T H E GENUS D O L A B E L L A
by
Dr H. ENGEL
Zoological Museum,
With
Amsterdam
16 text-figures
Diagnosis. Aplysiidae o f conical form, narrower
obliquely
truncate behind.
bearing v i l l i ; warts
Integument
more
or
i n front,
wide and
less w a r t y , the
warts
a n d v i l l i b e i n g w h o l l y retractile. P a r a p o d i a u n i t e d ,
save f o r a d o r s a l s l i t ; the free lobes c o v e r i n g the s l i t leave t w o c o n s p i c u o u s
r e s p i r a t o r y openings, one at the p o s t e r i o r a n d one at the a n t e r i o r e n d . T h e
posterior
respiratory
o p e n i n g , c o r r e s p o n d i n g to the mantle-siphon, lies
about i n the m i d d l e o f the p o s t e r i o r d i s c , the a n t e r i o r o p e n i n g outside the
disc, somewhat i n front o f the disc's a n t e r i o r m a r g i n . S h e l l s o l i d , hatchetshaped, the free s p i r e calloused. P e n i s a r m e d o r u n a r m e d . R a d u l a w i t h
i n c o n s p i c u o u s , o f t e n reduced r h a c h i d i a n tooth a n d m a n y laterals w i t h l o n g
s i m p l e cusps.
S h o r t h i s t o r y o f the genus
T h o u g h m o s t a u t h o r s consider it, w i t h C u v i e r (1804) a d i s c o v e r y o f
P e r o n , that R u m p h i u s ' a n i m a l o f plate X belongs to the s h e l l o f h i s plate
X L , this seems not true. I f w e compare the t e x t o n p. 38 o n
tertia w i t h that o n p. 122-3
o
n
t n e
Limax (marina)
Opercula callorum, i t i s clear that
R u m p h i u s w a s s p e a k i n g o f the same species o f m o l l u s c , that the o n l y
m i s t a k e Schijnvoet, the editor o f R u m p h ' s M . S . , made, w a s that he d i d
not refer to p. 38 w h e n R u m p h i u s o n p. 122 r e m a r k e d : " z i j n d e een slach
v a n den L i m a x m a r i n a " . T h e " t h i r d o p e r c u l u m " lies i n the
flesh and
looks l i k e a h a m ( " s c h o n k j e v a n een v a r k e n " , w h i c h m a y be a scapula,
as some have interpreted it, o r a h a m , as R u m p h i u s h i m s e l f s a i d o n p. 3 8 ) .
A l l details o n the taste a n d s m e l l a n d o n the habitat are the same o n p . 38
a n d p. 123. W e m a y conclude that R u m p h i u s i n t e n d e d the same a n i m a l ,
t h o u g h the d e s c r i p t i o n o f the a n i m a l a n d that o f the s h e l l w e r e
inserted i n a separate chapter
o f the b o o k b y Schijnvoet. T h e
h i s t o r y o f the genus w i l l be clear f r o m the f o l l o w i n g s h o r t
each
further
summary,
g i v e n w h i l e so m a n y o l d types w e r e so o f t e n renamed. F u r t h e r l i t e r a t u r e
w i l l be f o u n d i n the g e o g r a p h i c a l section.
H.
ENGEL
Limax marina tertia R u m p h i u s , 1705, p. 38, p i . X no. 5 ( a n i m a l f r o m
Ambon).
Derde slack van de Opercula callorum (resp. derde v a n de vijfde s o o r t
van klipkleevers)
Rumphius,
1705, p. 122-3, pi- X L f i g . N
(shell
from
Ambon).
Doris verrucosa B a r b u t , 1783, p i . I V f i g . 1 ( t h i s is R u m p h i u s ' p i . X f i g .
Doris); n o n D. verrucosa L i n n e ,
7 S 8 , p. 653, w h i c h is a n Oncidium? a n d a Phyllydia verrucosa.
(Patella) Scapula M a r t y n , 1786, v o l . I l l f i g . 9 9 ( a shell f r o m A m b o n ;
5, the t e x t (p. 36) describes a E u r o p e a n
I
f o r the date see D a l l , P r o c . U . S . N a t . M u s . , v o l . 33, p. 187).
Patella auricularia S o l a n d e r , 1786, p. 154 ( n a m e f o r R u m p h i u s * shell
p i . X L f i g . N , fide Iredale, 1929, p. 2 9 2 ) .
Dolabella c aliosa L a m a r c k , 1801, p. 62 ( n a m e f o r R u m p h i u s p i . 40, f i g .
w i t h the Dolabella s h e l l o n p i . X L n u m b e r e d N , n e v e r 12).
Dolabella . . . C u v i e r , 1804, p. 437, p i . 29 f i g . 1 ( a n a n i m a l , collected b y
12) ( s i c ! I have o n l y seen editions o f the " R a r i t e i t k a m e r "
P e r o n at H e de F r a n c e , d e s c r i b e d a n d
figured).
