Bursaphelenchus hunanensis sp.

n. differ s from all othe r specie s in th e genu s by

the lengt h of femal e stylet, 20-26 /um . Amon g

thos e specie s tha t lac k

a vulva l flap, B . huna-

nensis

appear s to b e closes t in spicul e shape to

B. eggersi, fro m whic h it differ s due t o shorte r

female body lengt h (580-79 0 vs. 990-1,12 2 ^m)

and mor e posteriorl y place d vulv a (76-8 0 vs. 69 -

70%) . The spicule s of B . hunanensis also resem -

ble thos e of B . talonus, B . tritrunculus, an d B .

teratospicularis. Th e ne w specie s differ s mark-

edly fro m the firs t 2 specie s by it s greate r style t

length as wel l as th e mor e posteriorl y positione d

vulv a

(71-7 3 vs. 76-80%) . Bursaphelenchus

hunanensis appears clos e

to B . teratospicularis

in al l essentia l measurement s excep t "c" ratio .

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The 2 specie s can b e furthe r differentiate d by Femal e tai l terminu s mostl y broadl y

variance s

in th e spicules . In B . hunanensis, th e rounded , rarel y wit h digitat e ter -

rostru m is mor e elongate-digitat e and th e lamin a minu s xylophilus (Steiner and

is mor e symmetricall y rounde d tha n for B . ter- Buhrer , 1934 ) Nickle , 1970 4

atospicularis. (syn. : lignicolus Mamiya and

TYP E DESIGNATIONS : HOLOTYPE : Female, Kiyohara , 1972)

collecte d

by Mrs . Qiu Shouwei , Octobe r 1984 . 3(2) . Female tai l taperin g to a pointe d ter -

Specime n number :

Bursaphelenchus hunanensis minus , femal e "c" rati o 12-1 7

holotyp e 1, Nematod e Collection , Departmen t borealis Korentchenko , 198 0

of Plan t Protection , Sout h China Agricultura l Femal e tai l definitel y mucronate , fe-

University , Guangzhou , China . mal e "c" rati o 20-3 0 4

ALLOTYPE : Male, sam e dat a as holotype . 4(3). Male spicul e 17-21 nm long , vulv a

Specime n number : Bursaphelenchus hunanensis 68-75 %

allotype 1. kolymensis Korentchenko, 198 0

PARATYPES : Same dat a as holotyp e and al- Mal e spicul e 21-29 jum long , vulv a

lotype . One mal e and femal e para type , slid e 73-77 % 5

T-3794p, USD A Nematod e Collection , Belts - 5(4) . Female style t 12-1 4 ^m long , mal e

ville, Maryland , U.S.A.; and slid e 86c/8/l , Nem- spicul e 21-22 /urn lon g

atod e Collection , Rothamste d Experimenta l fraudulentus Riihm, 1956

Station , Harpenden , Herts , England . Fiv e

femal e Femal e style t 14-16 /^m long , mal e

and 4 mal e paratype s in Nematod e Collection , spicul e 23-29 /zm lon g

Departmen t

of Plan t Protection , Sout h Chin a mucronatus Mamiy a and

Agricultura l University , Guangzhou , China . Enda , 1979 5

TYPE HABITAT : Dead woo d of Pinus masso- 6(1) . Female style t 20-26 /urn lon g

niana Lamb . hunanensis

sp . n .

TYP E LOCALITY: Hongjiang , Huaihua Coun - Femal e style t shorte r tha n 20 pm

ty, Huna n Province , China . (teratospicularis femal e style t 18-

22 fjan) 7 Definitio n and Curren t Status of th e Genu s y(6 ) S picular lamin a wit h a blunt-roun d Bursaphelenchus ^ D g

Since the revie w of th e genu s Bursaphelenchus Ti p o f spicula r lamin a mor e acut e to

b y Tarja n and Baeza-Arago n (1982), 6 ne w finel y rounde d 1 1

species hav e bee n describe d and 1 specie s has 8(7). Spicula r rostru m wit h acut e tip ; mal e

bee n transferre d int o

the genus , makin g a total "a " ratio 19-28 ; femal e lengt h

of 4 9 recognize d specie s (Tabl e 1). 390-83 0 /nm hunti (Steiner , 1935 )

Kakuliy a et al . (1980 ) referre d to Bursaphe- Gibli n and Kaya , 198 3

lenchus populneus and 5. tbilisensis an d Vosilit e Ti p o f spicula r rostru m digitat e to

