On ogeny and in aspeci ic a iabili y o he ail shape in Xiphinema mon eneg inum and X. hy caniense
E-mail: laslo.ba [email protected]. s
On ogeny and in aspeci ic a iabili y o he ail shape in
Xiphinema mon eneg inum and X. hy caniense
(Nema oda: Longido idae)
László BARSI
Uni e si y o No i Sad, Facul y o Sciences, Depa men o Biology and Ecology, T g Dosi eja Ob ado ića 2,
21000 No i Sad, Se bia
Biologia Se bica, 2025, 47: 18-26
O iginal pape
DOI: DOI 10.5281/zenodo.17679075
Summa y. The mic oscopic size o nema odes, hei mo phological simila i y, limi ed numbe o dis inguishable axo-
nomic cha ac e s and o e lapping mo phome ic measu emen s, make hem one o he mos di icul o ganisms o
iden i y. Mainly, a ious mo phological and mo phome ical ea u es a e used o hei p ima y iden i ica ion and
cha ac e iza ion. Knowledge o in e speci ic and in aspeci ic a ia ions o hese ea u es in di e en popula ions o
di e en o igins is essen ial and o en c ucial when dealing wi h species ha a e e y simila o each o he . Xiphinema
mon eneg inum om Mon eneg o and X. hy caniense om I an ep esen wo geog aphically dis an bu mo phologi-
cally close species ha can be sepa a ed by on ogeny o he ail shape and some mo phome ic indices. Females and
males o X. mon eneg inum ha e a sligh ly sho e ail, a sho e hyaline po ion o he ail, and g ea e ail ip a ia ion
compa ed o emales and males o X. hy caniense. The dis ance be ween he cloacal opening and he adanal pai o
supplemen s in males is a good diagnos ic cha ac e o sepa a ing hese wo species since he e is no o e lap in alues.
Keywo ds: adanal pai o supplemen s, cloacal opening, geog aphically dis an , I an, Mon eneg o.
Accep ed: 20 Janua y 2025 / Published online: 26 Janua y 2025
INTRODUCTION
Because o hei mic oscopic size, mo phological simi-
la i y, limi ed numbe o dis inguishable axonomic cha ac-
e s and o e lapping mo phome ic measu emen s, nema-
odes a e conside ed one o he mos di icul o ganisms o
iden i y (Oli ei a e al. 2011). Thei p ima y iden i ica ion
and cha ac e iza ion a e mainly based on a ious mo pho-
logical ea u es (Bha e al. 2022). Tail shape and leng h a e
some o hese ea u es, bu hey show conside able a ia ion
om long ili o m o sho blun ly ounded. Tail shapes in
ju eniles o a ious s ages is a good cha ac e o he di e -
en ia ion o Xiphinema species (Siddiqi 1997). The s udy o
he a iabili y o mo pho-ana omical cha ac e s wi hin and
be ween di e en popula ions o di e en o igins is essen ial.
In aspeci ic a iabili y o mo phology and mo phome y is
always p esen o a g ea e o lesse ex en , e en in pa heno-
gene ic species. Knowing his a iabili y is o en c ucial when
dealing wi h species ha a e e y simila o each o he . Se ing
aside molecula analysis as an in eg al pa o in eg a i e ax-
onomy and ocusing only on mo phology, he mo phological
cha ac e is ics o an indi idual a e wha we i s encoun e
du ing i s obse a ion. Fo he iden i ica ion o he popula ion
o a species, i is always a lucky ci cums ance i , in addi ion
o adul s, ju enile s ages a e a ailable in sa is ac o y numbe .
Howe e , in p ac ice his is o en no he case.
In Longido idae, he numbe o ju enile de elopmen-
al s ages (JDS) be o e becoming adul s is mos o en 4 (J1,
J2, J3, J4), and less o en 3 (JI, JII, JIII) (Robbins e al. 1995,
1996). Each subsequen JDS, s a ing om he i s s age (J1/
JI), shows a ce ain deg ee o di e ence in mo phology and
mo phome y, compa ed o he p e ious s age. This is also he
On ogeny and in aspeci ic a iabili y o he ail shape in Xiphinema mon eneg inum and X. hy caniense
case wi h he de elopmen o he ail. Al hough knowledge o
he on ogeny o he ail shape in one species is impo an o
knowing he species i sel ; o compa ison o wo species ha
a e simila in many ways, he on ogeny o he ail becomes
e en mo e impo an .