Dolabella rumphii C u v i e r , 1817, p. 398 (name f o r the f o r e g o i n g a n i m a l ,
w h i c h C u v i e r i d e n t i f i e d w i t h R u m p h ' s a n i m a l a n d shell, t h o u g h he w r o n g l y
gives p i . X f i g . 6, instead o f f i g . 5 ) .
Dolabella peronii B l a i n v i l l e , 1819, p. 395 ( n a m e f o r P e r o n ' s a n i m a l a f t e r
C u v i e r ' s d e s c r i p t i o n , a n d f o r R u m p h ' s shell w h i c h , o n P e r o n ' s a u t h o r i t y ,
B l a i n v i l l e c o n s i d e r s i d e n t i c a l w i t h that o f P e r o n ' s a n i m a l ) .
Dolabella rumphii B l a i n v i l l e , 1819, p. 395, n o n C u v i e r ! ( n a m e
for
R u m p h ' s a n i m a l p i . X f i g . E o r 5, w h i c h B l a i n v i l l e does not r e g a r d i d e n t i c a l w i t h R u m p h ' s shell, n o r w i t h P e r o n ' s a n i m a l as d e s c r i b e d b y C u v i e r ) .
Dolabella rumphii V a n H a s s e l t , 1824, p. 54 ( d e s c r i p t i o n o f a n a n i m a l
from
Peperbaai, Java.
The
drawing
after
the
living animal
was
later
r e p r o d u c e d b y R a n g a n d is s t i l l e x t a n t i n the L e i d e n M u s e u m . A n a n i m a l
collected b y V a n H a s s e l t a n d p r e s e r v e d i n the same M u s e u m c o u l d
be
examined).
Aplysia (Dolabella) hasseltii R a n g , 1828, p. 49, p i . X X I V f i g . 1 ( a f t e r
the d r a w i n g b y V a n
Hasselt).
Aplysia dolabella D u f o , 1840, p. 202 ( a n a m e f o r R u m p h ' s a n i m a l , after
L a m a r c k ' s generic n a m e ) .
Dolabella rumphii v a r . maculosa B e r g h , 1905, p. 18 (new name for
D. hasseltii).
etc.
etc.
Aplysia (Dolabella) gigas R a n g , 1828 ( s h e l l o n l y , the a n i m a l is described
hereafter).
THE
GENUS
199
DOLABELLA
T h e other s p e c i f i c names, as w i l l be s h o w n a l l s y n o n y m s o f
scapula
( M a r t y n ) , are g i v e n i n the g e o g r a p h i c a l section. T h e
Dolabella
following
a u t h o r s have g i v e n a s u r v e y o f the g e n u s : R a n g ( 1 8 2 8 ) , S o w e r b y ( 1 8 6 8 ) ,
Mazzarelli (1893), P i l s b r y (1896), Clessin (1899), B e r g h (1905), M a c F a r land (1918).
Dolabella contains
D. gigas ( R a n g ) .
It i s the a u t h o r ' s o p i n i o n that the genus
species, v i z . ,
D. scapula
( M a r t y n ) and
T h e Species o f the G e n u s
only two
Dolabella
Dolabella scapula ( M a r t y n ) (= D. agassizi, andersoni, auricularia,
callosa, dolabella, ecaudata, hasseltii, hemprichii, neira, peronii, rumphii,
teremidi, tongana, truncata, variegata). T h i s species can be d i s t i n g u i s h e d
f r o m the f o l l o w i n g b y the shell, w h i c h here has a h e a v i l y c a l l o u s e d s p i r e ,
a n d b y the penis w h i c h i s u n a r m e d . T h e glans i s r a t h e r short,
foliate
( b e a r i n g t w o folds o f s k i n , a s m a l l e r a n d a broader, each o n one s i d e o f
the s p e r m a t i c g r o o v e ) .
Dolabella gigas
( R a n g ) . T h i s species was t i l l n o w o n l y k n o w n as a
shell, w h i c h bears o n the s p i r e a v e r y large, t h i n , erect,
saucer-shaped
accessory p l a t e ; o n l y the a p e x o f the shell bears o n its i n n e r s i d e a s m a l l
l u m p o f callus. T h e penis o f this species is a r m e d , h a v i n g s m a l l tubercles
b e a r i n g spines, a r r a n g e d o n l o n g i t u d i n a l folds o f the p r a e p u t i u m , especially
o n the h e a v i e r folds b o r d e r i n g the s p e r m a t i c g r o o v e . T h e glans p e n i s has
the same spine b e a r i n g tubercles t i l l about h a l f w a y i t s l e n g t h a n d e s p e c i a l l y
o n the side opposite the s p e r m a t i c g r o o v e . T h e glans is l o n g , t h i n a n d c o n i c a l
a n d does not s h o w the s i d e flaps o f the p r e c e d i n g species.
Material examined ( A = M u s e u m Amsterdam, L = M u s e u m L e i d e n ) :
Dolabella gigas
1
3
1
1
3
5
(Rang)
specimen, Moluccas, 1858, leg. V a n der Hucht ( A ) .
shells, idem, idem ( A ) .
shell, Moluccas ( A ) .
shell, Indian Ocean ( A ) .
shells, without locality ( i A , 2L).
shells, Mauritius ( A ) .