(1980 ) to B . sexdentai which , to th e bes t of ou r almos t flattened ; mal e "a" rati o

knowledge , are nomin a nud a and ar e no t in - 27-45; femal e lengt h 690-1,35 0

eluded in th e ke y o r i n Table s 1 an d 2 . ju m 9

The followin g key t o vali d Bursaphelenchus 9(8) . Spicule 13 ^m long , spicula r rostrum

spp . list s synonymie s whic h wer e

propose d by almos t flattened

Tarjan and Baeza-Arago n (1982). It doe s not list gonzalezi Loof , 1964

B. conjunctus (Fuchs, 1930 ) and B. conurus Spicul e 18-2 7 jum long , spicula r ros-

(Steiner ,

1932) which wer e deeme d specie s in- tru m digitate 1 0

certae sedis , B. cryphali (Fuchs , 1930 ) place d in

specie s inquirendae , and B. ruehmi Baker , 196 2 /,noi \ A c. A- D / :,- ... . . 4 Wingfield et al . (1983 ) reporte d finding B . xylophi- whic h was regarde d as specie s indeterminate . lus in balsa m fir whic h had digitat e tails and whic h

woul d mate wit h B. xylophilus fro m pine , but no t wit h Key to Bursaphelenchus Specie s B muc ronatus Mamiy a and Enda , 1979 .

1. Spicul e with cucullu s (cf. Fig . 12) 2 5 Baujar d (1980) considere d B. mucronatus a syn - c .,..,., , £ onym of B . lignicolus Mamiya and Kiyohara , 197 2 Spicul e withou t cucullu s 6 (=R xylophilu de Guira n et al (1985 ) demon .

2(1). Femal e tail alway s with digitat e or strate d distinct difference s on th e basi s of enzym e elec- pointe d terminu s 3 trophoresis .

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OF WASHINGTON , VOLUM E 55, NUMBE R 1, JANUAR Y 1988

Table 1. Diagnosti c dat a on Bursaphelenchus females : essentia l measurement s and ratios .

Bursaphelenchus species

baker/bestiolusborealischitwoodicorneoluscrenaticryphalidigituluseggersieidmanniel viruseremuseucarpusfraudulentusfungivorusgeorgicusgonzalezihunanensis sp. n. huntiidiusincurvuskevinikolymensisleonilignicoluslignophilusmucronatusnaujacinewmexicanusnuesslinipinasteripiniperdaepityogenipoligraphiratzeburgiisachsiscolytiseanisilvestrissteineri

sutoricus

sychnustalonusteratospicularistritrunculusvaricaudawilfordixerokarterusxylophilus

Bod y lengt h(Mm)

924-1,23 2 830 507-1,01 8 619 650-70 0825-1,12 5 447 790-1,26 0990-1,12 2795-990890-960858-99 0825-1,27 5675-900770-1,16 0 656 690-1,100580-79 0390-830:) :610-885660-840810-980473-71 6580-860710-1,01 0992-1,25 7700-980730-1,30 01,500660-1,02 0550-650 551§940 605- 9 14§ 720-82 5660-763 840 770-1,35 0720-1,22 0456^75 930 634-66 2 800 580-67 5 650 710-89 0 900 600-93 0447-609 1

"a" rati o

38-413226-3 8212939-5 41931-4 437-3 925-3 235^14139-4 63228^ 33325-3 631-4121-2 932-3424-2 526-2 922-3633^t433^t 646-5 536-4635-514022-3 635^7403041-5 739-41344 025-3 235^* 722-264423-2 93329-3 42926-3 66026-3 837-48

"c" rati o

261 712-1 7161 826-3 21913-2 02070-7 3f 17-2 12320-2 621-2 610-152012-1 914-199-1323-2 519-22

17-1 9

17-2410-1 523-3220-2 520-3019-302824-2 919-2527264 521-2 214-152014-1 911-1 592511-1 22522-2 31115-2 52720-3 023-3 1

Style t lengt hGum)

18-19

13-1 4

14-161212 *111211-1 416-1814-1 8151312-1 4

12-1 4

14-161213-1 620-2 612-1 411-1415-19

15-1 7

12-1512-1 714-1811-1 414-1613-181512-1 5

11-1 3

1214-1 591112-1 41115-1 912-1 612-141312-1 31118-2 21312-1 714*12-1 513

Vulva l %

73-757471-7 7697 372-7 47184-8 869-7 072-7 87369-7 3

68-7 6

74-7 567-7 38271-7 676-80

67-8 3

76-8072-7 376-8168-7 5

69-7 4

67-7 870-7373-7 771-7 67371-7 669-73767 37671-7 471-727276-8 272-7 668-7 36473-747375-7 97173-8 270*73-8 073-77

Vulva lflap

9

++7

+9

- ----

79

+---

--9- -

+ +

+?

