A synopsis of the species of Myxobolus Bütschli, 1882 (Myxozoa: Bivalvulida) parasitising Indian...
21
A synopsis of the species of Myxobolus Bu ¨ tschli, 1882 (Myxozoa: Bivalvulida) parasitising Indian fishes and a revised dichotomous key to myxosporean genera Harpreet Kaur • Ranjeet Singh Received: 15 May 2011 / Accepted: 14 July 2011 Ó Springer Science+Business Media B.V. 2011 Abstract A synopsis of 131 nominal species of Myxobolus Bu ¨tschli, 1882 (Myxozoa: Myxosporea: Myxobolidae) reported from India is presented. For each species, the relevant morphometric and mor- phological data are indicated, as well as the host(s), site(s) of infection within the host and locality data. A revised dichotomous key of 59 genera of the class Myxosporea has also been included. This key incor- porates 10 additional genera to that proposed in 1991 by Lom & Dykova ´. Introduction Myxosporeans are an abundant and diverse group of parasites infecting fishes, amphibians and reptiles (they have also recently been reported from birds and mammals). According to Lom & Dykova ´(2006), the phylum Myxozoa includes four malacosporean and 2,180 myxosporean species in a total of 62 genera. However, three more genera, Soricimyxum Prunescu, Prunescu, Pucek & Lom, 2007, Gadimyxa Køie, Karlsbakk & Nylund, 2007 and Thelohanelloid Sarkar, 2009 have subsequently been described (Prunescu et al., 2007; Køie et al., 2007; Sarkar, 2009). Myxobolus Bu ¨tschli, 1882 (Myxobolidae) remains by far the largest genus within the Myx- osporea. The phylum Myxozoa has been studied by only a limited number of workers in the Indian subcontinent. Most of the work on this phylum has been carried out, on both freshwater and marine fishes, in two states, West Bengal and Andhra Pradesh. During the last decade, Basu and Haldar (1999, 2002a, b, 2003a, b, 2004), Bandyopadhyay et al. (2007), Acharya & Dutta (2007) and Basu et al. (2006, 2009) have contributed a significant number of new species from freshwater and marine fishes belonging to genera Myxobolus, Thelohanellus Kudo, 1933, Myxidium Bu ¨tschli, 1882 and Henneguya The ´lohan, 1892. Reports of myxosporean species from other parts of India are few, except for some from freshwater fishes of Manipur (Hemananda et al., 2009, 2010) and Kerala (Sheeja & Janardanan, 2006). Hemananda et al. (2006), during an investigation on the preva- lence of myxozoa parasites in the fish Cirrhina mrigala, recorded the pathogenicity and mortality due to Myxobolus haldi Hemananda, Mohilal & Vishwanath, 2006 infecting the gill filaments. In northern India, Gupta & Khera (1987, 1988a, b, c, d, 1989a, b, 1990, 1991) recorded 25 species belonging to the genera Myxobolus, Henneguya, Myxidium, Thelohanellus and Unicauda Davis, 1944 infecting freshwater fishes. Since 2007, there have been numerous other myxosporean species, particularly those belonging to Myxobolus, described in India. For example, Kaur & Singh (2008, 2009, 2010a, b, c, H. Kaur (&) Á R. Singh Department of Zoology, Punjabi University, Patiala 147002, India e-mail: [email protected] 123 Syst Parasitol (2012) 81:17–37 DOI 10.1007/s11230-011-9321-z
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Transcript of A synopsis of the species of Myxobolus Bütschli, 1882 (Myxozoa: Bivalvulida) parasitising Indian...
A synopsis of the species of Myxobolus Butschli,1882 (Myxozoa: Bivalvulida) parasitising Indian fishesand a revised dichotomous key to myxosporean genera
Harpreet Kaur • Ranjeet Singh
Received: 15 May 2011 / Accepted: 14 July 2011
� Springer Science+Business Media B.V. 2011
Abstract A synopsis of 131 nominal species of
Myxobolus Butschli, 1882 (Myxozoa: Myxosporea:
Myxobolidae) reported from India is presented. For
each species, the relevant morphometric and mor-
phological data are indicated, as well as the host(s),
site(s) of infection within the host and locality data. A
revised dichotomous key of 59 genera of the class
Myxosporea has also been included. This key incor-
porates 10 additional genera to that proposed in 1991
by Lom & Dykova.
Introduction
Myxosporeans are an abundant and diverse group of
parasites infecting fishes, amphibians and reptiles
(they have also recently been reported from birds and
mammals). According to Lom & Dykova (2006), the
phylum Myxozoa includes four malacosporean and
2,180 myxosporean species in a total of 62 genera.
However, three more genera, Soricimyxum Prunescu,
Prunescu, Pucek & Lom, 2007, Gadimyxa Køie,
Karlsbakk & Nylund, 2007 and Thelohanelloid
Sarkar, 2009 have subsequently been described
(Prunescu et al., 2007; Køie et al., 2007; Sarkar,
2009). Myxobolus Butschli, 1882 (Myxobolidae)
remains by far the largest genus within the Myx-
osporea.
The phylum Myxozoa has been studied by only a
limited number of workers in the Indian subcontinent.
Most of the work on this phylum has been carried out,
on both freshwater and marine fishes, in two states,
West Bengal and Andhra Pradesh. During the last
decade, Basu and Haldar (1999, 2002a, b, 2003a, b,
2004), Bandyopadhyay et al. (2007), Acharya &
Dutta (2007) and Basu et al. (2006, 2009) have
contributed a significant number of new species from
freshwater and marine fishes belonging to genera
Myxobolus, Thelohanellus Kudo, 1933, Myxidium
Butschli, 1882 and Henneguya Thelohan, 1892.
Reports of myxosporean species from other parts of
India are few, except for some from freshwater fishes
of Manipur (Hemananda et al., 2009, 2010) and
Kerala (Sheeja & Janardanan, 2006). Hemananda
et al. (2006), during an investigation on the preva-
lence of myxozoa parasites in the fish Cirrhina
mrigala, recorded the pathogenicity and mortality
due to Myxobolus haldi Hemananda, Mohilal &
Vishwanath, 2006 infecting the gill filaments. In
northern India, Gupta & Khera (1987, 1988a, b, c, d,
1989a, b, 1990, 1991) recorded 25 species belonging
to the genera Myxobolus, Henneguya, Myxidium,
Thelohanellus and Unicauda Davis, 1944 infecting
freshwater fishes. Since 2007, there have been
numerous other myxosporean species, particularly
those belonging to Myxobolus, described in India. For
example, Kaur & Singh (2008, 2009, 2010a, b, c,
H. Kaur (&) � R. Singh
Department of Zoology, Punjabi University,
Patiala 147002, India
e-mail: [email protected]
123
Syst Parasitol (2012) 81:17–37
DOI 10.1007/s11230-011-9321-z
2011a, b, c, d, e, f) have contributed 17 new species
of Myxobolus from freshwater fishes in the wetlands
of Punjab.
A synopsis of the species of Myxobolus parasitising
Indian fishes
Previous synopses include those of Eiras et al. (2005),
who provided a list of 744 valid species of Myxobolus
worldwide, and Kalavati & Nandi (2007), who
reported 104 species of this genus from Indian fishes
in addition to 175 species belonging to 29 myxosp-
orean genera in total. The present synopsis (presented
in Table 1) updates the number of Myxobolus species
infecting Indian fishes to 131 and presents data for
each species in a concise, tabulated format, giving
details on spore dimensions, morphology, the
site(s) of infection within the host, the host(s) and
the region(s) of India where the parasite has been
found. The data are obtained from the original
descriptions, sometimes supplemented by more
recent information.
Myxobolus sp. of Padma Dorothy & Kalavati
(1992) is included in the synopsis, although these
authors did not assign it a specific name for want of
adequate information. The following six species,
M. cartilaginis (Hoffman, Putz & Dunbar, 1965),
M. cerebralis (Hofer, 1903), M. encephalicus Muslow,
1911, M. episquamalis Egusa, Maeno & Sorimachi,
1990, M. parvus Shulman, 1962 and M. spinocurva-
tura Maeno, Sorimachi & Egusa, 1990, previously
reported from other parts of the world, have also been
reported from India by Ram et al. (1984), Das et al.
(1988), Kalavati & Anuradha (1992), Eiras and
D’Souza (2004) and Anuradha (1991), respectively.
Measurements given in Table 1 are in micrometres
unless otherwise stated, and for two- and three-
dimensional measurements, length is given first.
Geographical abbreviations used in this table are:
And, Andaman Islands; AP, Andhra Pradesh; Har,
Haryana; Kar, Karnataka; Ker, Kerala; Man, Mani-
pur; MP, Madhya Pradesh; Or, Orissa; Pun, Punjab;
Raj, Rajasthan; TN, Tamil Nadu; UP, Uttar Pradesh;
WB, West Bengal. Other abbreviations are: FC,
filament coils; ICN, intercapsular notch; ICP, inter-
capsular process; PC, polar capsule; PF, parietal
folds. In order to reduce space the authorities for the
taxa serve as references for the data sources. Ta
ble
1A
syn
op
sis
of
Myx
ob
olu
ssp
p.
fro
mIn
dia
nfi
shes
Spec
ies
Host
(s)
Sit
eof
infe
ctio
n
Loca
lity
Spore
size
PC
size
(s)
ICP
PF
FC
Cyst
size
PC
s
equal
M.
ali
garh
ensi
sB
hat
t&
Sid
diq
ui,
1964
[Syns
M.
ophic
ephali
Bhat
t&
Sid
diq
ui,
1964;
M.
punct
atu
sR
aych
audhuri
&C
hak
ravar
ty,
1970]
Ophio
cephalu
spunct
atu
sB
loch
Acc
esso
ry
resp
irat
ory
mem
bra
ne
WB
,
Pun,
UP
12–14
96–7.5
6.5
–8
92–2.5
and
6–7
92.5
Abse
nt
Abse
nt
–190–628
9192–533
Yes
/
No
M.
am
pull
ace
us
Lal
itha
Kum
ari,
1969
Barb
us
kolu
sS
kyes
Dors
alan
d
ven
tral
fin
AP
9.8
(8.6
–10.7
)9
7.1
(6.4
–7.9
)
5.0
–6.4
(5.8
)9
2.5
–2.9
(2.8
)
Abse
nt
55–6
–Y
es
M.
analfi
nus
Bas
uet
al.,
2009
Het
eropneu
stes
foss
ilis
Blo
ch
Anal
fin
WB
12.3
98.6
96.2
4.1
92.2
and
2.5
91.8
ICN pre
sent
Abse
nt
5–7
in
larg
er
PC
,3–4
in smal
ler
250
9200
No
M.
andhra
e(L
alit
ha
Kum
ari,
1969
)G
upta
&
Kher
a,1988(a
)[S
yn.