Dolabella scapula
3
1
1
1
(Martyn)
specimens, Telok Dalam, Nias, leg. D r . J . P. Kleiweg de Zwaan (A.).
specimen, Sabang, leg. G. Herman (A).
specimen, Java (probably Peperbaai 1823), leg. V a n Hasselt ( L ) .
specimen, Island Hoorn, Bay of Batavia, leg. Dr. J . Verwey et W . S. S. van Benthem
Jutting ( A ) .
200
H.
ENGEL
i specimen, Bay of Batavia, leg. D r . C. Ph. Sluiter (A).
I specimen, Island Kerkhof, Bay of Batavia, leg. Jhr. W . C. van Heurn, 31-VII-1927
(A).
1 specimen, Pulu Panggang, Thousand Islands, Java Sea, leg. D r . J . D . F . Hardenberg (A).
1 shell, Tjilaut Eureum, S. Coast of Java (Geol. Inst. Amsterdam).
3 specimens, Sailus Ketjil, Paternoster Islands, close to reef (Stat. 37 Siboga Exp.,
cf. Bergh, 1905, p. 18) (A).
2 specimens, Bay of Bima, near South fort and Bima anchorage (Stat. 47 Siboga
Exp., cf. Bergh, 1905, p. 13) (A).
1 specimen, Buka or Cyrus-bay, south coast of Rotti Island (Stat. 299 Siboga Exp.,
cf. Bergh, 1905, p. 18) (A).
4 specimens, Pepela-bay, east coast of Rotti Island (Stat. 301 Siboga Exp., cf.
Bergh, 1905, p. 13) (A).
2 specimens, Sanana-bay, east coast of Sula Besi (Stat. 193 Siboga Exp., cf. Bergh,
1905, P. 18) (A).
1 specimen, Moluccas, 1858, coll. V a n der Hucht (A).
5 shells, Moluccas, don. G. de Seriere (A).
2 shells, Moluccas (A).
1 specimen, Ambon, leg. Forsten (L.).
1 specimen, Bay of Ambon, leg. Ludeking (L).
1 specimen, Ambon, leg. Bleeker (A).
1 shell, Ambon, leg. Hoedt (L).
1 specimen, Banda, leg. V a n de Velde (A).
1 specimen and 1 shell, New Guinea, leg. D r . H . Macklot (L).
1 shell, Waigeu, ex museo F . van Heukelom (A).
1 specimen, Biak Island, near New Guinea, 1915, leg. W . K. H . Feuilleteau de
Bruyn (A).
2 specimens, Kaimana, Southwest New Guinea, 133 44" E . 3 41' S., reef, leg.
L . Hendriks, 28-VIII-1910 (A).
4 specimens, Pulu Sangursiapo, Tawi-Tawi Islands, Sulu Archipelago (Stat. 93 Siboga
Exp., cf. Bergh, 1905, p. 19) (A).
1 shell, Philippine Islands (L).
1 specimen, probably Japan (Mus. Berlin).
2 specimens, Tiop, Bougainville Island, Solomon Islands, 3-4-XI-1909, leg. H . Schoede
(Mus. Berlin).
1 shell, New Caledonia (A). N . B . This specimen was labelled: "Dolabella sulcata
Bernardi, Nw. Caledonia, ab auctore". A s far as I know this species was never
published by Bernardi, the name is only a museum name. The shell is that of the
common D. scapula.
2 shells, Indian Ocean (L).
4 shells, Mauritius (2A, 2L).
2 shells, Mauritius, coll. Schepman (A)3 specimens, Island Inhaca near Lorengo Marquez, July 1934, leg. Prof. D r . C. J .
van der Horst (A).
1 specimen,, idem, no date (A).
6 specimens without locality (5L, i A ) .
0
Dolabella gigas as
H i s t o r y o f the
0
a separate species
species.
Aplysia (Dolabella) gigas R a n g , 1828, p. 48, p l . I l l f i g . 4, after
shell o n l y , f r o m the " M e r des Indes".
the
THE
GENUS
Dolabella gigas Deshayes, 1863, p.
Dolabella gigas S o w e r b y i n R e e v e ,
201
DOLABELLA
53, m e n t i o n e d f r o m H e de l a R e u n i o n .
1868, X V I , p i . I f i g . 1 ( n e w f i g u r e s
after specimens f r o m " I n d i a n S e a s " ; i n C u m i n g ' s c o l l e c t i o n there w e r e t w o
y o u n g shells, p r o b a b l y this species, f r o m the R e d S e a ) .
Dolabella gigas L i e n a r d , 1877, p . 53 ( M a u r i t i u s ) .
Dolabella gigas M a r t e n s , 1880, p . 306 ( V o n P r o f .
M o b i u s lebend a u f
M a u r i t i u s gefunden. B i s 3 0 c m l a n g , v i o l e t t u n d g r a u , m i t k o n i s c h e n a n
der
S p i t z e abgerundeten
e i n f a c h e n W a r z e n , w a h r e n d d i e folgende A r t
o l i v e n g r i i n i s t , m i t s p i t z e n W a r z e n , w o r a u f spitze P a p i l l e n stehen. B e i d e
stossen v i o l e t t e n S c h l e i m aus d e m M a n t e l s p a l t a u s ) .