+7

+9

+-+7

-7

--+9-9?9 9

+-?

+

* Measure d from illustration . t A n obviou s error . $ Measurement s of lectotyp e materia l by Nickle , 1970. § Measurement s from Fuchs , 1937. || Measurement s of lectotyp e materia l by Nickl e ct al. , 1981 .

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Table 2. Diagnosti c dat a on Bursaphelenchus males: essentia l measurement s and ratios .

Bursaphelenchus species

bakeri bestiolus borealis

chitwoodiconjunctuscorneoluscrenaticryphalidigituluseggersieidmanniclytruseremuseucarpusfraudulentlyfungivorusgeorgicusgonzalezihunanensis sp. n. huntiidiusincurvuskevinikolymensisleonilignicoluslignophilusmucronatusnaujacinewmexicanusnuesslinipinasteripiniperdaepityogenipoligraphiratzeburgiisachsiscolytiseanisilvestrissteinerisutoricussychnustalonusteratospicularistritrunculusvaricaudawilfordixerokarterusxylophilus

Body lengt h(Mm)

858-99 0 670 548-81 3 54745 5 570-70 0825-1,08 0885-945f530-710450-99 0675-85 5830-1,04 0782-840750-1,02 0645-685570-1,03 0 720 580-97 0480-590360-740:) :619-643600-645890-1,12 0423-706510-1,060590-820912-1,14 7640-970630-1,2001,250780-90 0500-620 570f820 589-785 f630-675662-69 1 800 660-1,29 0730-895476-48 0 740 490-54 7 800 450-49 0 650 540-84 0670-75 0680-825520-60 1§

"a " rati o

38-453225-4 3343 13539-5 151-5433-4121-3 132-3941-5 240-4943^* 933-3731^t53930-3 838-4 519-283421-2 427-3 624-4636-5636-4748-6 439-5132-5 24743-4 542-5 1383 135-6 538-3 9

26-2 9

3827-3 337-4423-2 63424-3 04728-3 02930-4 16131-4 135^t 5

"c" rati o

25-2 61816-2 924151829^ 225-3 715-19

19-2 8

19-2 3

21-2 6

31-3 6

33^t520-2 718-252421-2 9

15-1 9

18-213324-2 615-19

18-2 9

16-2621-3 118-2326-3620-2 72628-3 220-2 52422342 825-2 72518-2 7

18-2 2

211422-2 42523-3 32315-2 32624-3 421-2 9

Style t lengt h(urn)

1813-1 4

12-1 6

1216. 512*1112-1 4

11-1 2

14-1614-181511-1 2

12-1 4

12-1 5

14-159

12-1 519-2112-1 31112-1 7

14-1 6

12-1 5

13-1 7

14-1711-1 414-1613-1 815

1 5

11-1 31214-1 5109-1 112-1 41113-1 8

12-1 5

121412-1 41118-2 21315-1 714*12-1 8

13-1 4

Spicul e lengt h 0*m)

19-2 218*16-2 01614. 513*17-1 814-1516*18-2 418-2 128*14-1 9

15-1 6

21-2 2

18*1913 *13-2 016-1 71417-1 818-2217-2110-2025-3 021-2 323-2913-2024*15-1 6

12-1 4

1417 *13-1 4

11-1 2

17-1 8

7*18-2 728-31172 219-2 315*15-1 814*13-1 710*14-1 519-23

* Measure d from illustration . t Measurement s from Fuchs , 1937. £ Measurement s of lectotyp e materia l by Nickle , 1970. § Measurement s of lectotyp e materia l by Nickl e et al. , 1981 .

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OF WASHINGTON , VOLUM E 55, NUMBE R 1, JANUAR Y 198 8

22(18) .

23(22) .

24(21) .

10(9). Vulv a 67-73% , male "a" rati o 31-

45 fungivorus Frankli n and

Hooper , 196 2 20(19) .

Vulv a

76-82% , male "a" rati o 27-

33 seani Giblin and Kaya , 198 3

11 (7) . Femal e length 456-47 5 /um, "c" mea -

suremen t

8.6-8.8, spicule thin -

elongat e

steineri (Riihm , 1956)

Goodey, 1960

Femal e length 580 /um an d greater , 21(20) .