Xiphinema mon eneg inum Ba si, Lambe i & Agos i-
nelli, 1998 om Mon eneg o (Ba si e al. 1998) and X. hy -
caniense Fadaka , Pou jam, Ba si & Ped am, 2021 om I an
(Fadaka e al. 2021) ep esen wo geog aphically dis an
bu mo phologically close species ha can be sepa a ed by
on ogenesis o ail shape and some mo phome ic indices.
In he o iginal desc ip ion o X. mon eneg inum (Ba si e
al. 1998) ail shapes o emales we e illus a ed wi h h ee
d awings ( ig. 4B-D) and one pho omic og aph ( ig. 5B), o
males wi h one d awing ( ig. 4G) and one pho omic og aph
( ig. 5C), and o ou JDS wi h only one d awing o each
s age ( ig. 4H-K). In he o iginal desc ip ion o X. hy cani-
ense (Fadaka e al. 2021) ail shapes o emales, males and
ou JDS a e well illus a ed ei he wi h d awings ( igs 1-2)
o pho omic og aphs ( igs 3-5).
The goals o he p esen s udy we e: i) o show he in a-
speci ic a iabili y o ail shape in emales, males and ju eniles
o X. mon eneg inum; ii) o compa e he cha ac e is ics o he
ail egion o emales and males o X. mon eneg inum and X.
hy caniense; iii) o gi e a compa a i e accoun o he on oge-
ne ic de elopmen o he ail shape in hese wo species.
MATERIAL AND METHODS
In his s udy o X. mon eneg inum, pe manen slides
o he ype popula ion and unpublished ma e ial om h ee
o he popula ions ound in di e en locali ies in Mon ene-
g o we e used. These we e: ype ma e ial: hizosphe e o
Ca pinus o ien alis Mill. a Mo inj (Ad ia ic coas ), 08-06-
1994, 05-04-1997; unpublished ma e ial: hizosphe e o C.
o ien alis, Dob čin, Canyon o Cije na Ri e , 16-06-2001;
hizosphe e o C. o ien alis, K uše ice, O jen M ., 24-09-
2008; hizosphe e o Ace campes e L. and Co ylus a ellana
L., Ja ljen (ea lie Ja ljem), Golija M ., 13-08-2010.
Pho og aphs we e aken using a Zeiss Axio Image A1
compound mic oscope equipped wi h an AxioCam MRc 5
digi al came a. D awings we e made based on o iginal pho-
og aphs o emale ails and ju enile s ages o X. mon eneg i-
num and X. hy caniense.
RESULTS AND DISCUSSION
In aspeci ic a iabili y o ail shape in emales, males
and ju enile s ages o X. mon eneg inum
A compa a i e s udy o he ails o emales and males
in 4 popula ions e ealed he exis ence o ce ain a iabili y
in shape (Figs 1 and 2). The posi ion, shape and size o he
peg is he mos ob ious exp ession o his a iabili y. Peg-
less ails a e ex emely a e. Some mu ually e y simila o
almos iden ical o ms appea mo e o en, and some o ms
less o en in he popula ions. In he ype popula ion, each
JDS has a cha ac e is ic ail shape, which shows he deg ee
o a iabili y wi hin each s age (Fig. 3).
In emales, ail sho , ounded o do sally mo e con ex,
wi h a sub-cen al peg o a iable leng h and blind canal;
excep ionally wi hou peg and blind canal (Fig. 1).
In males, ail is simila o ha o emales, wi h a sub-
cen al peg o a iable leng h and blind canal; excep ionally
wi hou peg and blind canal (Fig. 2).