Dolabella gigas
P i l s b r y , 1896, p. 152, p i . 65 figs. 4-6. T h e s e f i g u r e s a r e
new, b u t P i l s b r y does not m e n t i o n t h e i r locality.
G e o g r a p h i c a l d i s t r i b u t i o n (see m a p , f i g . 1 6 ) : M a u r i t i u s ( b ) , R e d S e a
(a), Moluccas (c) (described below).
D e s c r i p t i o n o f the a n i m a l o f Dolabella gigas.
A s m e n t i o n e d above w e c o u l d investigate one s p e c i m e n o f t h i s species
f r o m the M o l u c c a s (1858) i n o u r M u s e u m f r o m the V a n d e r H u c h t c o l lections. I t w a s c o m p a r e d w i t h M a c F a r l a n d s b e a u t i f u l d e s c r i p t i o n o f h i s
D. agassizi.
E x t e r n a l characters.
G e n e r a l f o r m a n d c o l o u r . A s m y f i g u r e 1 s h o w s , the b o d y i n i t s p r e s e r v e d
c o n d i t i o n has the same o u t l o o k as D. scapula. T h e b o d y c a r r i e s a l o t o f
tubercles, w h i c h i n t h e i r centre c l e a r l y s h o w the t i p o f the retracted p a r t .
A c c o r d i n g to M a r t e n s (1880, p. 3 0 6 ) these appendages a r e s i m p l e ( n o t
b r a n c h i n g as i n
D. scapula).
T h e y a r e m o r e c r o w d e d o n the disc t h a n o n the rest o f the b o d y . L i k e
in
D. scapula
between
the
some p a r t s o f the s k i n a r e q u i t e s m o o t h a n d even. T h e s k i n
nodosities
looks
l i k e shagreen
leather, a c u r i o u s
feature
w h i c h I d i d not see i n a n y s p e c i m e n o f D. scapula. I n the s m o o t h e r p a r t s
o f the s k i n , h o w e v e r , these v e r y s m a l l shagreen tubercles m a y disappear
a l s o ; they m a y be present o r not o n the l a r g e r nodosities. T h e m a r g i n o f
the d i s c is v i l l o s e , its v i l l i a r e not a l l a n d not completely retracted. T h e
surface o f the foot i s s m o o t h ; as u s u a l i n the genus, the foot shows n o
c l e a r l y d e f i n e d m a r g i n s , o n l y i n front there i s a g r o o v e between the foot
a n d the head, w h i l e its r i g h t a n t e r i o r c o r n e r has a tentacle-like f o r m , as'
the f i g u r e shows. T h e a n t e r i o r tentacles are s t r o n g l y contracted, ear-like.
T h e l i k e w i s e contracted r h i n o p h o r e s stand close together a n d o n l y s h o w
a s m a l l g r o o v e o n t h e i r top, not c o n t i n u i n g a l o n g the w h o l e length. T h e
202
H. ENGEL
body colour is that of all Dolabella's after preservation in alcohol, a dirty
yellow, but the remains of dark pigment blotches are clearly visible here
and there, as the figure shows.
The length of the preserved body is 108 mm, its greatest transverse
diameter 64 mm, its greatest height about 40 mm. T h e posterior disc is
54 mm broad and 51mm long. T h e total length of the epipodial slit is
Fig. 1. Dolabella gigas (Rang) from the
Moluccas, from above. Natural size.
F i g . 2. Dolabella gigas (Rang),
diagrammatically to show the pallial
complex. F o r explanation see text, o,
osphradium; g, genital pore.
60 mm (i.e., 0.5 of the body length), beginning about 41 mm (i.e., 0.38 of
the body length) from the anterior end of the body and 26 mm from the
rhinophores. M y figure 1 shows the epipodial slit continuing far into the
disc. In this figure the posterior part of the disc is shortened in perspective and, moreover, it is strongly contracted. Smoothing out as far
as possible the corrugations one can measure a distance of 25 mm between
THE
GENUS
203
DOLABELLA
its p o s t e r i o r b o r d e r a n d the p o s t e r i o r end o f the body, so that w e m a y
say, that the p o s t e r i o r o p e n i n g lies i n the c e n t r e o f the d i s c . T h e e p i p o d i a l
slit
is w i d e l y open. I do not see a n y t h i n g o f i n t e r l o c k i n g grooves a n d
ridges,
nor
o f the
l o w sharp-edged
fold
round
the p o s t e r i o r o p e n i n g
described by M a c F a r l a n d .
