On the Nematode Genus Proteptus.

325

X X X I I I . - - O n the Nematode Genus Proleptus. B y H. A.

BAYLIS, M.A., D.Sc., Department of Zoology, British
Museum (Natural History).
THE genus Proleptus Dujardin, 1845, is not yet at all
well known. It is proposed in this paper to review
briefly what is known of the various species which have
been or m a y be attributed to it, adding a description
of one new species and some account of another species
(P. robustus) which has never been adequately described.
1. Proleptus acutus Dujardin, 1845.
S y n . ? Spiroptera dacnodes Creplln, 1851 ; ? Histiocephalus dacnode~
Molin, 1860 ; ? Spiropterina daenodes D i e s i n g , 1861.
Of this species, which is regarded as the genotype,
the original description is very brief and was based on a
single male specimen. No more recent description
of it appears to exist. The host from which Dujardin
recorded it was Raja clavata.
For measurements see table below (p. 333).
2. Proleptus obtusus Dujardin, 1845.
Syn. Coronilla scillicola v. B e n e d e n , 1871 ; Spiropterina scillicola
v. L i n s t o w , 1901 ; Proeptus scillicola L l o y d , 1920 ; ? Spiropterina
coronata v. B e n e d e n , 1858 (vide inJra).
This is the only well-known species of the genus.
Detailed accounts of it have been given b y v. Linstow
(1901), Seurat (1919), Lloyd (1920) and Mtiller (1925).
It is very common in the stomach and intestine of dog-
fish (Scyllium catulus, S. canicula) round the British
coasts and elsewhere. Every specimen of these two
species examined by the writer at Plymouth was more
or less heavily infested. P. obtusus has been recorded
by Taylor (1925) from Acanthias vulgaris off Ceylon
and from some South African host. Among the collection
of Nematodes belonging to the School of Public Health
and Tropical Medicine, Sydney, I~.S.W., the writer has
also found two sets of specimens of this species, collected
off South Africa b y Dr. H. P. Bayon. One of these
came from a " mole-shark " [? Rhinobatus], the other
from a " shark " off Robben Island. The latter specimens
are of exceptionally large size, the females measuring
up to 69 ram. in length, but otherwise there seems to be
nothing to distinguish them from European examples.
326 ])r. H. A. Baylis o~
Lloyd (1920) points out that there is great variation
in this species, not only in the actual measurements of
individuals but in the proportions of different parts.
Some of the measurements given in the recent descriptions
are summarized in the table below. A very characteristic
feature of P. obtusus is the shape of the right spicule, the
tip of which is bent towards the left, almost at right
angles to the main shaft. This has been more or less
clearly shown in some of the existing figures, but Mtiller
(1925) is almost the only author who has accurately
described it.
3. Proleptus inflatus (v. Linstow, 1890).
Syn. Spiropterina inflata v. L i a s t o w , 1890.
This species, which is recorded from Scyllium immoratu~n,
differs from P. obtusus, but rather closely resembles
P. acutus (according to Dujardin's description), in the
length of the spicules of the male. The size of the worms,
however, as given by v. Linstow, is nearly three times
that of P. acutus.
For measurements see table below.
4. Proleptus africanus (v. Linstow, 1899).
Syn. Spiropterina africana v. L i n s t o w , 1899.
This form, recorded from Anguilla sp. in South Africa,
may be provisionally referred to Proleptus. It differs
from any of the other known species in the lengths of the
spicules of the male.
For measurements see table below.
5. Proleptus robustus (v. Beneden, 1871) Seurat, 1916.
Syn. Coronilla robusta v. B e n e d e n , 1871 ; Spiropterina robusta
v. L i n s t o w , 1903 ; ? Coronilla minuta v. B e n e d e n , 1871 (vide infra).
This worm was recorded b y v. Beneden from Raja
circularis and R. clavata. He did not describe it, but
gave some figures from which it appears possible to
recognize the species. During a short stay at the
laboratory of the Marine Biological Association at Ply-
mouth in May 1932 the writer had the good fortune
on three occasions to find examples of a species of Pro-
leptus in the stomach of Raja clavata which, on comparison
with v. Beneden's figures, are considered to belong to this
species. Further, a few specimens obtained by Mr. L. A.
 the Ne,matode Genus Prolept.us. 327
H a r v e y from Mustelus loevis at Plymouth, and previously
recorded b y the writer (1928) as P. obtusus, prove to be
of the same species. They are, however, considerably
smaller than the specimens from Raja, and are not quite
mature.
Among the specimens from Raja, most of which are
immature, there are one mature male and two mature
females, but only the larger of these contains fully-
formed eggs. The male measures about 33 ram. in length
and 0"65 mm. in maximum thickness. The two females
measure respectively about 40 and 44 ram. in length
and 0"83 and 0'97 mm. in maximum thickness. The
Fig. 1.