Myx
oso
ma
andhra
eL
alit
ha
Kum
ari,
1969]
Ophio
cephalu
spunct
atu
sB
loch
Surf
ace
wal
l
of
inte
stin
e
AP
13.5 (1
2.1
–15.7
)9
6.4
(5.7
–8.6
)
9.0
(8.6
–10.0
)9
1.7
(1.4
–2.1
)an
d
7.1
–9.3
(8.3
)9
1.4
–2.1
(1.7
)
Abse
nt
4–6
91
mm
No
18 Syst Parasitol (2012) 81:17–37
123
Ta
ble
1co
nti
nu
ed
Spec
ies
Host
(s)
Sit
eof
infe
ctio
n
Loca
lity
Spore
size
PC
size
(s)
ICP
PF
FC
Cyst
size
PC
s
equal
M.
anil
iS
arkar
,1989
Rhin
om
ugil
cors
ula
Ham
.,
Liz
am
acr
ole
pis
Sm
ith
Duoden
al
mes
ente
ry,
inte
stin
al
vil
li
WB
,
AP
9.8
–11.0
6
(10.7
4)
97.9
2–9.8
4
(8.6
9)
4.7
4–5.0
5
(4.8
2)
92.4
8–3.1
6
(3.0
5)
Pre
sent
Abse
nt
4–5
560–615
9206–288
Yes
M.
att
ui
Sar
kar
1985(a
)W
all
ago
att
uB
leek
erG
ut
WB
12.8
–15.2
(13.8
9)
95–9.6
(8.5
3)
4.8
–7.2
(5.9
2)
92.4
–4
(3.0
)
Abse
nt
Abse
nt
6–7
–Y
es
M.
banki
mi
Sar
kar
,1999
Sic
am
ugil
casc
asi
aH
am.
Inner
wal
lof
gal
lbla
dder
WB
10–11
(10.6
)9
8–9
(8.7
)3.5
–4.5
(3.9
7)
92.5
–3
(2.7
2)
Abse
nt
Abse
nt
4–6
–Y
es
M.
barb
iT
ripat
hi,
1952
Barb
us
tict
oH
am.
Skin
WB
12.6
–13.5
99.0
3.6
–4.5
92.7
Pre
sent
Abse
nt
––
Yes
M.
bata
eK
aram
chan
dan
i,
1970
Labeo
bata
Ham
.G
ill
fila
men
tsO
r9.6
–11.2
97.2
–8.8
4.0
–4.8
91.6
Pre
sent
Abse
nt
–1.5
mm
Yes
M.
ben
gale
nsi
sC
hak
ravar
ty&
Bas
u,
1948
Catl
aca
tla
Ham
.G
ills
WB
,
AP
8.5
6–9.3
69
6.4
–6.8
4.2
8–5.4
09
2.5
–3.2
Abse
nt
Abse
nt
–2–4.1
mm
Yes
M.
bhadre
nsi
sS
eenap
pa
&
Man
ohar
,1981
Labeo
rohit
aH
am.
Musc
les
Kar
8–11
(9.5
)9
7–8
(7.1
4)
3–4
(3.5
)9
2–3
(2.2
)
and
2–4
(2.5
)9
1–2
(1.7
5)
Pre
sent
in
form
of
thic
ken
ing
Not clea
r
––
No
M.
bhaduri
us
(Sar
kar
,
1985(a
))G
upta
&
Kher
a,1988(a
)[S
yn.
Myx
oso
ma
bhaduri
aS
arkar
,1985(a
)]
Wall
ago
att
uB
leek
erG
all
bla
dder
WB
8.8
–11.2
(10.5
9)
94.8
–6.7
2
(6.2
8)
4–6.4
(5.3
1)
9
2.4
–3.2
(2.7
8)
Abse
nt
Abse
nt
5–6
–Y
es
M.
biv
acu
ola
tus
Nar
asim
ham
urt
i&
Kal
avat
i,1986
Cla
rias
batr
ach
us
L.
Inte
stin
alw
all
AP
8–11
(9.0
)3–4.5
(4.2
)9
2.6
–4
(3.0
)A
bse
nt
36–7
2–4
92–3
mm
Yes
M.
bucc
oro
ofu
sB
asu
&
Hal
dar
,2004
Labeo
bata
Ham
.R
oof
of
bucc
al
cavit
y
WB
11.6
–12.7
(12.1
)9
6.4
–8.1
(7.1
)
4.5
–5.3
(4.9
)9
2.7
–3
(2.9
)an
d2–2.9
(2.5
)9
1.3
–1.7
(1.5
)
Abse
nt
Abse
nt
6–8
in
larg
er
PC
,
4–5
in
smal
ler
–N
o
M.
calb
asu
iC
hak
ravar
ty,
1939
Labeo
calb
asu
Ham
.,L
.ro
hit
aH
am.,
Cir
rhin
am
rigala
Ham
.,L
.bata
Ham
.,L
.dyo
chei
lus
McC
lell
and
Gil
ls,
fins,
gal
l
bla
dder
,
scal
es
WB
,
Pun
12.4
–15
98.2
–10
6.1
89
4.1
2an
d
4.1
29
3.0
9
Abse
nt
Abse
nt
–300–350
No
M.
calc
ari
feru
mB
asu
&
Hal
dar
,2003(b
)
Late
sca
lcari
fer
Blo
chG
ill
fila
men
tsW
B6.6
(6.1
–7.1
)9
6.2
(5.7
–6.5
)
3.8
–4.5
(4.2
)9
2–2.7
(2.3
)
Abse
nt
Abse
nt
4–5
–Y
es
M.
carn
ati
cus
See
nap
pa
&
Man
ohar
,1980(a
))
Cir
rhin
am
rigala
Ham
.In
ner
bas
eof
hem
ibra
nch
s
Kar
8.6
96.8
3.8
92
and
2.1
91.5
Pre
sent
Abse
nt
––
No
M.
catl
ae
Chak
ravar
ty,
1943
Cir
rhin
am
rigala
Ham
.,
Labeo
bata
Ham
.,L
.dyo
chei
lus
McC
lell
and
Gil
lsW
B,
Pun
14.5
–16.5
(14.9
)9
6.1
810.3
–12.3
69
2.0
6–3.0
1A
bse
nt
Abse
nt
–45–150
Yes
Syst Parasitol (2012) 81:17–37 19
123
Ta
ble
1co
nti
nu
ed
Spec
ies
Host
(s)
Sit
eof
infe
ctio
n
Loca
lity
Spore
size
PC
size
(s)
ICP
PF
FC
Cyst
size
PC
s
equal
M.
catl
iK
aur
&S
ingh,
2011(d
)C
atl
aca
tla
Ham
.G
ill
lam
ella
eP
un
7.9
–8.7
(8.3
)9
4.2
–5.2
(4.7
)
2.8
–3.8
(3.3
)9
1.2
7–2.0
7
(1.6
7)
Abse
nt
Abse
nt
5–7
–Y
es
M.
catm
rigala
eB
asu
&H
aldar
,
2004
Catl
aca
tla
Ham
.9
Cir
rhin
am
rigala
Ham
.
(hybri
d)
Gil
lla
mel
lae
WB
20.4
916.3
11.9
92.3
and
is
11
92.3
Abse
nt
Abse
nt
20–25
in
larg
er
PC
,
20–23
in
smal
ler
–N
o
M.
cart
ilagin
is(H
off
man
etal
.,
1965)
Lan
dsb
erg
&L
om
,1991
Lep
om
ism
acr
och
irus
Raf
.,
Mic
ropte
rus
salm
oid
esL
ac.
Gil
lca
rtil
age,
bas
eof
fin
rays
Har
10–12
(10.8
)9
9–11
(9.5
)
5–6
(5.3
)9
3–4
(3.1
)P
rese
nt
6–9
5–7
–N
o
M.
cere
bra
lis
Hofe
r,1903
Cir
rhin
us
spp.,
Labeo
spp.,
Channa
spp.
Car
tila
ge
of
hea
dan
d
ver
tebra
l
colu
mn
Har
7.4
–9.7
97–10
4.2
–6
93–3.5
Pre
sent
(insi
gnifi
cant)
Abse
nt
––
No
M.
chakr
ava
rtyi
Hal
dar
etal
.,
1983
Catl
aca
tla
Ham
.In
tern
aley
e
musc
ula
ture
WB
12.1
–14.3
(12.3
)9
7.7
–10.5
5.5
–6.6
93.3
–5
and
4.4
–5
92.2
–4.4
Abse
nt
Abse
nt
8–9
in
larg
er
PC
,6–7
in smal
ler
390–440
9
220–280
No
M.c
hannai
(Kal
avat
iet
al.,
1981)
Gupta
&K
her
a,1988(a
)[S
yn.
Myx
oso
ma
channai
Kal
avat
i
etal
.,1981
]
Channa
punct
atu
sB
loch
Fin
s,body
musc
les,
kid
ney
,
liver
Kar
14.5
–18.0
(17.2
)9
6.0
–6.5
(6.3
)
9–10.8
(10.2
)9
2.8
–3.2
(3.0
)an
d7.2
–8.8
(7.5
)9
2.8
–3.2
(3.0
)
Abse
nt
Abse
nt
8in
larg
er
PC
0.8
–1.2
mm
No
M.
chil
kensi
s(K
alav
ati
etal
.,
1992)
Kal
avat
i&
Nan
di,
2007
[Syn.