T h e s k i n is not so v e r y t h i c k , i n the m i d d l e o f the foot V 2
m m
>
thicken-
i n front t i l l 2 m m a n d b e h i n d t i l l 5 m m .
ing
T h e p a l l i a l c o m p l e x . T h i s has been d r a w n i n f i g u r e 2. U n h a p p i l y it
was, o f course, o n l y after
I h a d r e m o v e d a n d studied the s h e l l , that I
r e c o g n i z e d t h i s a n i m a l to be D. gigas. S o the t r i a n g u l a r
shell-foramen
s h o w n i n f i g u r e 1 has been reconstructed after the traces i t h a d left o n
the surface
o f the s h e l l . L i k e w i s e , the mantle-flap, w h i c h encloses
s h e l l , is s h o w n i n f i g u r e 2 s h r u n k after
the
the r e m o v a l o f the s h e l l . O r i g -
i n a l l y it w a s l a r g e r . H e r e i t is i n d i c a t e d b y a h e a v y l i n e , e n d i n g b e h i n d
i n the mantle s i p h o n w h i c h contains the a n a l o p e n i n g i n its w a l l . A l i n e
of
dots a n d stripes r u n n i n g o v e r the m i d d l e o f the mantle indicates
the
place w h e r e i t is fastened to the b o t t o m o f the c a v i t y . T h e r i g h t h a l f is
free
and
cavers
the
ctenidium
(especially w h e n
the
shell is
present,
w h i c h i n o u r f i g u r e is not the case), the left h a l f ' s underside is one w i t h
the b o t t o m o f the m a n t l e c a v i t y . T h e t r i a n g u l a r s h e l l - f o r a m e n i n the r o o f
of
the mantle, d r a w n i n f i g u r e 1, has not been r e f i g u r e d i n f i g u r e 2.
T h e left
half
o f the
mantle
contains
i n its widened hinder
part
" s a u c e r - l i k e a p p e n d a g e " o f the s h e l l , w h i c h i s t y p i c a l f o r this
the
species.
T h e c t e n i d i u m , as m a y be seen i n f i g u r e 2, has the c o m m o n f o r m
(cf.
M a c F a r l a n d ) . Its o s p h r a d i u m ( 0 ) is seen, as a l i t t l e slit, near i t s a n t e r i o r
end. T h e c i r c u m f e r e n c e o f the mantle c a v i t y is here i n d i c a t e d b y a b r o k e n
line. T h e left p a r t o f its f l o o r ( p a r t l y c o v e r e d b y the m a n t l e )
contains
the k i d n e y , i n d i c a t e d i n f i g u r e 2 b y a fine c o n t i n u o u s l i n e . I c o u l d not
f i n d the r e n a l pore. T h e f l o o r o f the mantle c a v i t y r i g h t o f the c t e n i d i u m
contains the g e n i t a l pore (g), f r o m w h i c h the g e n i t a l g r o o v e r u n s f o r w a r d
to the r i g h t side o f the head, w h e r e the penis lies i n v a g i n a t e d i n t o
the
b o d y c a v i t y . A s is w e l l k n o w n , t h i s e x t e r n a l g e n i t a l g r o o v e consists o f
t w o s m a l l folds, w h i c h f o r m a gutter, t h r o u g h w h i c h , as H i r a s e (1929, p.
no)
observed, the s p e r m a as w e l l as later o n the s p a w n take t h e i r w a y .
C u r i o u s l y enough
a somewhat
narrower
g r o o v e r u n s f r o m the
genital
pore b a c k w a r d . I n f i g u r e 2 i t has been i n d i c a t e d b y t w o dotted lines.
As
can be
seen
this
groove
branches o n the
fold
of
the
saucer-like
appendage, one b r a n c h f o l l o w i n g its outer contour, one e n d i n g into
siphon.
As
M a c F a r l a n d showed,
the
knob
formed
by
the
the
saucer-like
204
appendage ( o r i n
H.
D. scapula b y
ENGEL
the c a l l u s ) serves the purpose o f s h u t t i n g
the s i p h o n w h e n necessary. T h e use o f the d e s c r i b e d accessory grooves i s
not clear, they m a y s e r v e the e x c r e t i o n o f some g e n i t a l w a s t e
product.
T h e f l o o r o f the m a n t l e c a v i t y r i g h t o f the g r o o v e i s g l a n d u l a r , l i k e w i s e
the free r i g h t p a r t o f the mantle.
Fig. 3 . Shells of Dolabella gigas (Rang), a, b, from the animal here, described,
Moluccas, 1858, V a n der Hucht-collection; a, from above, b, from below; c-f, alsofrom the Moluccas; c-e, belonging to the V a n der Hucht-collection; g, without
locality; h-l, from Mauritius. X ^ .
T h e shell (see f i g u r e 3 a a n d b) is conspicuous i n t h i s species, because
o f the m e n t i o n e d s a u c e r - l i k e appendage, instead o f a callous k n o b . A s m a l l
callus is present o n the i n n e r side o f the apex. T h e calcareous l a y e r i s o f
the same size as the c u t i c u l a r p o r t i o n , i t is m o r e s o l i d t h a n i n
and beautifully porcelain-white within. P i l s b r y
D. scapula
(1896, p. 152) g i v e s a
205
T H E GENUS D O L A B E L L A
v e r y g o o d d e s c r i p t i o n o f i t . O u r s h e l l ( f i g . 3 a a n d 6) has a l e n g t h o f
46 m m a n d a b r e a d t h o f 32 m m .