In ju eniles, ail o J1 conical-subdigi a e, o J2 do sally
con ex conoid wi h subdigi a e e minus, o J3 do sally con-
ex conoid, digi a e wi h en al peg, o J4 do sally con ex
conoid, digi a e wi h almos en al o sub-cen al peg, a ely
pegless (Fig. 3)
Commen s on he ail egions o emales and males o X.
mon eneg inum and X. hy caniense
A i s glance, he ail shapes o emales on he one
hand, and males on he o he hand, in hese wo species a e
e y simila o each o he (Fig. 4).
In emales, ail sho , ounded o do sally mo e con ex,
wi h a sub-cen al peg and blind canal, excep ionally wi hou
peg and blind canal (Fig. 4A-B, C-D and Fig. 5). Females
o X. mon eneg inum ha e a sligh ly sho e ail, a sho e
hyaline po ion o he ail, and g ea e ail ip a ia ion com-
pa ed o emales o X. hy caniense (Table 1, Fig. 5).
Males o X. mon eneg inum ha e a sligh ly sho e ail,
a sho e hyaline po ion o he ail, and g ea e ail ip a ia-
ion compa ed o males o X. hy caniense (Table 1, Fig. 4E-F,
G-H). The mos signi ican di e ence in males is he dis ance
be ween adanal pai o supplemen s and cloacal opening.
The dis ance be ween he cloacal opening and adanal pai
o supplemen s in X. mon eneg inum (n = 41) 14.7 (12.3-
16.8) µm, and in X. hy caniense (n = 9) 25.3 (22.6-27.0) µm
wi hou o e lapping. Spicule leng hs mos ly o e lap, 80.7-
92.9 µm in X. mon eneg inum (n = 24), and 72-95 µm in X.
hy caniense (n = 16).
The dis ance be ween he cloacal opening and he ad-
anal pai o supplemen s is a good diagnos ic cha ac e o
sepa a ing hese wo species, because he e is no o e lap in
alues, whe eas he leng h o he spicules is no sui able due
o signi ican o e lap.
On ogeny o he ail shape in X. mon eg inum and X.
hy caniense
In emales o X. mon eg inum and X. hy caniense, he
Biologia Se bica 47 19
L. Ba si On ogeny and in aspeci ic a iabili y o he ail shape in Xiphinema mon eneg inum and X. hy caniense
ails a e ela i ely simila and o someone wi h less expe i-
ence hey may e en seem he same (Fig. 5). When i comes o
he i s (J1), second (J2), hi d (J3) and ou h (J4) ju enile
s ages o hese wo species, he si ua ion is somewha di e -
en (Fig. 6).
Tail o J1 conical-subdigi a e in X. mon eg inum and
elonga e conoid, do sally con ex, en ally conca e, wi h
blun ly ounded ip in X. hy caniense (Fig. 6).
Tail o J2 do sally con ex conoid wi h subdigi a e e -
minus in X. mon eg inum and do sally con ex and en ally
conca e, ending o subdigi a e ip in X. hy caniense (Fig. 6).
Tail o J3 do sally con ex conoid, digi a e wi h en al
peg in X. mon eg inum and conical do sally con ex subdigi-
a e in X. hy caniense (Fig. 6).
Tail o J4 do sally con ex conoid, digi a e wi h almos
en al o sub-cen al peg, a ely pegless in X. mon eg inum
and sho do sally mo e con ex wi h a small muc o in X.
hy caniense (Fig. 6).
Ju enile s ages o X. mon eg inum ha e somewha
sho e ail and sho e hyaline po ion o he ail compa ed
o ju enile s ages o X. hy caniense (Table 1).
Coomans e al. (2001), in hei capi al wo k, highligh ed
ha “The ela ions be ween he a ious ail ypes a e complex
and can be assessed h ough he s udy o he pos emb yonic
de elopmen ”. A e analyses o he ail mo phology o ju-
eniles and adul s and ail de elopmen du ing emb yogen-
esis in he genus Xiphinema, hese au ho s classi ied he ail
shapes in ju eniles and adul s in o 13 ypes, anging om
ili o m (Type 1) o b oadly ounded (Type 13), and p o-
ided an illus a ion wi h some ep esen a i es o e e y ype
(Coomans e al. 2001, p. 31, ig. 11). They also de ined he
pa e ns o changes in he ail shape du ing pos emb yonic
de elopmen and p esen ed a scheme o illus a e which ail
ypes could be de i ed om o he ypes (Coomans e al.