Rudic
apsu
lach
ilke
nsi
sK
alav
ati
etal
.,1992]
Labeo
rohit
aH
am.
Gal
lbla
dder
Or
7.2
–8
(7.7
7)
95.6
–6.6
(6.2
)
3.2
–4.8
(4.2
)9
1.8
–2.2
(1.9
8)
and
1–1.2
(1.0
8)
Abse
nt
Abse
nt
6–8
in
larg
er
PC
500–800
No
M.
chin
sura
hen
sis
Bas
u&
Hal
dar
,2003(b
)
Anabas
test
udin
eus
Blo
ch
Sca
les
WB
8–9.7
(8.4
)9
5.1
–6.1
(5.4
)
3.9
–6.6
(4.4
)9
1.8
–2.5
(2.1
)
Pre
sent
Abse
nt
5–6
–N
o
M.
chit
tali
iK
aur
&S
ingh,
2011(a
)
Punti
us
sophore
Ham
.
Gil
lla
mel
lae
Pun
99
6.1
84.5
92.4
Pre
sent
Abse
nt
4–5
–Y
es
M.
clari
ae
Hem
anan
da
etal
.,
2009
Cla
rias
batr
ach
us
L.
Corn
eaM
an10.5
3(1
0.2
–11.5
)9
7
(6.8
–7.7
)
3.4
–4.3
(3.6
7)
91.7
–2.6
(2.3
8)
Pre
sent
Abse
nt
––
Yes
M.
clari
iC
hak
ravar
ty,
1943
Cla
rias
batr
ach
us
L.
Gal
lbla
dder
liver
,te
stes
,
ovar
y
WB
11.3
–12.4
910.3
6.8
93.0
9A
bse
nt
Abse
nt
–775–780
9
604–877
Yes
M.
coel
iH
aldar
etal
.,1996
Chanos
chanos
Fors
skal
Gal
lbla
dder
Or
8.3
–14.9
(10.6
)9
4.1
–8.4
(5.4
9)
4.1
–9.1
(5.7
6)
91.7
–2.3
(2.8
5)
and
1.7
–6.6
(4.2
)9
1.7
–4.1
Abse
nt
Abse
nt
––
No
20 Syst Parasitol (2012) 81:17–37
123
Ta
ble
1co
nti
nu
ed
Spec
ies
Host
(s)
Sit
eof
infe
ctio
n
Loca
lity
Spore
size
PC
size
(s)
ICP
PF
FC
Cyst
size
PC
s
equal
M.
cruci
filu
s(Q
adri
1962(b
))
Lan
dsb
erg
&L
om
,1991
[Syn.
Gyr
osp
ora
cruci
filu
sQ
adri
1962(b
)]
Labeo
fim
bri
atu
sB
loch
Gil
lsA
P9–10
98–8.5
4–4.5
Abse
nt
Abse
nt
––
No
M.
curm
uca
eS
eenap
pa
and
Man
ohar
1980(a
)
Punti
us
curm
uca
Ham
.
Ben
eath
scal
es
Kar
8–11
(9.8
)9
7–8
(7.6
)4.5
–5
(4.9
)9
2–3
(2.5
)
and
3–4
(3.9
)9
2–3
(2.4
)
Pre
sent
Abse
nt
––
Yes
/
No
M.
cutt
ack
iH
aldar
etal
.,1996
Cyp
rinus
carp
ioL
.G
ills
Or
13–21.1
17.0
4)
94.9
–8.1
(6.4
8)
6.5
–13.0
(8.6
4)
91.6
–4
(2.8
)
Abse
nt
Abse
nt
5–8
–Y
es
M.
cyli
ndri
cus
(Sar
kar
etal
.,1985)
Gupta
&K
her
a,1988(a
)
Channa
gach
ua
Ham
.
Kid
ney
WB
12.8
–16.3
2
(14.3
7)
94.4
8–6.4
(4.9
1)
4–5.2
(4.4
)9
1.1
2–2.2
4
(1.6
8)
Abse
nt
Abse
nt
––
Yes
M.
dasg
upti
Hal
dar
etal
.,1996
Mugil
tade
Fors
skal
Gil
ls,
body
musc
les
Or
11.4
–19.5
(14.1
)9
4.9
–8.1
(6.4
)
7.3
–11.4
(9.1
1)
91.6
–4
(2.6
)
Pre
sent
Abse
nt
––
Yes
M.
deb
santu
sB
asu
etal
.,2009
Catl
aca
tla
Ham
.9
Labeo
rohit
aH
am.
(hybri
d)
Tai
lfi
nW
B9
98.4
96.3
4.3
92.3
and
2.8
91.8
Pre
sent
89–10
inla
rger
PC
,4–5
in
smal
ler
–N
o
M.
der
mit
is(H
aldar
etal
.,1981)
Gupta
&K
her
a,1988(a
)[S
yn.
Myx
oso
ma
der
mati
sH
aldar
etal
.,
1981)]
Labeo
rohit
aH
am.
Sca
les
WB
,
Ker
9–11
(10.3
)9
8–10
(9.4
)4–5
(4.4
)9
2–3
(2.2
)P
rese
nt
Abse
nt
60.5
–1
mm
Yes
M.
dhanach
andi
Ban
dyopad
hyay
etal
.,2007
Channa
ori
enta
lis
Sch
nei
der
Dors
al,
ven
tral
and
caudal
fins
Man
18.4
(17.0
–19.5
)9
6.0
5
(5.1
–6.8
)
7.7
–8.5
(8.2
6)
90.8
–1.7
(1.4
3)
Abse
nt
Abse
nt
–270
Yes
M.
duoden
ali
sK
aur
&S
ingh,
2011(e
)W
all
ago
att
uB
leek
er
Inte
rnal
inte
stin
al
wal
l
Pun
99
3.2
4.7
09
1.7
6an
d
2.7
69
1.0
6
Abse
nt
Abse
nt
6–7
inla
rger
PC
,not
vis
ible
in
smal
ler
40–50
No
M.
edel
lae
Sar
kar
,1999
Cte
nophary
ngodon
edel
laH
am.
Kid
ney
sW
B10.5
–11.5
(10.7
)9
9–10
(9.4
)
5.5
–6.5
(6.1
4)
92.5
–3.5
(3.1
2)
Abse
nt
Abse
nt
––
Yes
M.
eeli
Man
dal
&N
air,
1975
Mast
ace
mbel
us
arm
atu
sL
ac.
Inte
rnal
inte
stin
al
wal
l
WB
10.5
–14
98.2
–11.5
4.5
–5.5
92.5
–3
Abse
nt
2–3
6–
Yes
M.
eira
siK
aur
&S
ingh,
2009
Cir
rhin
am
rigala
Ham
.
Cau
dal
fin
Pun
8.6
96.7
3.2
91.5
7A
bse
nt
Abse
nt
3–4
7–100
990–300
Yes
M.
ence
phali
cus
Musl
ow
,1911
[Syn.
Len
tosp
ora
ence
phali
caM
usl
ow
,
1911]
Labeo
bata
Ham
.B
rain
,
blo
od
ves
sels
Or
6.2
–12.6
95.8
–10.6
––
––
35–110
–
Syst Parasitol (2012) 81:17–37 21
123
Ta
ble
1co
nti
nu
ed
Spec
ies
Host
(s)
Sit
eof
infe
ctio
n
Loca
lity
Spore
size
PC
size
(s)
ICP
PF
FC
Cyst
size
PC
s
equal
M.
epis
quam
mali
sE
gusa
etal
.,
1990
Mugil
cephalu
sL
.,L
iza
macr
ole
pis
Sm
ith
Gil
ls,
vis
cera
l
per
itoneu
m,
inte
stin
e,
scal
es
AP
,O
r6–7.6
(7.2
)9
5.1
6–5.8
4
(5.6
)
3.2
–3.6
(3.4
5)
92–2.8
(2.6
)an
d3.2
–3.6
(3.4
5)
91.4
–1.6
(1.5
)
Pre
sent
Abse
nt
4–6
–N
o
M.
esom
i(K
alav
ati
&
Nar
asim
ham
urt
i,1984
)
Lan
dsb
erg
&L
om
,1991
[Syn.
Rudic
apsu
laes
om
iK
alav
ati
&N
aras
imham
urt
i,1984
]
Eso
mus
sp.
Cau
dal
fin
AP
11.2
–12.4
(11.8
)9
6.8
–7.2
(7.0
)
4.8
–5.2
(5.2
)9
3–3.2
(3.1
6)
and
1.6
–2
91.0
Abse
nt
Abse
nt
9–10
in
larg
er
PC
2–3
mm
No
M.
etro
pli
Raj
endra
net
al.,
1998
Etr
oplu
ssu
rate
nsi
sB
loch
Bulb
us
arte
riosu
s
TN
10.2
–14.5
(12.2
)9
8.5
–11.9
(10.5
)
3.8
–6
(5.0
)9
1.7
–3.4
(2.4
)
Abse
nt
Abse
nt
4–5
0.1
–2
mm
Yes
M.
fila
men
tosu
s(H
aldar
etal
.,
1981)
Gupta
&K
her
a,
1988(a
)[S
yn.M
yxoso
ma
fila
men
tosa
Hal
dar
etal
.,1981
]
Punti
us
fila
men
tosu
sV
al.
Bra
inca
rtil
age
WB
11.2
–17.3
(13.7
)9
8.1
–12.2
(9.5
)
4–7.1
(3.6
)9
2–4
(3.1
)A
bse
nt
Abse
nt
5–6
–N
o
M.
foss
ilii
(Gupta
etal
.,2001)
Kal
avat
i&
Nan
di,
2007
[Syn.