Proleptus robustus. Anterior end of female ; dorsal view.

c.p., cervical papillse ; n.r., nerve-ring ; oes. 1, oes. 2, a n t e r i o r
a n d posterior portions of t h e oesophagus.

cuticular striations are about 4/~ apart. The anterior

end (fig. 1) is surrounded by a well-developed euticular
inflation. The lips are large and rounded, and each
bears on its inner surface a single conical tooth. The
anterior portion of the (esophagus is 0"6-0"65 mm. in
length (measured from the cephalic extremity). The
entire (esophagus, measured from the same point, is
6-6"7 mm. long. The cervical papillae, which are pointed
and bristle-like, and contained in small depressions in
328 Dr. H. A. Baylis on
the cuticle, are situated at 0"39-0"45 ram. from the anterior
end. The nerve-ring is situated at 0"42-0"5 ram., and
the excretory pore at 0"85-0"9 mm., from the cephalic
extremity.
The caudal end of the male (fig. 2) is curled ventrally.
The tail is about 0"8 mm. long. The caudal aloe are
very broad and give the posterior end a spoon-like shape.

Fig. 2.

Proleptus robustus. P o s t e r i o r end of male ; lateral view.

1., left spicule ; r., right spicule.

They contain eight pairs of rib-like papillm, three of which

are preanal and five postanal. Two pairs of subventral
papillm are also present, one just behind and one just
in front of the cloacal aperture. The right spicule is
broad at its base and tapers rapidly towards the tip,
which has a hyaline cap. I t measures about 0"4 mm.
in length. The left spicule is 1"5 mm. long, and has a
 tl,.e Nematode Genus Proleptus. 329

tubular shaft and a membranous ala which appears to

widen considerably and narrow again rather suddenly
a little before the tip. The cuticle of the ventral surface
in the caudal region is tessellated.
The caudal end of the female (fig. 3) is curved towards
the dorsal side. The tail is rather blunt and measures
0"42-0"44 mm. in length. It bears a pair of subterminal
papillm. The vulva is situated near the posterior end,
in the larger specimen at 0"36 mm. and in the smaller

:Fig. 3.

Proleptus robustus. P o s t e r i o r e n d of f e m a l e ; l a t e r a l v i e w .
a., a n u s ; i., i n t e s t i n e ; u., u t e r u s ; v., v u l v a .

at 0"5 ram. from the anus. The vagina, together with

the common trunk of the uterus, is very short (0"55-
0"7 ram.), and there is a muscular ovejector immediately
before the vulva. The two uterine branches run forward
parallel to each other. The eggs are roundish-ovoid,
measure on the average about 0"0450"032 mm.,
and contain coiled embryos when ready for laying. The
egg-shell is relatively very thick (about 6/~).
330 Dr. H. A. Baylis on

6. Proleptus australis, sp. n.

The hetminthological collection of the School of Public
Health and Tropical Medicine, Sydney, N.S.W., includes
some specimens of Proleptus from the intestine of a
" tiger-shark " (probably Galeocerdo tigrinus). These
were obtained off the Flinders Islands, North Queensland,
by Mr. Bassett-Hull, and appear to represent a new
species.
The male measures approximately 14-16 ram. in lengbh
and 0"46-0'48 ram. in maximum thickness, the female
about 18"5 mm. and 0"75 ram. respectively. The cuti-
cul~r striations are at intervals of about 4"5 ~. The

Fig. 4.