T h o s h o w the great v a r i a t i o n o f t h i s o r g a n , the f i g u r e s 3 c to / a r e
g i v e n , a f t e r loose shells i n o u r M u s e u m .
T h e penis. A s i n a l l A p l y s i i d a e the penis is i n v a g i n a t e d at the r i g h t
side o f
the head u n d e r
the
r i g h t a n t e r i o r f i r s t tentacle.
The
genital
groove r u n s f r o m the g e n i t a l pore ( f i g . 2 g), w h i c h , as s a i d , here l i e s
u n d e r the a n t e r i o r p a r t o f the c t e n i d i u m , a l o n g the m i d l i n e o f the back
a n d t h e n to the r i g h t , e n t e r i n g the p e n i a l o p e n i n g . T h e g r o o v e continues
a l o n g the w a l l o f the p r a e p u t i u m ( f i g . 4). T h e p r a e p u t i u m consists o f a
n a r r o w , s m o o t h , s i m p l e p a r t a n d a w i d e n e d p a r t , w h i c h last contains the
Fig. 4. Dolabella gigas (Rang). The penis and its praeputium.
a, the opening in the body wall ; a-b, the narrow, smooth part
of the praeputium; b-c, the wider, folded and armed part of
the praeputium; c-d, the retractor muscle; c-e, the glans
penis. Natural size.
glans a n d ends i n the r e t r a c t o r muscle. T h i s w i d e r p a r t s h o w s m a n y
longitudinal
folds a n d i s covered
w i t h tubercles
bearing small conical
spines o n t h e i r top. T h i s a r m a t u r e is especially s t r o n g o n the t w o
folds
b o r d e r i n g the spermatic groove. It continues a l o n g the free glans to about
h a l f w a y i t s l e n g t h , especially o n the side opposite the groove. A s
the
f i g u r e shows, the glans is long, t h i n a n d c o n i c a l a n d does not s h o w the
side-flaps, w h i c h a r e d e s c r i b e d f o r the other species. T h e t w o p a r t s o f
the penis sheath are each about
30 m m l o n g , the glans has a
length
o f 50 m m .
I n t e r n a l characters.
C o m p a r i n g t h i s a n i m a l w i t h the d e s c r i p t i o n g i v e n b y M a c F a r l a n d f o r
h i s D. agassizi, I c a n o n l y a d d :
T h e m a n d i b l e s are 8 já m m l o n g a n d 4j4 m m b r o a d , t h e i r m i c r o s c o p i c a l
s t r u c t u r e as d e s c r i b e d b y M a c F a r l a n d .
T h e r a d u l a i s m o r e t h a n 12 m m l o n g a n d 14 m m b r o a d . I counted m o r e
t h a n 55 r o w s a n d t i l l 203 teeth o n one side i n the 28th r o w . T h e f o r m o f
206
H.
ENGEL
the teeth i s best u n d e r s t o o d f r o m f i g . 5. It is clear that the m i d d l e tooth
l o o k s l i k e that f i g u r e d b y B e r g h f o r D. rumphii, but i t s h o w s no denticles.
It
differs
somewhat
i n the
different
rows. T h i s specimen
is a
good
example o f the v a r i a t i o n i n the r a d u l a . I f w e compare the f i r s t left a n d
the f i r s t r i g h t tooth, w e see that the f i r s t r i g h t resembles m u c h m o r e the
m i d d l e tooth, w h i l e the left has a h e a v i e r denticle. It is l i k e w i s e clear
f r o m o u r f i g u r e that each tooth, as m i g h t be expected, agrees w i t h its
3
25,
2
1
3
^
Fig. 5. Dolabella gigas (Rang). Radula. T h e median part of the 25th to the 28th row
of teeth. Some of the side teeth of the 27th row, indicated by their number, are
given to show their form in different aspects. A s the middle tooth of D. scapula
seems to form a point of discussion among the authors, I have given here the median
tooth of one of the last (54th) rows, as this is the least worn, somewhat more
enlarged than the others, to show the irregular form of the blade.
predecessors a n d its f o l l o w e r s i n a l o n g i t u d i n a l d i r e c t i o n . F o r c o m p a r i s o n
I have g i v e n a l a r g e r f i g u r e o f one o f the last, least w o r n , m i d d l e teeth,
to s h o w the exact f o r m o f the r a t h e r i r r e g u l a r blade.
T h e layer o f palatal spines seems to detach q u i t e easily f r o m the p a l a t a l
folds, f o r I c o u l d o n l y f i n d some traces o f it, after m u c h s e a r c h i n g . T h e s e
s h o w e d the same m i c r o s c o p i c a l s t r u c t u r e as that d e s c r i b e d b y M a c F a r l a n d .
T h e rest o f the intestine agrees w i t h M a c F a r l a n d ' s d e s c r i p t i o n , t h o u g h
some features s h o w e d v a r i a t i o n s w h i c h most p r o b a b l y w e r e i n d i v i d u a l , e.g.,
the f o r m a n d the d i r e c t i o n o f the folds o f the oesophagus a n d the f o r m o f
the tooth bases i n the stomach.