Myx
oso
ma
foss
ilii
Gupta
etal
.,
2001
]
Het
eropneu
stes
foss
ilis
Blo
ch
Bra
inca
rtil
age
UP
9–12
(10.2
5)
96.2
–8.5
(7.4
4)
4.5
–7
(6.4
3)
91.7
5–2.7
5
(2.2
)
Abse
nt
Abse
nt
––
Yes
M.
ganguly
i(S
arkar
etal
.,1982)
Gupta
&K
her
a,1988(a
)[S
yn.
Myx
oso
ma
ganguli
iS
arkar
etal
.,1982
]
Sil
lago
macu
lata
Quoy
&G
aim
ard
Car
tila
ge
of
hea
d
skel
eton
And
8–10.5
(8.7
3)
94.8
–6.5
(5.0
3)
2.8
–4.6
(3.6
)9
1.5
–2.5
(1.9
)an
d2.2
–3
(2.4
)9
1.3
–2
(1.5
)
Abse
nt
Abse
nt
5–6
100
No
M.
goen
sis
Eir
as&
D’S
ouza
,
2004
Mugil
cephalu
sL
.G
ill
raker
sG
oa
9.5
–10.5
(9.7
)9
6.0
–7.5
(6.6
)
4.5
–6
(5.3
)9
2–3
(2.4
)
and
2–3
(2.4
)9
1.5
–2
(1.8
)
Abse
nt
65
in larg
er
PC
,3
in smal
ler
80–120
9280–360
No
M.
gorh
am
iG
upta
&K
her
a,1991
Catl
aca
tla
Ham
.,T
or
puli
tora
Ham
.
Gil
ls,
oper
culu
m
Pun
11
(10–12)
910.0
3
(10–10.5
)
4.5
–5
(4.8
)9
3–4
(3.5
5)
Abse
nt
53–4
–Y
es
M.
hald
iH
eman
anda
etal
.,2006
Cir
rhin
am
rigala
Ham
.G
ill
fila
men
tsM
an–
––
––
––
M.
hald
ari
Gupta
&K
her
a,
1989(a
)
Cir
rhin
am
rigala
Ham
.,
Labeo
dyo
chei
lus
McC
lell
and,
L.
rohit
aH
am.,
L.
bata
Ham
.
Fin
san
dgil
lsP
un
9–10
(9.3
1)
97.0
–8.5
(7.9
5)
4–5
(4.3
1)
92.5
–3
(2.9
7)
and
2.5
–3
(2.9
5)
91.5
–2
(1.9
8)
Pre
sent
Abse
nt
–152–168
9102–532
No
M.
hari
kensi
sK
aur
&S
ingh,
2011(c
)
Cir
rhin
am
rigala
Ham
.C
audal
fin
(bet
wee
nfi
n
rays)
Pun
10.1
98.5
4.8
–5.2
(5.0
)9
2.6
–3.6
(3.1
)an
d1–2.4
(1.7
)9
0.8
–2
(1.4
)
Abse
nt
Abse
nt
4–5
in
larg
er
PC
,
2–3
in
smal
ler
600–800
No
22 Syst Parasitol (2012) 81:17–37
123
Ta
ble
1co
nti
nu
ed
Spec
ies
Host
(s)
Sit
eof
infe
ctio
n
Loca
lity
Spore
size
PC
size
(s)
ICP
PF
FC
Cyst
size
PC
s
equal
M.
hosa
durg
ensi
sS
eenap
pa
&M
anohar
,
1981
Cir
rhin
am
rigala
Ham
.
Gil
ls,
musc
les
Kar
9–11
(10.5
)9
5–8
(6.2
5)
4–6
(5.3
7)
92–3
(2.3
)
and
2.5
–4
(3.3
)9
1–2.5
(1.4
3)
Pre
sent
Abse
nt
––
No
M.
hyd
erabaden
sis
(Lal
itha
Kum
ari,
1969)
Gupta
&K
her
a,1988(a
)[S
yn.
Myx
oso
ma
hyd
erabaden
seL
alit
ha
Kum
ari,
1969]
Barb
us
pin
naura
tus
Day
,P
unti
us
fila
men
tosu
sV
al.
Gil
l fila
men
ts
AP
,K
er9.3
–11.5
(10.1
)9
5–8
(5.9
)
5–7.3
(5.8
)9
1.4
–3
(2.2
)A
bse
nt
4–6
8–9
–Y
es
M.
india
e(L
alit
ha
Kum
ari,
1969)
Gupta
&K
her
a,1988(a
)[S
yn.
Myx
oso
ma
india
eL
alit
ha
Kum
ari,
1969]
Barb
us
sara
na
Ham
.
Gil
l fila
men
ts
AP
12.4
–15
(13.7
)9
6.4
–8.6
(7.3
)
5.7
–7.1
(5.9
)9
1.4
–2.5
(2.1
)an
d5–6.4
(5.2
)9
1.4
–2.5
(2.1
)
Abse
nt
78–10
1–2
mm
No
M.
indic
us
Tri
pat
hi,
1952
Cir
rhin
am
rigala
Ham
.
Musc
les,
inte
stin
al
wal
l,li
ver
WB
9.5
–10.8
97.5
–8.2
2.7
–3.6
91.8
and
1.8
91
Abse
nt
Abse
nt
––
No
M.i
ndir
ae
(Kundu,1985
)G
upta
&K
her
a,
1988(a
)[S
yn.
Myx
oso
ma
indir
ae
Kundu,
1985]
Cir
rhin
am
rigala
Ham
.
Hea
d
cart
ilag
e,
scal
esof
tail
fin
WB
11–14
(12.6
)9
9–11
(9.6
)
4–6
(4.7
)9
2–2.5
(2.2
)A
bse
nt
Abse
nt
8–10
0.5
–1
mm
No
M.
kalm
ani
Kau
r&
Sin
gh,
2011(d
)C
irrh
ina
reba
Ham
.
Gil
l lam
ella
e
Pun
9.5
–10.5
(10.0
)9
4.2
–5.2
(4.7
)
3–3.8
(3.4
)9
1.3
–1.9
(1.6
7)
Pre
sent
(sm
all)
Abse
nt
6–7
–Y
es
M.
kanja
liK
aur
&S
ingh,
2011(b
)C
irrh
ina
mri
gala
Ham
.
Sca
les
Pun
9.5
97.7
4.8
91.8
Abse
nt
Abse
nt
6–7
1–2
mm
Yes
M.
karn
ata
kae
(Hag
argi
&A
moji
,1981
)
Lan
dsb
erg
&L
om
,1991
[Syn.
Myx
oso
ma
Kar
eH
agar
gi
&A
moji
,
1981]
Barb
us
chola
Ham
.&
Buch
.
Cau
dal
musc
les
Kar
16.3
–19.0
4
(17.5
8)
910.8
8–13.6
(11.1
)
7.3
–10.8
8
(11.1
1)
93.5
8–5.4
4
(4.7
8)
Abse
nt
Abse
nt
6–7
–N
o
M.
koli
Lal
itha
Kum
ari,
1969
Barb
us
kolu
sS
kyes
,
Punti
us
fila
men
tosu
sV
al.
Dors
alfi
n,
ven
tral
fin
Ker
7.1
–9.6
(8.4
)9
5–6.4
(6.0
)
3.9
–4.6
(4.3
)9
2.1
–3.1
(2.8
)an
d1.4
–2.1
(2.0
)9
0.7
–1.4
(1.2
)
Abse
nt
6–8
5–6
–N
o
M.
labeo
sus
Sar
kar
,1995
Labeo
fim
bri
atu
sB
loch
Mes
ente
ries
nea
r
sple
en
WB
8–10
(9.2
5)
96.5
–9
(7.6
2)
6–6.5
(6.1
5)
92–4
(2.7
)
and
3–4.5
(4.0
2)
91.8
–2.5
(2.3
3)
Abse
nt
Abse
nt
5–7 an
d
3–5
–N
o
M.
lali
thae
Gupta
&K
her
a,1988(d
)L
abeo
calb
asu
Ham
.
Gil
ls(d
ista
l
lip
of
fila
men
ts)
Pun
9–11
(10.0
)9
8–9
(8.4
)5–6
(5.8
)9
3–3.5
(3.0
5)
and
4–0
(4.8
)9
2.5
–3
(2.8
5)
Pre
sent
Abse
nt
––
No
M.
liza
e(N
aras
imham
urt
i&
Kal
avat
i,
1979)
Lan
dsb
erg
&L
om
,1991
[Syn.
Myx
oso
ma
lair
di
Nar
asim
ham
urt
i&
Kal
avat
i,1979
]
Liz
am
acr
ole
pis
Sm
ith
Inte
stin
al
wal
l
AP
4.6
–5.2
(4.9
)9
9–9.5
(9.3
)
3–4.6
(3.2
)9
1.8
–2.6
(2.0
)
Abse
nt
Abse
nt
5–7
1.5
–3
mm
Yes
Syst Parasitol (2012) 81:17–37 23
123
Ta
ble
1co
nti
nu
ed
Spec
ies
Host
(s)
Sit
eof
infe
ctio
n
Loca
lity
Spore
size
PC
size
(s)
ICP
PF
FC
Cyst
size
PC
s
equal
M.
ludhia
nae
Gupta
&K
her
a,
1991
Labeo
dyo
chei
lus
McC
lell
and
Fin
sP
un
10
(10.0
)9
9.2
(9.0
–9.5
)5
93
Abse
nt
Abse
nt
–0.6
–1.2
90.5
–0.7
mm
Yes
M.
macr
ole
pi
Pad
ma
Doro
thy
&
Kal
avat
i,1992
Liz
am
acr
ole
pis
Sm
ith
Inte
stin
eA
P,
Ker
5.2
–6.9
(6.2
8)
94.3
–6.9
(5.2
6)
1.7
2–3.4
4
(2.7
8)
91.7
2–2.5
8
(2.0
)
Abse
nt
Abse
nt
3–4
188–200
Yes
M.
magauddi
(Baj
pai
etal
.,1981)
Gupta
&K
her
a,1988(a
)[S
yn.