"c" measuremen t 10 o r more ,

spicule more stou t 1 2

12(11). Male spicul e 26-31 /um lon g 1 3

Male spicul e 24 /u m or les s in lengt h

15

13(12) . Femal e withou t vulv a flap, "c " ra -

tio = 17-2 1 elytrus Massey , 196 0

Femal e wit h vulva l

flap, "c " rati o

eithe r mor e tha n 21 or les s tha n

17 1 4

14(13). Condylu s (cf . Fig . 12 ) of spicula r ca-

pitulu m with a conspicuou s pos -

terio r

flexure , male length 730-89 5

/um silvestris (Lieutie r and

Laumond , 1978) Baujard , 198 0

Condylu s of spicul e withou t poste -

rio r

flexure , male lengt h 1,250 /um

newmexicanus Massey , 197 4

15(12) . Vulva 82-88% 1 6

Vulva 78% or les s (varicauda vulv a

73-82% ) 1 7

16(15). Spicul e relativel y slende r in width ,

condylu s digitate , mal e "c" rati o

11-12 digitulus Loof, 196 4

Spicul e relativel y broade r

in width ,

condylu s broadly conical , mal e "c"

rati o 24

georgicus Devdariani , Kakuliya ,

and Khavtasi , 1980

17( 15) . Condylu s of spicula r capitulu m wit h

a posterio r flexur e 27(25) .

leoni Baujard, 1980

Condylu s of spicul e withou t poste -

rio r

flexur e 18 28(27).

18(17). Spicul e lengt h (measure d alon g arc )

11-1 3 /um or les s 1 9

Spicule lengt h 14 /u m or greate r (var-

icauda spicul e 13-17 /um) 2 2

19(18). Femal e with vulv a flap, lengt h 650- 29(27) .

700 /u m long ; "c" valu e of mal e

18.5 corneolus Massey , 1966

Femal e withou t vulv a flap, lengt h 30(29) .

25(22).

26(25).

720 /u m or greater ; "c" valu e of

male s 21-2 9 2 0

Female style t 12-16 /um long , spic -

ular lamina e strikingl y widene d or

"mitten-shaped " at midpoin t

wilfordi Massey, 1964

Female style t 9-11 /um long , spicula r

lamina e

not excessivel y widene d

21

Mal e lengt h 800 /um , spicul e 7 /um

lon g scolyti Massey, 1964

Male lengt h 630-67 5 /um, spicul e 11-

12 /u m lon g

ratzeburgii (Riihm , 1956)

Goodey , 196 0

Female s wit h vulva l

fla p 2 3

Females withou t vulva l flap, o r

presenc e of fla p no t determine d 25

Male style t 15-17 /um long , spicul e

length 13-1 7 /um

varicauda Thon g and

Webster, 1983

Mal e style t

13-15 /um long , spicul e

length 17-1 8 /um 2 4

Lateral incisure s present , mal e lengt h

820 /um , femal e lengt h 940 /um

pityogeni Massey, 1974

Latera l incisure s absent , mal e lengt h

670 /um , femal e lengt h 830 /um

bestiolus Massey, 1974

Male lengt h 450-54 7 /um (eggersi

male lengt h 450-99 0 /um) 2 6

Male lengt h 547 /um or mor e 2 7

Male style t 12-14 /um long , vulv a

73-74% , female "c" rati o 11-1 2

sychnus (Riihm , 1956)

Goodey ,

1960

Male style t 18-2 2 /um long, vulva

75-79% , female "c" rati o 22-2 3

teratospicularis Kakuliy a and

Devdariani , 1965

Mal e style t

18-19 /on 2 8

Male style t 18 /u m or less , averag e

length 16 /u m or les s 2 9

Female style t 18-1 9 /um , vulva 72-

74% bakeriRuhm, 196 4

(syn. : naujaci Baujard , 1980 )

Femal e style t

16-18 /um vulv a 75-

76% piniperdae Fuchs, 1937

Spicul e lengt h

18-24 /um 3 0

Spicule lengt h no greate r tha n 18 /u m

33

Style t lengt h of bot h sexe s 11-14 /um,

Copyright © 2011, The Helminthological Society of Washingto n

PROCEEDINGS OF TH E HELMINTHOLOGICA L SOCIETY

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. Spicule s of Bursaphelenchus spp. (partiall y afte r Tarja n and Baeza-Aragon , 1982) .

Copyright © 2011, The Helminthological Society of Washingto n

OF WASHINGTON , VOLUME 55, NUMBE R 1, JANUAR Y 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

Figur e 13. Continued.