THE
The
species
GENUS
207
DOLABELLA
Dolabella scapula
(Martyn)
H i s t o r y o f the species.
For
the e a r l i e r h i s t o r y see the f i r s t page o f this a r t i c l e , w h e r e a s h o r t
h i s t o r y o f the genus i s g i v e n . A s i t is the a u t h o r ' s o p i n i o n , that, except f o r
D. gigas, a l l the other a n i m a l s f o r m b u t one species, w h i c h m u s t be called
D. scapula, the names there m e n t i o n e d , except D. gigas, c a n be r e g a r d e d
as s y n o n y m s o f D. scapula. T h e f u r t h e r s y n o n y m s a r e f o u n d i n the geo-
D. agassizi, andersoni, auricularia,
callosa, dolabella, ecaudata, hasseltii, hemprichii, neira, peronii, rumphii,
teremidi, tongana, truncata, variegata.
g r a p h i c a l s e c t i o n here after, they a r e :
D e s c r i p t i o n o f the a n i m a l o f
D. agassizi
External
by M a c F a r l a n d and of
D. scapula as
D. gigas here
compared w i t h
that o f
above.
characters.
G e n e r a l f o r m a n d c o l o u r . I a m f i r s t d e s c r i b i n g here the three a n i m a l s
c a p t u r e d b y P r o f . K l e i w e g de Z w a a n at T e l o k D a l a m , i s l a n d N i a s . T h e y
l o o k l i k e the figures g i v e n b y R a n g f o r
1) a n d those o f
1832,
D. hasseltii
D. hasseltii
(1828, p i . X X I V f i g .
i n the A s t r o l a b e A t l a s ( Q u o y & G a i m a r d ,
M o l l . , p i . 23 f i g . 1) b o t h i n the f o r m a n d i n the m a n y fleshy p r o -
cesses.
T h e a n i m a l s are b e a u t i f u l l y preserved.
A l l three s h o w
brown
blotches i r r e g u l a r l y scattered o v e r head, back a n d disc, as m y f i g . 6 s h o w s .
The
tentacles are a u r i c u l a t e , the r h i n o p h o r e s have a slit a l o n g t h e i r upper
outer h a l f . T h e r e t r a c t i l e c i r r h i o r v i l l i a r e f o u n d e v e r y w h e r e , except o n
the foot, even o n the tentacles a n d r h i n o p h o r e s . O n e c a n d i s t i n g u i s h larger,
o f t e n m u l t i p l e o r b r a n c h e d processes, m o s t l y o n tubercles, a n d s m a l l e r ,
s i m p l e ones, scattered between these a n d also o n tubercles. I n a l l three
specimens I see a p a i r o f large a n d c o n s p i c u o u s b r a n c h e d c i r r h i to the
r i g h t a n d the left o f the a n t e r i o r o p e n i n g o f the m a n t l e slit, b u t t h e i r
s p e c i f i c i t y seems v e r y d o u b t f u l . F o r the rest I do not see a n y r e g u l a r i t y .
T h e r e are tubercles o n the head, between the r h i n o p h o r e s , w h i c h a r e r a t h e r
w i d e apart, a n d e v e r y w h e r e o n the back. O n the d i s c they seem to be m o r e
crowded.
T h e d i s c s are a l l three s t r o n g l y c o r r u g a t e d
C o m p a r i n g these specimens w i t h that o f
by contraction.
D. gigas d e s c r i b e d
above, w h i c h
has a m u c h t h i n n e r s k i n (here i t ranges f r o m 4 m m , i n the m i d d l e o f the
foot, to 8 m m , i n front, a n d t i l l 12 m m a n d m o r e , b e h i n d ) , i t seems to me,
that i n
D. gigas
there a r e less s m a l l c i r r h i , w h i l e the tubercles a r e m o r e
numerous a n d m o r e conspicuous., L i k e i n D.
gigas
the fringe o f the disc here
consists o f m a n y i r r e g u l a r lobes. I n f i g . 6 I have d r a w n the largest a n i m a l
after the m a n t l e c a v i t y h a d been l a i d open. T h e w h i t e r i n g r o u n d this c a v i t y
208
H.
ENGEL
Fig. 6. Dolabella scapula (Martyn). The 153I mm long animal collected by Prof.
Kleiweg de Zwaan at Nias. The disc has been cut away, so that the mantle cavity
lies open. F o r further explanation see text, a, anus; g, genital opening.
THE
GENUS
2Q9
DOLABELLA
shows the v e r y t h i c k s k i n i n transverse section w i t h its caverns. T h e c o n t o u r
of the disc, as i t o r i g i n a l l y w a s , has been d r a w n i n a fine dotted l i n e .