Myx
oso
ma
magauddi
Baj
pai
etal
.,1981
]
Tri
chogast
erfa
scia
tus
Sch
nei
der
&
Blo
ch
Gil
lsW
B10.8
–11.7
(11.2
)9
8.3
––10
(9.2
)
3.3
–5
(4.0
)9
2.5
–3.3
(3.0
)an
d2.5
–3.3
(2.6
)9
1.7
–2.5
(1.8
)
–A
bse
nt
6–7
–N
o
M.
maguri
iS
arkar
,1993
Cla
rias
magur
L.
Acc
esso
ry
resp
irat
ory
org
an
WB
13–15
(14.1
3)
96.5
–8
(7.7
5)
7–8
(7.5
3)
92–3
(2.3
4)
Abse
nt
Abse
nt
7–9
-Y
es
M.
mahen
dra
eS
arkar
,1986
Catl
aca
tla
Ham
.G
ill
arch
epit
hel
ium
,
bucc
o-
phar
yngea
l
cavit
y
WB
,
AP
,
Pun
11.5
2–13.9
6
(12.7
)9
9.7
7–10.4
7
(9.2
)
6.2
8–7.3
3
(6.9
8)
93.4
9–4.1
9
(3.7
3)
and
4.1
9–6.9
8
(5.4
4)
93.1
4–3.4
9
(3.4
2)
Abse
nt
Abse
nt
6–7
in
PC
,
4–5
in
smal
ler
2–3
mm
No
M.
manora
mae
Bas
u&
Hal
dar
,
2002(b
)
Catl
aca
tla
Ham
.9
Labeo
rohit
aH
am.
(hybri
d)
Tai
lfi
nW
B11.8
(10.7
–13.1
)9
5.6
(4.8
–6.3
)
5.5
–6.6
(6.2
)9
2.0
–2.5
(2.2
)
Abse
nt
Abse
nt
5–6
in
larg
er
PC
,
4–5
in
smal
ler
200–310
Yes
/
No
M.
maru
lien
sis
(Sar
kar
etal
.,
1985)
Gupta
&K
her
a,
1988(a
)[S
yn.
Myx
oso
ma
maru
lien
sis
Sar
kar
etal
.,1985
]
Channa
maru
lius
Ham
.
Kid
ney
sW
B12–17.5
(14.3
7)
92.5
–5
(4.2
3)
7.5
–13
(10.0
5)
91–2
(1.5
5)
Abse
nt
Abse
nt
9–11
–N
o
M.
math
uri
Jayas
riet
al.,
1981
Punti
us
sara
na
Ham
.
Gil
lsR
aj8.7
–23.5
95.1
–10.1
2.7
–11.9
91.8
–4.6
and
2.7
–7.8
91.8
–4.6
Abse
nt
Abse
nt
8–9
in
larg
er
PC
,
3–4
in
smal
ler
–N
o
M.
meg
lits
chus
Sar
kar
,1996
Noto
pte
rus
noto
pte
rus
Pal
las
Lam
ella
r
epit
hel
ium
of
gil
ls
WB
8–9.5
(8.8
)9
6–7.5
(6.9
6)
3.5
–5
(4.4
2)
91.5
–3
(2.0
7)
Pre
sent
55–7
90–100
945
Yes
M.
meh
lhorn
iK
aur
&S
ingh,
2011(a
)
Cir
rhin
am
rigala
Ham
Gil
lla
mel
lae
Pun
8.9
96.8
3.7
92.5
and
2.6
91.5
Abse
nt
Abse
nt
4–6
–N
o
M.
mola
eS
arkar
,1993
Am
bly
phary
ngodon
mola
Ham
.
Kid
ney
sW
B8.5
–9.5
(9.0
)9
7–8.5
(7.4
)
5.0
–6.5
(6.0
)9
3.8
–4.5
(4.0
)
Pre
sent
Abse
nt
4–6
–Y
es
M.
mri
gala
eC
hak
ravar
ty,
1939
Cir
rhin
am
rigala
Ham
.
Sca
les
WB
17.2
1–8.2
4in
dia
m.
5.1
59
3.0
9an
d
3.0
99
2.0
6
Abse
nt
Pre
sent
–0.7
5–1.5
90.7
5–1
mm
No
24 Syst Parasitol (2012) 81:17–37
123
Ta
ble
1co
nti
nu
ed
Spec
ies
Host
(s)
Sit
eof
infe
ctio
nL
oca
lity
Spore
size
PC
size
(s)
ICP
PF
FC
Cyst
size
PC
s
equal
M.
mri
galh
itae
Bas
u&
Hal
dar
,2003(b
)
Cir
rhin
am
rigala
Ham
.9
Labeo
rohit
aH
am.
(hybri
d)
Gil
lsW
B10.8
–11.3
(10.8
)9
7.6
–8.1
(7.9
)
4.3
–5.2
(4.8
)9
2.7
–3.2
(2.1
)
and
2.9
–3.2
(3.0
)9
2–3.2
(2.1
)
Pre
sent
2–5
5–6
inla
rger
PC
3–4
in
smal
ler
–N
o
M.
mugce
phalu
s(N
aras
imham
urt
iet
al.,
1980)
Lan
dsb
erg
&
Lom
,1991
Mugil
cephalu
sL
.G
ill
fila
men
tsA
P,
Or
4.8
–5.2
(5.1
)in
dia
m.
1.6
–2
(1.8
)9
1–1.2
(1.1
)A
bse
nt
Abse
nt
5–6
0.5
–1
mm
Yes
M.
mugil
liH
aldar
etal
.,
1996
Mugil
cephalu
sL
.G
ills
Or
8.1
–16.3
(11.7
)9
4–7.3
(5.5
)
2.4
–8.1
(6.1
)9
1.6
–4
(2.7
)A
bse
nt
Abse
nt
––
Yes
M.
mult
ivader
isM
ukhopad
hyay
&
Hal
dar
,1998
Catl
aca
tla
Ham
.G
ills
,pal
ate,
inte
stin
e,bra
in
tiss
ue
WB
5.5
2–8.9
89
3.4
4–6.7
74.8
4–4.9
91.2
–3.3
and
1.1
4–2.5
79
1.1
2–1.7
2
Abse
nt
Abse
nt
8in
larg
er
PC
,6
in
smal
ler
–N
o
M.
mys
tusi
us
Sar
kar
,
1986
Mys
tus
vitt
atu
sB
loch
Sca
les
WB
12.2
3–13.9
6
(13.1
8)
98.7
3–10.4
7
(9.3
9)
6.2
8–7.6
8
(7.1
2)
92.7
9–4.1
9(3
.59)
and
3.4
9–4.1
9
(3.8
)9
1.4
–1.7
5(1
.5)
Abse
nt
Abse
nt
8–9
inla
rger
PC
,3–4
in
smal
ler
–N
o
Myx
obolu
ssp
.of
Pad
ma
Doro
thy
&K
alav
ati
(1992)
Liz
am
acr
ole
pis
Sm
ith,
Mugil
cephalu
sL
.
Inte
stin
eA
P6.9
–8.6
(7.7
4)
95.6
–6.5
(6.0
2)
2.6
–3.5
(3.3
2)
91.7
–2.6
(2.0
6)
Abse
nt
Abse
nt
5–6
–Y
es
M.
nain
iK
aur
&S
ingh,
2008
Cir
rhin
am
rigala
Ham
.
Gil
lsP
un
12.9
98.2
4.9
93.1
and
3.3
39
1.6
3P
rese
nt
Abse
nt
6–7
inla
rger
PC
,4–5
in
smal
ler
–N
o
M.
nara
sii
(Nar
asim
ham
urt
i,
1970)
Lan
dsb
erg
&
Lom
,1991
[Syn.
Myx
oso
ma
inte
stin
ali
sN
aras
imham
urt
i,1970]
Mugil
waig
ensi
sQ
uoy
&G
aim
ard
Inte
stin
eA
P12.5
–13.5
(12.8
)9
8.6
–9.5
(9.0
)
2.9
–3.6
(3.2
)9
1.6
–1.8
(1.7
)A
bse
nt
Abse
nt
4–5
2m
mY
es
M.
noble
i(S
arkar
,1982
)
Gupta
&K
her
a,
1988(a
)[S
yn.
Myx
oso
ma
noble
iS
arkar
,1982]
Ophio
cephalu
sst
riatu
sB
loch
Gal
lbla
dder
WB
11.5
–14.3
(12.7
)9
8.3
–10.5
(9.5
)
5.8
–8.5
(7.0
)9
3–4.5
(3.7
)A
bse
nt
Abse
nt
–121
998
Yes
M.
nodula
ris
South
wel
l
&P
rash
ad,
1918
Rasb
ora
danoco
niu
sH
am.
Fin
,m
usc
les,
pan
crea
s,
inte
stin
e,gal
l
bla
dder
Ker
6–12
96–9
1.8
5–4.5
90.7
5–3.0
Abse
nt
Abse
nt
3–5
260–450
Yes
M.
opth
alm
usc
ula
taB
asu
&H
aldar
,2002(a
)
Cir
rhin
am
rigala
Ham
.
Eye
musc
les
WB
12.4
–13.9
7
(13.1
3)
97.4
5–9.1
8
(8.0
4)
4.9
–6.1
2(5
.47)
92.6
5–3.3
7
(3.0
6)
and
2.3
5–3.3
7
(3.0
3)
91.8
4–2.1
4(1
.99)
Abse
nt
Abse
nt
7–9
inla
rger
PC
,2–3
in
smal
ler
0.6
7–1.0
mm
No
Syst Parasitol (2012) 81:17–37 25
123
Ta
ble
1co
nti
nu
ed
Spec
ies
Host
(s)
Sit
eof
infe
ctio
n
Loca
lity
Spore
size
PC
size
(s)
ICP
PF
FC
Cyst
size
PC
s
equal
M.
ori
ssae
Hal
dar
etal
.,1996
Cir
rhin
am
rigala
Ham
.