Proleptus australis. A n t e r i o r end of male ; lateral view.

anterior end (fig. 4) is, as usual, surrounded by a cuticular

inflation. Each lip, in addition to the usual median
lateral tooth, bears on its inner surface two pairs of
rather long, spine-like teeth, situated subdorsally and sub-
ventrally, and between these and the median tooth a
few small denticles. The anterior portion of the oeso-
phagus, measured from. the extremity of the head, is
about 0"5-0"6 mm. long in the male and 0"65 mm. in the
female. The entire (esophagus, measured from the same
point, is 2"4-2"7 mm. long in the male and 3"4 mm.
in the female. The nerve-ring is situated at 0"33-0"38 mm.,
 tl~e Nematode Genvs Proleptus. 331
and the excretory pore at 0"54-0"6 mm., from the anterior
extremity. The cervical papillee are at about the level
of the nerve-ring or immediately behind it.
The caudal end of the male (fig. 5) is spirally coiled,
and its ventral cuticle is tessellated. The tail is about
0 55 ram. long. The caudal alee are only moderately
developed. Nine pairs of papilke project into them.

Fig. 5.

r.
Proleptus a~istralis. Posterior end of male ; lateral view.
l., left spicule ; r., right spicule.

Of these the anterior seven are long and rib-like, and four
are preanal. The edges of the cloacal aperture appear
to be produced for some little distance as a delicate
membranous tube, through which the spicules m a y be
protruded. The right spicule is relatively stout, measures
0"26-0"27 ram. in length, and has a hyaline cap at the tip.
J u s t before the tip the abrupt termination of a mem-
branous ala sometimes produces the appearance of a
332 On the Nematode Genus Proleptus.
lateral barb. The left spicule is slender and about
1"2 ram. long. It has a rather narrow ala.
The caudal end of the female is curled towards the
dorsal side. The tail is blunt and about 0"4 ram. long.
The vulva is situated exceptionally far forward, at
8"2 mm. from the posterior end. It opens on the
posterior surface of a rounded prominence which appears
to be caused by the muscular ovejeetor. The vagina
runs posteriorly from the vulva, and after a course
of only 0"15 ram. gives off the two uterine branches,
which at first run parallel posteriorly. The eggs are
of very uniform size, measuring about 0"038 mm. x
0"027 ram. Their shells are relatively very thick (about
4"5 ~), and they contain coiled embryos when ready
for laying.
Co-type specimens of this species will be deposited
in the British Museum (Natural History) as well as in the
Sydney collection.

7. Proleptus coronatus (v. Beneden, 1858).

Syn. Spiropterina coronata v. Beneden, 1858.
This species is recorded from Scyllium canicula and
Raja radians. The male, according to van Beneden's
description, is 25-30 mm. long, the female 60-65 ram.
No other measurements are given, and the worm is not
figured. It seems possible that this m a y be a synonym
of P. obtusus.

8. Proleptus (?) minutus (v. Beneden, 1871).

Syn. Coronilla minuta v. Beneden, 1871.
This form is recorded b y v. Beneden from Raja rubu8
and R. davata, b u t no description of it is given. Possibly
it m a y be a synonym of P. robustus.

9. Proleptus elegans (0rley, 1885}.

Syn. Splropterina elegans Orley, 1885.
The host from which this species is recorded is Hexan-
chus griseus. The length of the worms, according to
0rley's description, is 15-20 mm. in both sexes. The
description appears to contain no other characters of
specific value except the statement that the male has
five pairs of rib-like preanal papillae.
 I
P. acutus. P. obtusus. P . inflatus. P. africanus. P. robus~us. P . australis.

Describer . . . . Dujardin, v. L i n s t o w , 1901 ; v. L i n s t o w , v. L i n s t o w , Baylis Baylis

...... 1845. Seurat,Lloyd,19191920.; 1890. 1899. (present paper). (present paper).