Copyright © 2011, The Helminthological Society of Washingto n

10 PROCEEDING S OF TH E HELMINTHOLOGICA L SOCIETY

"a" rati o 48 o r mor e

lignophilus (Korner , 1954)

Meyl, 1960

Style t lengt h of bot h sexe s 14 /ur n or

greater , "a" rati o 39 o r les s 3 1

31(30). Femal e lengt h 991-1,12 2 /um, vul-

va 69-70% , female "a" rati o 37-

39 eggersi (Riihm , 1956)

Goodey , 196 0

Femal e length 990 /um or less , vulv a

72-81% , female "a" rati o 26-

32 3 2

32(31). Male lengt h 675-85 5 /urn; spicule

narro w wit h thornlik e pointe d

rostru m

eidmanni (Riihm , 1956 )

Goodey ,

1956

Male lengt h 890-1,12 0 /urn, spicul e

broa d wit h narro w digitat e ros -

tru m kevini Giblin, Swan ,

andKaya , 1984

33(29) . Male lengt h 547 /um , femal e length

619 M m chitwoodi (Riihm , 1956 )

Goodey , 196 0

(syn.: pinasteri Baujard , 1980)

Male lengt h 570 /um an d greater , fe-

mal e lengt h 650 /um or greate r (xe-

rokarterus range 600-90 3 /um,

idius range 610-88 5 /um) 3 4

34(33). Spicule lengt h 14-1 5 yum (exc . poli-

graphi spicul e 13-18 /um) 3 5

Spicule length 15 /u m or mor e (ere-

mus rang e 14-1 9 /um ) 3 8

35(34). Male lengt h 650 /um , vulv a 71%

tritrunculus Massey , 1974

Male lengt h greate r tha n 680 /um or

les s tha n 643 /um , vulv a 73% o r

mor e 36

36(35) . Condylu s of spicul e a reduce d pro -

tuberance , mal e lengt h

619-64 3

/urn

idius (Riihm , 1956 ) Goodey , 196 0

Condylu s

of spicul e prominent , mal e

lengt h 680-82 5 /um 3 7

37(36). Vulva 73-80% , stylet lengt h bot h sexes 12-1 8 /^m

xerokarterus (Riihm , 1956 )

Goodey , 196 0

(syn. : sutoricus Devdariani , 1974 )

Vulva 73% , style t lengt h bot h sexe s

11 /u m talonus (Thorne, 1935 )

Goodey, 1960

38(34) . Rostru m of capitulu m sharpl y

pointe d

3 9

Rostru m of capitulu m digitat e 4 2

39(38). Condylu s rectangula r to rectangu -

lar-elongat e

4 0

Condylus digitate to broadl y digitat e

41

40(39) . Femal e style t 12-1 4 /urn long , mal e

lengt h 750-1,02 0 /um, male "c"

valu e 3 3-4 5

eucarpus (Riihm , 1956 )

Goodey , 196 0

Femal e stylet 15-19 /um long , mal e

lengt h 600-64 5 /um, male "c" val-

ue 24-2 6 incurvus (Riihrn , 1956)

Goodey , 196 0

41(39) . Male lengt h 780-90 0 /um, "c" valu e

of femal e 24-29 , spicula r lamina e

with symmetricall y curve d dorsa l

surface nuesslini (Riihm , 1956 )

Goodey , 196 0

Male lengt h 662-69 1 um, "c " valu e

of femal e 14-15 , spicula r lamina e

with dorsa l surfac e formin g an 80 °

angl e sachsi (Riihm, 1956)

Goodey, 1960

42(38) . Femal e style t 9-11 /um long , femal e

"c" valu e at leas t 26, spicula r ca-

pitulu m slightl y depresse d 4 3

Female style t 13 /u m long , femal e "c"

valu e 23, spicula r capitulu m

moderatel y depresse d

eremus (Riihm, 1956 )

Goodey , 196 0

43. Femal e style t 9 /um long , mal e lengt h

589-785/um , spicule lengt h 13-1 4

/urn poligraphi Fuchs , 1937

Femal e style t 11 /u m long , mal e

lengt h

825-1,08 0 yum, spicul e

length 17-1 8 /um

crenati (Riihm , 1956)

Goodey , 1960

The essentia l measurement s and ratio s (Table s

1, 2 ) an d spicul e shape s foun d withi n the specie s

(Fig .

13 ) whic h follo w are a n amplificatio n of

thos e presente d in Tarja n and Baeza-Arago n

(1982), which are brough t up t o dat e wit h the

inclusio n of th e 6 recentl y describe d specie s and

additiona l dat a on th e olde r species . It i s incum -

ben t upo n

the author s to cautio n potentia l user s

tha t severa l

of th e specie s are poorl y describe d

and lackin g essentia l data . Almos t no mentio n

of nematod e variatio n is made , henc e the ke y

and diagnosti c dat a mus t be relie d upo n only

with adequat e reservation . Spicul e shap e has bee n

assumed

t o exhibi t little , if any , variation . Be-

Copyright © 2011, The Helminthological Society of Washingto n

x

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