Gil
lsO
r13.0
–19.5
(15.7
1)
94.9
––8.1
(6.8
)
7.3
–11.8
(8.8
)9
2.4
–3.2
(1.7
8)
and
6.5
–11.4
(7.5
8)
91.6
–3.4
(2.5
7)
Pre
sent
Abse
nt
––
No
M.
osm
ania
eL
alit
ha
Kum
ari,
1969
Barb
us
punja
ben
sis
Day
Liv
er,
inte
stin
e
AP
12.4
–15
(13.5
)9
7.1
–10
(8.6
)
5–7.1
(5.6
)9
2–3.9
(3.2
)an
d
2.1
–3.6
(2.6
)9
1.4
–2.9
(2.5
)
Abse
nt
8–10
5–6
1–1.5
mm
No
M.
pars
iD
as,
1996
Liz
apars
iaH
am.
Gil
lsW
B9–9.5
(9.1
)9
8–8.5
(8.1
)4–5
(4.4
)9
2.5
–3
(2.8
)A
bse
nt
Abse
nt
5–
No
M.
parv
us
Shulm
an,
1962
Mugil
cephalu
sL
.G
ills raker
s
Goa
6–7
(6.3
)9
4.5
–6
(4.8
)3–4
(3.6
)9
1.5
–2
(1.6
)–
Abse
nt
6–7
40–50
9640–280
Yes
M.
pati
ale
nsi
sK
aur
&S
ingh,
2011(e
)
Labeo
rohit
aH
am.
Cau
dal
fin
Pun
11.2
89
6.6
74.8
93.1
and
1.7
09
1.5
1A
bse
nt
4–5
5–6
inla
rger
PC
,
2–3
insm
alle
r
0.8
–1
mm
No
M.
pin
naura
tiL
alit
ha
Kum
ari,
1969
Barb
us
pin
naura
tus
Day
Gil
l fila
men
ts
AP
8–11.4
(9.6
)9
6.5
–7.9
(7.0
)
3.6
–6.4
(4.4
)9
1.1
–2.1
(1.9
)
and
2.9
–5
(3.1
)9
1.1
–2.1
(1.6
)
Abse
nt
Abse
nt
––
No
M.
pota
ili
Lal
itha
Kum
ari,
1969
Labeo
pota
ilS
kyes
Liv
er,
inte
stin
e
AP
7.2
95.4
3.3
92
Pre
sent
Pre
sent
3–
Yes
M.
psi
lorh
ynch
iL
alit
ha
Kum
ari,
1969
Psi
lorh
ynch
us
bali
tora
Ham
.
Gil
l fila
men
ts
AP
9.3
–10.7
(10.0
)9
8.6
–10
(9.4
)
4.3
–5.7
(4.8
)9
2.9
–3.9
(3.2
)
and
2.9
–4.3
(3.5
)9
2.1
–3
(2.5
)
Pre
sent
3–4
6–7
0.5
–1
mm
No
M.
punja
ben
sis
Gupta
&K
her
a,
1989(b
)
Labeo
dyo
chei
lus
McC
lell
and
Fin
s,gil
lsP
un
10–12
(10.8
6)
910–11
(10.6
4)
5–6
(5.7
2)
and
3.9
–5.0
Abse
nt
Abse
nt
–76–133
No
M.
punja
bii
Kau
r
&S
ingh,
2010(c
)
Labeo
rohit
aH
am.
Cau
dal
fin
Pun
8.9
96.9
4.4
92.7
and
3.2
91.7
Abse
nt
Abse
nt
6–7
inla
rger
PC
,
4–5
insm
alle
r
0.0
89
1.0
mm
No
M.
punti
usi
Shee
ja
&Ja
nar
dan
an,
2006
Punti
us
tict
opunct
atu
sD
ay
Hea
rtw
all
Ker
13.5
–16.5
(15.0
)9
10.5
–12
(11.0
2)
6.7
5–7.5
(7.0
8)
93–4.5
(3.6
7)
and
5.2
5–6.7
5(5
.7)
Pre
sent
12
7–9
inla
rger
PC
,
5–7
insm
alle
r
249.6
–296.4
9109.2
–140.4
(269.1
9120.9
)
No
M.
rew
ansi
sS
rivas
tava,
1979
Cir
rhin
am
rigala
Ham
.
Sca
les
MP
9.6
69
8.0
54.8
29
3.2
Pre
sent
Abse
nt
–2.0
mm
Yes
M.
rigid
aS
arkar
,
1999
Am
bly
phary
ngodon
mola
Ham
.
Kid
ney
sW
B10–11
(10.8
)9
7.5
–9.5
(8.6
)
5.5
–7
(6.3
)9
3–4
(3.6
)P
rese
nt
Abse
nt
7–8
–Y
es
M.
roca
tlae
Bas
u
&H
aldar
,
2002(b
)
Catl
aca
tla
Ham
.9
Labeo
rohit
aH
am.
(hybri
d)
Gil
ls,
gut
WB
Spore
sw
ith
equal
PC
16.9
–19.3
(18.3
)9
5.6
–6.9
(6.0
);
spore
sw
ith
uneq
ual
PC
17.5
–19.3
(18.5
)9
5.6
–6.2
(5.9
)
Equal
PC
s10.4
–13.7
(12.6
)9
2.2
–3.1
(2.8
);
uneq
ual
PC
s11.8
–13.7
(12.9
)9
2.5
–3
(2.8
5)
and
10–12.2
(11.3
)9
2–2.4
(2.2
)
Abse
nt
Abse
nt
17–19
inla
rger
PC
,15–18
in
smal
ler
1100
9560
or
440
9218
Yes
/
No
26 Syst Parasitol (2012) 81:17–37
123
Ta
ble
1co
nti
nu
ed
Spec
ies
Host
(s)
Sit
eof
infe
ctio
n
Loca
lity
Spore
size
PC
size
(s)
ICP
PF
FC
Cyst
size
PC
s
equal
M.
rohit
ae
Hal
dar
etal
.,
1983
Labeo
rohit
aH
am.,
L.
dyo
chei
lus
McC
lell
and,
L.
bata
Ham
.,L
.calb
asu
Ham
.,P
unti
us
sara
na
Ham
.
Sca
les
WB
,
Pun
9.9
–12.1
(10.6
)9
8.8
–9.9
(9.0
)
6.6
93.3
Abse
nt
Abse
nt
5–6
200–300
Yes
M.
ropari
Kau
r&
Sin
gh,
2011(b
)
Cir
rhin
am
rigala
Ham
.G
ill fila
men
ts
Pun
12.5
89
4.5
4.9
69
1.5
0P
rese
nt
Abse
nt
9–10
–Y
es
M.s
ara
nae
Gupta
&K
her
a,
1990
Punti
us
sara
na
Ham
.,L
abeo
calb
asu
Ham
.
Gil
lsP
un
6–9
(7.7
2)
96–7
(6.2
)4–5
(4.2
4)
92.5
–4
(3.0
4)
and
1.5
–3
(1.9
8)
91–2
(1.3
)
Abse
nt
Abse
nt
–1.7
85–2.8
50
9
0.9
86–1.0
64
mm
No
M.
sara
nai
(Tri
pat
hi,
1952
)
Lan
dsb
erg
&L
om
,1991
[Syn.
Myx
obolu
sbra
nch
iali
sT
ripat
hi,
1952]
Barb
us
sara
na
Ham
.G
ills
WB
6.4
–7
94.5
–5
3.5
91.5
and
1.5
91
Abse
nt
Abse
nt
––
No
M.
sara
swati
iG
upta
&
Sar
asw
at,
1993
Cla
rias
batr
ach
us
L.
Kid
ney
sU
P10–20
97.8
–11
3.5
–9.5
92–3.5
Abse
nt
Abse
nt
––
Yes
M.
saugati
Kau
r&
Sin
gh,
2011(f
)
Labeo
rohit
aH
am.
Sca
les
Pun
8.3
96.6
49
2.4
Abse
nt
Abse
nt
4–5
1–1.5
mm
Yes
M.
scato
phagi
Hal
dar
etal
.,
1996
Sca
tophagus
arg
us
L.
Gil
lsO
r11.4
–17.9
(14.0
)9
4.0
–8.1
5.6
–9.8
(7.1
3)
91.6
–3.2
(2.3
6)
and
4.9
–8.1
(6.0
)9
1.6
–2.4
(2.0
)
Abse
nt
Abse
nt
––
No
M.
scle
rii
Kau
r&
Sin
gh,
2010(a
)
Catl
aca
tla
Ham
.E
ye
bal
l
(scl
era)
Pun
8.7
95.0
4.7
91.8
Pre
sent
Abse
nt
4–5
1–2
mm
Yes
M.
serr
atu
sP
agar
kar
&
Das
,1993
(em
end.)
Cyp
rinus
carp
ioL
.G
ill
arch
WB
8.5
–10
(9.0
7)
98–9
(8.5
)4–5
(4.4
7)
92.5
–3
(2.6
)A
bse
nt
–5–6
–Y
es
M.
sesh
adri
Lal
itha
Kum
ari,
1969
Labeo
fim
bri
atu
sB
loch
Gil
lsA
P11.4
–12.9
(12.2
1)
98.6
–10
(9.0
4)
5.7
92.9
–4.3
(3.6
6)
and
4.3
–5
(4.9
)9
2.5
–3.6
(2.9
9)
Pre
sent
5–
–N
o
M.
shanti
puri
Bas
u&
Hal
dar
,2002(b
)
Catl
aca
tla
Ham
.9
Labeo
rohit
aH
am.