~. ~. ?.
Length .................... 12 31.73-38 32.17-59 35-5 39.7-57 19.7 23-3 33 40-44 14-16 18.5

Thickness (max.) ............ 0.25 0.54-0.55 0"73-0"91 0.54 0.54-0-~ 0.51 0.71 0"65 0"83-0"97 0-46-0.48 0.75

A n t . e n d to e n d of o e s o p h a g u s . -- 3.6-4.79 3"69-6.9 [5.07] [7.4] [3.18] [4] 6-6-7 2.4-2.7 3.4

[4-9]

A n t . e n d to n e r v e - r i n g . . . . . . . 0.4--0.645 0.48 0'42-0'5

0.3~0.38
m
A n t . e n d to cervical p a p i l h e . . . 0-46-0.54 0.65 0"39-0.45

A n t . e n d to e x c r e t o r y p o r e . 0.67-1.17 0'85-0"9 0.54-0-6

Tail, l e n g t h ............... 0.24 1.03-1.560.395--0.71 [0.6] [0.77] [0.318] [0-12] 0-8 0 . 4 2 - 0 44 0.55 0.4
[1.76] [1.27]

~ 0.46, 1-32-2.125, f 0"44, 0.76, 1.5, 1.2,

Spicules, l e n g t h s . . . . . . . . . . . . i 0" 12 0.36-0.41 0"19 0.29 0.4 0.26-0.27

Eggs ...................... -- 0.047-0.052 0.043 0.029 0.049 X 0.031 0.045 0-032 0.038 0.027
0"02875-0-037
i
 334 On the Nematode Genus Proleptus.
10. Proleptus (?) rajce (Diesing, 1851).
Syn. Spiroptera rajaz I)iesing, 1851 ; Splroptera Rajce batis Diesing,
1851 ; Spiropterina rajae P a r o n a , 1894 ; Spiroptera raja~ clavat~
Molin, 1860 ; Spiropterina rajaz clavatcv v. L i n s t o w 1890.
All the names q u o t e d a b o v e seem to h a v e been based
on larval forms recorded b y B c l l i n g h a m and b y Dujardin,
and do not refer to a n y identifiable species.

11. " Proleptus " tortus v. Linstow, 1906.

This species, recorded f r o m a tortoise, Cistudo ornata,
does not a p p e a r f r o m v. L i n s t o w ' s description to be a
Proleptus, b u t m a y be a Physaloptera.
12. " Proleptus " gordioides v. Beneden, 1858.
This w o r m was f o u n d in the wails of the u t e r u s a n d
in the chorion of the eggs of Galeus canis. From
v. Beneden's description a n d figures (1871) it seems almost
certain t h a t it was n o t a Proleptus, and more probable
t h a t it was a species of Philometra.

F r o m the preceding review it appears t h a t six species

m a y be recognized with some degree of p r o b a b i l i t y as
belonging to the genus Proleptus. Some of the more
i m p o r t a n t m e a s u r e m e n t s of these are given in the table on
p. 333. The m e a s u r e m e n t s are in millimetres. The figures
in square brackets h a v e been calculated f r o m p r o p o r t i o n a l
m e a s u r e m e n t s given b y v. Linstow.

