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
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