(hybri
d)
Gil
l lam
ella
e
WB
Spore
sw
ith
equal
PC
6.3
–8.2
(7.3
)9
5.2
–6.1
(5.8
);
spore
sw
ith
uneq
ual
PC
6.2
–8.1
(7.4
)9
5.1
–6.2
(5.8
)
Equal
PC
s3.5
–4.1
(4.0
)9
2–2.8
(2.4
);uneq
ual
PC
s3.2
–4.2
(3.9
)9
2–3
(2.2
)an
d2.5
–4
(3.4
)9
1.9
–2.8
(2.2
)
Abse
nt
Abse
nt
4–5
100
990;
140
Yes
/
No
M.
shet
tiS
ennap
pa
&
Man
ohar
,1981
Cir
rhin
am
rigala
Ham
.G
ills
Kar
,
Ker
8–9
(8.8
)9
7–8
(7.4
)3–4
(3.4
)9
2–3
(2.3
)A
bse
nt
Abse
nt
––
Yes
M.
slen
dri
iK
aur
&S
ingh,
2010(b
)
Cir
rhin
am
rigala
Ham
.G
ills
Pun
14.8
79
3.4
5.7
49
1.4
8A
bse
nt
Abse
nt
6–8
–Y
es
M.
sophora
eJa
yas
ri,
1982
Punti
us
sophore
Ham
.G
ills
,
kid
ney
s
Raj
6.4
–26.6
(14.9
)9
5.9
–10.1
(7.7
)
–A
bse
nt
Abse
nt
––
No
M.
sphaer
ali
sP
adm
a
Doro
thy
&K
alav
ati,
1992
Liz
am
acr
ole
pis
Sm
ith
Gil
l fila
men
ts
AP
13–15
(14.1
3)
96.5
–8
(7.7
5)
7–8
(7.5
3)
92–3
(2.3
4)
Abse
nt
Abse
nt
4–5
96–215
980–165
Yes
Syst Parasitol (2012) 81:17–37 27
123
Ta
ble
1co
nti
nu
ed
Spec
ies
Host
(s)
Sit
eof
infe
ctio
nL
oca
lity
Spore
size
PC
size
(s)
ICP
PF
FC
Cyst
size
PC
s
equal
M.
spin
ocu
rvatu
raM
aeno
etal
.,1990
Mugil
cephalu
sL
.B
ranch
ial
cart
ilag
e,
mes
ente
ry,
bra
in,
sple
en
AP
10.6
–13.6
(11.5
)9
9.6
–11.2
(10.5
)
2.5
8–4.8
(3.8
)9
1.7
2–3.2
(2.6
)
Abse
nt
Abse
nt
4–6
–Y
es
M.
sush
mii
Kau
r&
Sin
gh,
2010(c
)
Labeo
rohit
aH
am.
Eye
bal
lP
un
9.6
98.3
4.7
93.2
and
3.1
92
Pre
sent
6–8
4–6
inla
rger
PC
,3–4
in
smal
ler
2–3
mm
No
M.
szek
eli
Kau
r&
Sin
gh,
2011(f
)
Wall
ago
att
uB
leek
er
Inte
rnal
wal
lof
stom
ach
Pun
8.7
94.1
4.4
91.5
Abse
nt
Abse
nt
5–6
600–800
Yes
M.
tict
oS
hee
ja&
Janar
dan
an,
2006
Punti
us
tict
opunct
atu
sD
ay
Gil
ls,
musc
les
inte
stin
e,li
ver
Ker
12.7
5–15
(14.5
5)
96.7
5–9
(7.8
)
4.5
–7.5
(6.6
3)
92.5
–3
(2.9
2)
Abse
nt
6–8
6–8
223–452
9215–405
(317–284)
Yes
M.
tric
hogast
eri
(Sar
kar
,
1985(b
))G
upta
&K
her
a,
1988(a
)[S
yn
.M
yxoso
ma
tric
hogast
eri
Sar
kar
,
1985(b
)]
Tri
chogast
erfa
scia
tus
Sch
nei
der
&
Blo
ch
Gal
lbla
dder
WB
14–17
(15.5
5)
98.7
–9.8
5
(9.3
5)
9–10.4
7(1
0.1
)9
3–3.8
4
(3.3
2)
Abse
nt
Abse
nt
5–6
–Y
es
M.
trip
ath
iiK
alav
ati
etal
.,1981
Cla
rias
sp.
Wal
lof
gut
and
vis
cera
lorg
ans
AP
9.8
–10.2
(10.1
)9
12.0
–13.5
(13.0
)
5–6
(5.5
)9
2.5
Abse
nt
Abse
nt
80.5
–1
mm
Yes
M.
undasu
tura
eS
arkar
,1994
Am
bly
phary
ngodon
mola
Ham
.
Uri
nar
ybla
dder
WB
10–11
(10.6
)9
5–6
(5.7
)5.0
5–6.5
(6.1
5)
92–2.5
(2.3
5)
and
3–3.5
(3.2
)9
1–1.2
(1.1
)
Abse
nt
Abse
nt
7–8
inla
rger
PC
,3–4
in
smal
ler
–N
o
M.
utl
ouen
sis
Hem
anan
da
etal
.,2009
Cla
rias
batr
ach
us
L.
Corn
eaM
an12.2
8
(11.9
–12.6
)9
8.9
2
(8.5
–9.2
)
1.7
–2.4
(2.3
4)
Pre
sent
Abse
nt
––
Yes
M.
vaniv
ilasa
eS
eenap
pa
&
Man
ohar
1980(b
)
Cir
rhin
am
rigala
Ham
.
Ben
eath
scal
esK
ar8–10
97–9
3–4
92.5
7(i
funeq
ual
,
smal
ler
2.9
89
2.5
)
Pre
sent
Abse
nt
–0.4
5–2.1
89
0.3
3–1.9
5m
m
Yes
/
No
M.
vari
form
is(H
aldar
etal
.,
1996)
Kal
avat
i&
Nan
di,
2007
[Syn.
Myx
obolu
sva
riabil
isH
aldar
etal
.,
1996]
Mys
tus
guli
oH
am.
Body
musc
les,
gil
ls
Or
13–17.9
(15.2
)9
4.9
–8.1
(5.6
5)
8.1
–11.4
(9.6
7)
91.6
–4
(2.8
4)
and
6.5
–11.4
(8.6
)9
1.6
–3.2
––
––
No
M.
vedava
tien
sis
See
nap
pa
&
Man
ohar
,1981
Cir
rhin
am
rigala
Ham
.
Gil
lsK
ar13–15
(13.8
)9
8–10
(9.2
)
6–7
(6.2
)9
3–4
(3.4
)
and
3–5
(3.9
)9
2–3
(2.6
)
Pre
sent
Abse
nt
––
No
28 Syst Parasitol (2012) 81:17–37
123
A revised key to the genera of the Myxosporea
A revised dichotomous key of the 59 genera of the
class Myxosporea is presented below. This new key
includes 10 additional genera erected since the
appearance of the key to this group given by Lom
& Dykova (1991). Furthermore, some genera e.g.
Pentacapsula Naidenova & Zaika, 1970, Hexacap-
sula Arai & Matsumoto, 1953 and Septemcapsula
Hsieh & Chen, 1984 have been synonymised and
their species assigned to Kudoa Meglitsch, 1947
based on phylogenetic analyses of their comparative
rDNA (Whipps et al., 2004). Likewise, Leptotheca
Thelohan, 1895 is no longer considered valid (Gunter
& Adlard, 2010) and its species have been transferred
to Ceratomyxa Thelohan, 1892 and Sphaerospora
Thelohan, 1892. Lastly, Davisia Laird, 1953 is
preoccupied by Davisia Del Guercio, 1909 (Insecta:
Hemiptera) and has been replaced by Myxodavisia
Zhao, Zhou, Kent & Whipps, 2008 by Zhao et al.
(2008).
1 Mature spore contains only one polar capsule
(PC)………..……………………………………2
– Mature spore contains more than one
PC……………………………………………….9
2 Spores pyriform, tear-shaped or elongate
ellipsoidal………………………...……………..3
– Spores rounded to spherical or ellipsoidal
……………………..……………………………7
3 Suture line of two shell valves sigmoidal
…………….…Coccomyxa Leger & Hesse, 1907
– Suture line of two shell valves straight or slightly
bent…………………..….………………………4
4 Spores with bifurcate caudal process…………………………………..Phlogospora Qadri, 1962(a)
– Spores without bifurcate caudal process…….…5
5 Spores with slightly bent sutural line; PC with
anterior recurved spine……..……………………………………......…Theohanelloid Sarkar, 2009
– Spores with straight sutural line; PC lacking
process(es)………………………………………6
6 Spores with PC discharging apically and axi-
ally……………….…Thelohanellus Kudo, 1933
– Spores with PC discharging subapically and to
one side…………………………..………………………....Neothelohanellus Das & Haldar, 1986
Ta
ble
1co
nti
nu
ed
Spec
ies
Host
(s)
Sit
eof
infe
ctio
n
Loca
lity
Spore
size
PC
size
(s)
ICP
PF
FC
Cyst
size
PC
s
equal
M.
venka
tesh
iS
eenap
pa
&M
anohar
,
1981
Labeo
bata
Ham
.,L
.ca
lbasu
Ham
.,L
.ro
hit
aH
am.,
Cir
rhin
am
rigala
Ham
.
Gil
ls,
ovar
y
Kar
,
Ker
,
Pun
9–10
(9.3
1)
97–8.5
(7.9
)
4–5
(4.3
1)
and
2.5
–3
(2.9
)9
2.5
–3
(2.9
)
and
1.5
–2
(1.9
)
Pre
sent
Abse
nt
––
Yes
M.
yogen
dra
i(T
ripat
hi,
1952
)
Lan
dsb
erg
&L
om
,1991
[Syn.