I~EFEREIqC ES.
BAXLIS, I t . A. 1928. " R e c o r d s of some Parasitic W o r m s f r o m
British V e r t e b r a t e s . " Ann. & Mag. Nat. Hist. (10) i. pp. 329-
343.
BENEDE~, P. J. VAN. 1858 (1861). " Mdmolre s u r les vers intesti-
n a u x . " Compt. Rend. Acad. Sci., Paris, Suppl. ii. pp. 1-376,
pls. i.-xxvii.
1871. " Les Poissons des c6tes de Belgique, leurs Parasites et
leurs C o m m e n s a u x . " M6m. Acad. R o y Sci. Belg. xxxviii.
4 mdm., x x + i00 pp., pls. i.-vi.
DUJAI~DII~, F. 1845. ' Histoire naturelle des H e l m i n t h e s . ' Paris.
LI~STOW, O. vo. 1890. " B e i t r a g z u r K e n n t n i s s d e r V o g e l t ~ i n i e n , & c . "
Arch. f. N a t u r g . lvi. 1, pp. 171-188, pl. x.
----. 1899. " N e m a t o d e n aus der Berliner Zoologischen S a m m l u n g . "
Mitt. Zool. Mus. Berlin i. pp. 3-28, pls. i.-vi.
1901. " B e o b a c h t l m g e n an H e l m i n t h e n des Senckenbergischen
n a t u r h i s t o r i s c h e n Museums, &c." Arch. f. mikr. Anat, 1viii.
pp. 182-198, pls. viii., ix.
1906. " Neue H e l m i n t h e n . " Centralbl f. Bakt. &e., Abt. I.
Orig., xli. pp. 749-752, 1 pl.
 On new or little-l,mown Tipulid~e. 335
LLOYD, J. ]c[. 1920. " Some Observations on the Structure and
Life-History of the Common Nematode of the Dogfish (Scyllium
canicula)." Prec. Zool. Soc. Lend. pp. 449-456.
Mt~ZLEI~.,3. F. 1925. " Some new Features of Nematode Morphology
in Proleptus obtusus Dujardin." Journ. Parasitot. xii. pp. 84-90,
pl. x.
(~RLEY, L. 1885. " Die Entozoen der Haien und Rochen." Terrors.
Fiiz., Budapest, ix. pp. 97-126 {Hungarian), 216-220 (German
rdsumd), pls. ix., x.
S ~ A T , L.G. 1919. " Sur la morphologie du Proleptus obtusus Duj.
(Acuariidee)." Bull. Mus. Hist. Nat., Paris, pp. 166-172.
TAYLOR, E. L. 1925. "/qotes on some Nematodes in the Museum
of the Liverpool School of Tropical Medicine.--II." Ann.
Trop. Med.& Parasitol. xix. pp. 57-69.

XXXIV.--New or little-known Tipulidm (Diptera).--L.

Australasian Species. B y CI~&RLES P. ALEXANDER,
P h . D . , F.E.S., M a s s a c h u s e t t s S t a t e College, A m h e r s t ,
M a s s a c h u s e t t s , U.S.A.
THE crane-flies discussed h e r e w i t h are all f r o m E a s t e r n
Australia, where t h e y were collected b y m y colleagues,
Messrs. H e r o n , Illingworth, Tonnoir, a n d Wilson, to
w h o m I a m g r e a t l y i n d e b t e d for this f u r t h e r aid in m a k i n g
k n o w n t h e v a s t l y rich f a u n a of the C o m m o n w e a l t h .
T h e m a t e r i a l collected b y T o n n o i r is p r e s e r v e d in the
S t a t i o n Collection a t C a n b e r r a ; the Wilson specimens
are p r e s e r v e d in t h e c a b i n e t of t h e c o l l e c t o r ; t h e
r e m a i n i n g specimens are r e t a i n e d in m y own collection.
L i m o n i a (Dicranomyia) $unetulatella, sp. n.
Closely allied a n d generally similar to L. (D.) punctulata
(de Meijere) of t h e G r e a t e r S u n d a Islands, differing
especially in the s t r u c t u r e of the m a l e h y p o p y g i u m .
M a l e . - - L e n g t h a b o u t 5 m m . ; wing 5.8 ram.
R o s t r u m , palpi, a n d a n t e n n m b l a c k t h r o u g h o u t . Meso-
n o t a l preescutum reddish brown, w i t h t h r e e d a r k e r
b r o w n stripes, of which only the m e d i a n one is b r o a d
a n d distinct. F e m o r a d a r k e n e d s u b t e r m i n a l l y . Wings
w i t h a b o u t five or six d a r k spots in cell C b e t w e e n
arculus a n d s t i g m a ; a large oval d a r k s p o t on vein M
j u s t b e y o n d o n e - t h i r d t h e l e n g t h ; r e m a i n d e r of wing-
p a t t e r n as in the group. Male h y p o p y g i u m with t h e
c a u d a l m a r g i n of tergite b r o a d l y e m a r g i n a t e , each lateral
lobe w i t h a b o u t eighteen to t w e n t y coarse set~e. V e n t r o -
m e s a l lobe of basistyle v e r y s t o u t a n d b r o a d - b a s e d .
Dorsal dististyle g e n t l y curved, a little dilated j u s t