Fig. 1. In aspeci ic a iabili y o ail shape in emales o Xiphinema mon eneg inum om Mon eneg o. Based on ype popula ion om
Mo inj (Ad ia ic coas ), and popula ions om Dob čin (Canyon o Cije na Ri e ), K uše ice (O jen M .), and Ja ljen (Golija M .).
20 Biologia Se bica 47
On ogeny and in aspeci ic a iabili y o he ail shape in Xiphinema mon eneg inum and X. hy caniense
Fig. 2. In aspeci ic a iabili y o ail shape in males o Xiphinema mon eneg inum om Mon eneg o. Based on ype popula ion om
Mo inj (Ad ia ic coas ), and popula ions om Dob čin (Canyon o Cije na Ri e ), K uše ice (O jen M .), and Ja ljen (Golija M .).
2001, p. 32, ig. 12). The whole pa e n was based on pub
-
lished da a o mo e han 80 species o which he ail shape
o di e en ju enile s ages was known a ha ime. One o
hese species was X. mon eneg inum in which he ail shapes
we e illus a ed wi h one d awing each o J1, J2, J3 and J4,
and h ee d awings o he emale. Among many o he s, hese
d awings we e used o cons uc ion o ail shape sco es o
Xiphinema species o which mo phological da a o a leas
h ee ju enile s ages we e known (Coomans e al. 2001).
The ail shape sco e o X. mon eneg inum was: J1 (7), J2
(8), J3 (9), J4 (9), emale (10). The same ail shape sco e could
apply o X. hy caniense as well. Based on he s udied ma e ial
om Mon eneg o and I an, he in aspeci ic and in e speci ic
a iabili y o he ail shape o emales and ou ju enile s ages
in X. mon eneg inum and X. hy caniense is shown he e (Figs
5-6). I can be no ed ha emales wi h pegless ails a ely
appea in bo h species. Likewise, wi h X. mon eneg inum,
Table 1. Compa ison o ail leng h and hyaline po ion o
he ail o ju eniles and adul s in Xiphinema mon eneg i-
num and X. hy caniense.
Tail leng h and hyaline po ion o ail, min-max (µm)
X. mon eneg inum X. hy caniense
J1 47.1-57.8; 9.4-13.8 60-67.5; 16-20
J2 42.8-52.9; 11.9-17.3 55-67.5; 17.5-25
J3 37.5-50; 8.8-13.7 45-60; 15-21.3
J4 35.0-44.6; 7.5-11.9 42.5-55; 12.5-17.5
Female 31.4-42.7; 7.5-13.8 35-47.5; 12.5-21.3
Male 36.9-47.5; 6.9-14.4 41-55; 11.5-17.5
Sou ces: Ba si e al. 1998; Fadaka e al. 2021.
Biologia Se bica 47 21
L. Ba si On ogeny and in aspeci ic a iabili y o he ail shape in Xiphinema mon eneg inum and X. hy caniense
Fig. 3. In aspeci ic a iabili y o ail shape in ou ju enile de elopmen al s ages o Xiphinema mon eneg inum. A1-A4 = J1; B1-B6 = J2;
C1-C7 = J3; D1-D11 = J4. Based on ype popula ion om Mo inj (Ad ia ic coas ), Mon eneg o.
22 Biologia Se bica 47
On ogeny and in aspeci ic a iabili y o he ail shape in Xiphinema mon eneg inum and X. hy caniense
indi iduals o he J4 s age wi h a pegless ail a ely appea .
These ac s lead o he addi ion o he ail shape sco e in bo h
species (Fig. 7). Addi ional ail shape sco es o X. mon ene-
g inum a e: J1 (7), J2 (8), J3 (9), J4 (9), emale (12); J1 (7), J2
(8), J3 (9), J4 (12), emale (12) (Fig. 7). Addi ional ail shape
sco e o X. hy caniense is: J1 (7), J2 (8), J3 (9), J4 (9), emale
(12) (Fig. 7).