M.
spher
icum
Tri
pat
hi,
1952
]
Cir
rhin
am
rigala
Ham
.In
ner
side
of
scal
es
WB
9–9.5
97.2
2.8
–3.6
Pre
sent
––
–Y
es
Syst Parasitol (2012) 81:17–37 29
123
7 Spores club-like, with broad anterior region and
narrow posterior region……………………………………………Auerbachia Meglitsch, 1968
– Spores spherical…………………………..…….8
8 Spores with three unequal shell valves (two small
and one), difficult to discern using light micro-
scope; exclusively histozoic………………………………………………Unicapsula Davis, 1924
– Spores with two shell valves adhering to curved,
delicate sutural line; exclusively coelozoic
……..…..Globospora Lom, Noble & Laird, 1975
9 Mature spore with two polar capsules (one much
smaller than other)……………………………..10
– Mature spore with more than two polar
capsules……………………………..…………53
10 PCs widely separated……...……...……………11
– PCs close together……………………….……29
11 PCs located at ends of spindle-shaped or elongate
spore………………………..….............………12
– PCs not terminal at opposite ends, nor close to
each other…………...............…………………16
12 Polar filament tapers strongly from base to
tip, does form a regular coil within PC but is
rather folded and bends several times…………….....……………..Sphaeromyxa Thelohan, 1892
– Polar filament forms thin spiral tube within
capsule, of about same diameter throughout its
length……………………………..……………13
13 Spores generally with more or less pointed ends
and pyriform PCs…….Myxidium Butschli, 1882
– Spores ovoid, semicircular or slightly crescent-
shaped, with rounded or bluntly pointed
ends……………………………………………14
14 Spores slightly crescent-shaped, with elongated
PCs……Enteromyxum Palenzuela, Redondo &
Alvarez-Pellitero, 2002
– Spores ovoid to semicircular with spherical
PCs……………………………………….……15
15 Spores semicircular or slightly curved in valvular
view………………..Zschokkella Auerbach, 1910
– Spores ovoid……...………………Soricimyxum
Prunescu, Prunescu, Pucek & Lom, 2007
16 Spores elongate, with central transverse sutural
line; PCs near opposite ends, discharge later-
ally….……Fabespora Naidenova & Zaika, 1969
– Spores rounded, ovoid, spherical or pyramidal
with longitudinal sutural line…………….........17
17 PCs set widely apart in sutural plane; sutural line
straight………………………..............…..……18
– PCs in plane perpendicular to sutural line, sutural
line mostly sinuous……………………………22
18 Spores triangular, with anterior end wider, flat-
tened parallel to sutural plane……………………………………Triangula Chen & Hsieh, 1984(a)
– Spores spherical to ovoid……………….………9
19 Spores spherical or subspherical; in marine
fishes………....………Ortholinea Shulman, 1962
– Spores ovoid in valvular view, wider at anterior
end, flattened towards sutural plane…..........…20
20 Spores with sutural markings or projections.…21
– Spores without sutural markings or projection
………….....Neomyxobolus Chen & Hsieh, 1960
21 Spores ovoid, with sutural markings along pos-
terior border……………………………………………..Cardimyxobolus Ma, Dong & Wang, 1982
– Spores elongate, with pointed projections extend-
ing posteriorly…………………………………………….……Kentmoseria Lom & Dykova, 1995
22 Spores spherical or subspherical, with or without
any caudal or lateral appendages………………23
– Spores inversely pyramidal with or without
projections…………………………......………24
23 Spores spherical; each shell valve bears hollow,
mainly horn-like projection in about its centre
………...………………Myxodavisia Laird, 1953
– Spores spherical or subspherical, without append-
ages….............……….…Sinuolinea Davis, 1917
24 Spores inversely pyramidal, with rounded
outline…...............….Myxoproteus Doflein, 1898
– Spores inversely pyramidal, with pointed end
extending posteriorly and projections or keel-like
membranous extensions………………….........25
25 Anterior surface of each valve with wing-like
projection attached….....………………..………..
....Bipteria Kovaleva, Zubchenko & Krasin, 1983
– In addition to valvular projections, stiff, keel-like
membrane runs along sutural line…….………26
26 Keel-like membranous extensions run along
straight or oblique sutural line…………..….…27
– Keel-like membranous extensions run along sin-
uous or slightly wavy sutural line……….……28
27 Keel-like membranous extensions run along
straight sutural line………………………………Schulmania Kovaleva, Zubchenko & Krasin,
1983
– Keel-like membranous extensions run meridio-
nally along oblique sutural line……….…………………….…Paramyxoproteus Wierzbicka, 1986
30 Syst Parasitol (2012) 81:17–37
123
28 Keel-like membranous extensions run along
meridian of sinous sutural line……Neobipteria
Kovaleva, Gaevskaya & Krasin, 1986
– Keel-like membranous extensions run along
slightly wavy sutural line………………………...
…………………..………Noblea Kovaleva, 1989
29 Spores essentially bilaterally symmetrical…….30
– Spores asymmetrical……………………..……51
30 PCs spherical, tandem in position and distant
from anterior end; two fine projections at both
posterior and anterior ends of spindle-shaped
spore…………..…Neohenneguya Tripathi, 1953
– PCs spherical to subspherical, closely situated on
sutural plane…………...………………………31
31 Spores flat in median plane……………………....
……Gadimyxa Køie, Karlsbakk & Nylund, 2007
– Spores of various shape, with rounded or bluntly
pointed ends……………………………………32
32 Spores without projections………………...…..33
– Spores with projections…………………….….36
33 Shell valves generally smooth……….……..…34
– Shell valves smooth or ridged often with pos-
terior or lateral protuberances or bumps……….35
34 Sutural line straight...………………………….....
……………………......Myxobolus Butschli, 1882
– Sutural line distinctly sinuous…………………………………Spirosuturia Chen & Hsieh, 1984(b)
35 Spores with two uninucleate sporoplasms ………...………………Sphaerospora Thelohan, 1892
– Spores with many uninucleate sporoplasms ………………………………………Polysporoplasma
Sitja-Bobadilla & Alvarez-Pellitero, 1995
36 Spores with shape of double, flattened isosceles
triangle in sutural view……….………… ……37
– Spores rounded, ellipsoidal, ovoid, pyramidal,
spindle-shaped, crescent-shaped or V-shaped………………………………………………………39
37 Spores reniform, with laterally alate processes
adhering to all sides of valves…………………………………Renispora Renispora Kalavati,
Longshaw & MacKenzie, 1996
– Spores extremely elongate with respect to plane
perpendicular to straight, central sutural line ………………………………………………………38
38 Shell valves with wing-like membranous projec-
tions……………………………Alatospora Shul-
man, Kovaleva & Dubina, 1979
– Shell valve projections are doubled to form
parachute-like pockets……………………………Pseudoalatospora Kovaleva & Gaevskaya, 1983
39 Spores projections extend in various directions
…………………………………………………40
– Spores elongate, drawn out to enormous length……………………………………………………49
40 Spores with multiple projections………………41
– Spores with 1-4 projections……………………42
41 Fine ridges on surface of valves extend into
fringe-like processes at posterior end………………………………………Wardia Kudo, 1919
– Spores with numerous thread- or pin-like projec-
tions…………….Hoferellus Berg in Kudo, 1919
42 Spores with single caudal projection or enveloped
in membranous veil……………………………43
– Spores with more than one caudal projection
…………………………………………………44
43 Single caudal projection is not extension of shell
valve………………..……Unicauda Davis, 1944
– Spores enveloped in large membranous veil.……Palliatus Shulman, Kovaleva & Dubina, 1979
44 Shell valves often smooth or with ridges extend-
ing posteriorly into two caudal projections …..45
– Spores with two or more caudal projections which
extend in various directions….………… ……46
45 Spores with usually ridged shell valves which
extend into two caudal projections……………………………………...Myxobilatus Davis, 1944
– Spores with smooth shell valves which continue
as caudal, often slightly, divergent projections……………………..…Henneguya Thelohan, 1892
46 Spores with two caudal projections which extend
in opposite directions……………..…………………………...…Dicauda Hoffman & Walker, 1978
– Spores with two or four caudal projections which
extend posterolaterally………...………………47
47 Four caudal projections extend posterolaterally….………Tetrauronema Wu, Wang & Jiang, 1988
– Two caudal projections or filamentous processes
extend posterolaterally………………………...48
48 Two projections extend laterally from one side of
posterior end of spore……………………………………Laterocaudata Chen & Hsieh, 1984(b)
– Valves of broadly triangular spores drawn into
filamentous processes on each side; each pair
connected by filament…………………………
Syst Parasitol (2012) 81:17–37 31
123
………………...…Trigonosporus Hoshina, 1952
49 Spores crescent-shaped or arcuate…………………………………….Ceratomyxa Thelohan, 1892
– Spores oval to V-shaped………………………50
50 Spores V-shaped………………………………….
…………………..…Meglitschia Kovaleva, 1988
– Spores oval………….…Ellipsomyxa Køie, 2003
51 Spores ellipsoidal or spindle-shaped, with non-
axially attached caudal appendage………………Hennegoides Lom, Tonguthai & Dykova, 1991
– Spores elongate, with unequal shell valves meet-
ing at curved suture……………………………52
52 PCs two, located anteriorly but discharge laterally
………………….…Parvicapsula Shulman, 1953
– PCs four, arranged in cruciform shape at spore
apex …Neoparvicapsula Gaevskaya et al., 1982
53 Mature spore contains three PCs…...…………54
– Mature spore contains more than three PCs….55
54 PCs situated centrally, in form of triradiate star in
apical view……………...Trilospora Noble, 1939
– PCs situated peripherally and discharge later-
ally…………………...Trilosporoides Køie, 2005
55 Mature spore with two shell valves, contains four
PCs…………………………………………..…56
– Mature spore with four shell valves, contains
more than four PCs……………………………58
56 Spore spindle-shaped, with two shell valves but
without caudal appendage…………………………………………………Agarella Dunkerly, 1915
– Spore spherical or subspherical, with two shell
valves, with or without caudal appendage……57
57 Spores spherical, rarely with caudal projections
…….…………Chloromyxum Mingazzini, 1890
– Spores subspherical, with one or two caudal
projections …….……Caudomyxum Bauer, 1948
58 Spore sutural ridge extends anteriorly and poste-
riorly in form of spines…………………………………………Octospina Hsieh & Xiao, 1993
– Spore sutural ridge without spines……………………………..…………Kudoa Meglitsch, 1947
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