The capi al wo k o Coomans e al. (2001) on cha ac e
analysis, phylogeny and biogeog aphy o he genus Xiphi-
nema is an indispensable sou ce o in o ma ion o esea ch-
e s dealing wi h Xiphinema species. O e he pas 23 yea s, a
la ge numbe o new Xiphinema species ha e been desc ibed
wi h de ailed desc ip ions and illus a ions o adul and ju-
enile s ages, which ep esen s an i eplaceable upg ade o
he wo k o Coomans e al. (2001). The eali y is ha he
wo k o Coomans e al. (2001) needs o be upda ed based
on new desc ip ions, bu he big ques ion is whe he anyone
will e e do i .
LITERATURE
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om Bosnia and He zego ina and X. mon eneg inum sp. n. om
Mon eneg o, Yugosla ia (Nema oda: Do ylaimida). Nema ologia
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Khan MN, Sayyed RZ, Poczai P, e al. 2022. Ad ances in nema ode
iden i ica ion: A jou ney om undamen als o e olu iona y aspec s.
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eny and biogeog aphy o he genus Xiphinema Cobb, 1913 (Nema
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oda: Longido idae). Annales Sciences Zoologiques, Musée Royal de
l’A ique Cen ale, Te u en. 287:1–239.
Fadaka S, Pou jam E, Ba si L, Ped am M. 2021. A new species o he genus
Xiphinema Cobb 1913 (Nema oda: Longido idae) om no h and
Fig. 4. Compa ison o ail egions. Xiphinema mon eneg inum: A-B, emale ail egion; E-F, mail ail egion. X. hy caniense: C-D, emale
ail egion; G-H, mail ail egion. (Abb e ia ions: a = anal opening; as = adanal pai o supplemen s; bc = blind canal; co = cloacal opening;
p = peg; spic = spicule.)
Biologia Se bica 47 23
L. Ba si On ogeny and in aspeci ic a iabili y o he ail shape in Xiphinema mon eneg inum and X. hy caniense
no hwes I an. Eu opean Jou nal o Plan Pa hology. 160:557–574.
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Oli ei a CMG, Mon ei o AR, Blok VC. 2011. Mo phological and molecula
diagnos ics o plan -pa asi ic nema odes: wo king oge he o ge he
iden i ica ion done. T opical Plan Pa hology. 36(2):65–73.
Robbins RT, B own DJF, Halb end JM, V ain TC. 1995. Compendium o
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Fig. 5. Compa a i e p esen a ion o in aspeci ic a iabili y o ail shape in emales o Xiphinema mon eneg inum om Mon eneg o and X.
hy caniense om I an. (Scale ba s = 20 μm.) (D awings wi h dashed line ed awn om Fadaka e al. 2021, ig. 1h, i, j, n.)
24 Biologia Se bica 47
On ogeny and in aspeci ic a iabili y o he ail shape in Xiphinema mon eneg inum and X. hy caniense
Fig. 6. Compa a i e p esen a ion o in aspeci ic a iabili y o ail shape in ou ju enile de elopmen al s ages (J1, J2, J3, J4) o Xiphinema
mon eneg inum om Mon eneg o and X. hy caniense om I an. (Scale ba s = 20 μm.)
Biologia Se bica 47 25
L. Ba si
Fig. 7. Compa a i e p esen a ion o he on ogeny o ail shape in Xiphinema mon eneg inum om Mon eneg o and X. hy caniense om
I an. Types o he ail shapes a e Coomans e al. 2001: 7 (conoid o do sally con ex conoid), 8 (do sally con ex conoid wi h subdigi a e
e minus), 9 (do sally con ex conoid, digi a e wi h en al peg), 10 (con ex conoid, digi a e wi h cen al [on he e minus] peg o bulge),
12 (conoid wi h b oadly ounded ip).
26 Biologia Se bica 47