T h e l e n g t h o f the three a n i m a l s is 153, 131 a n d 120 m m respectively, their
greatest b r e a d t h 100, 76 a n d 77 m m . T h e d i s c is l o n g 67, 6 0 a n d 6 0 m m
a n d b r o a d 87, 72 a n d 64 m m . T h e total l e n g t h o f the e p i p o d i a l slit i s 6 0 m m
(i.e., 0.39 b o d y l e n g t h ) , 4 7 m m (i.e., 0.36 b o d y l e n g t h ) a n d 53 m m (i.e.,
0.44 b o d y l e n g t h ) . T h e distance f r o m the a n t e r i o r end o f the slit to ths
r h i n o p h o r e s i s 38, 34 a n d 31 m m , a n d to the a n t e r i o r t i p o f the b o d y
55 m m (i.e., 0.42 b o d y l e n g t h ) , 57 m m (i.e., 0.43 b o d y l e n g t h ) , 4 6 m m
(i.e., 0.38 b o d y l e n g t h ) , l h e p o s t e r i o r o p e n i n g o f the slit lies i n the f i r s t
species 23 m m f r o m the a n t e r i o r b o r d e r o f the d i s c a n d 4 3 m m f r o m the
p o s t e r i o r border. I n the second specimen these measurements are 2 0 and
40 m m a n d i n the t h i r d about the same. F o r c l o s i n g the slit, the m i d d l e
p a r t of the left b o r d e r lies o b l i q u e l y w i t h i t s s m o o t h u n d e r s u r f a c e o v e r
the s m o o t h a n d o b l i q u e u p p e r surface o f the r i g h t b o r d e r , e x a c t l y f i t t i n g
together o v e r a s u r f a c e o f about 50 m m breadth. T h e a n t e r i o r o p e n i n g i s
o v a l w i t h n a r r o w , s o m e w h a t u p s t a n d i n g borders. T h e t h i n n e r i n n e r b o r d e r s
o f the s l i t pass r o u n d the o v a l p o s t e r i o r o p e n i n g , w h i c h they c a n close, as
I see i n m y 131 m m s p e c i m e n . T h i s p o s t e r i o r o p e n i n g lies e x a c t l y above
the s i p h o n o f the mantle.
T h e p a l l i a l c o m p l e x . I n the same f i g u r e 6, I have d r a w n t h i s , s o m e w h a t
d i a g r a m m a t i c a l l y . T h e h e a v y b r o k e n l i n e gives the l i m i t s o f the
mantle
c a v i t y . A s the f i g u r e s h o w s , the s k i n s u r r o u n d i n g the c a v i t y i s v e r y t h i c k ,
f r o m 7-23 m m , h a v i n g a spongy texture,
the p r i n c i p a l holes o f
which
w e r e d r a w n . T h e h e a v y continuous l i n e gives the c o n t o u r o f the mantle,
w h i c h i s fastened w i t h its left h a l f to the b o t t o m o f the c a v i t y , f r o m the
t h i c k c o n t i n u o u s l i n e to the l i n e o f dots a n d stripes. T h e p a r t r i g h t o f this
l i n e is free a n d covers the g i l l . T h i s mantle contains the shell. T h e s h e l l
f o r a m e n has been d r a w n i n a t h i n continuous l i n e ; t h r o u g h this o p e n i n g
the shell i s v i s i b l e . T h e callous s p i r e o f the shell is a n c h o r e d i n the b o t t o m
of
the mantle c a v i t y ; above i t I have i n d i c a t e d b y a dotted area
the
" t h i c k e n i n g i n the f l o o r o f the p a l l i a l chamber", a d i s c o r b u t t o n , m e n t i o n e d b y M a c F a r l a n d , w i t h w h i c h the a n i m a l m i g h t close the p o s t e r i o r
mantle o p e n i n g . A b o v e i t the free mantle b o r d e r s r i s e to f o r m the s i p h o n ,
w h i c h i n its p o s t e r i o r p o r t i o n contains the a n a l o p e n i n g , here i n d i c a t e d b y
a s m a l l cross
( a ) . T h e k i d n e y lies i n the left- p a r t o f the f l o o r o f the
mantle c a v i t y , i t has been i n d i c a t e d i n the d r a w i n g b y a t h i n c o n t i n u o u s
l i n e ; its r i g h t angle reaches u n d e r the left p a r t o f the mantle. I d i d not
see the r e n a l pore. A b o v e the k i d n e y , i n the left a n t e r i o r corner o f
Zoologische Mededeelingen X X I V
the
210
H.
ENGEL
mantle c a v i t y , lies the p e r i c a r d . T h e p a r t o f the mantle c a v i t y u n d e r the
mantle flap m a y be c a l l e d the g i l l c a v i t y ; i t i s l i m i t e d o n the left b y the
heavy line o f dots a n d stripes a n d o n the r i g h t b y the h e a v y b r o k e n l i n e .
L i k e i n D. gigas, there is a system o f grooves o n the f l o o r o f the mantle
c a v i t y . T h e s e are l i m i t e d b y s m a l l folds o f s k i n , here i n d i c a t e d b y double
l i n e s . O n e o f these grooves l i m i t s the " b u t t o n " . A second r u n s f r o m the
genital o p e n i n g (g), w h i c h lies under the a n t e r i o r p a r t o f the g i l l to the
p o s t e r i o r b o r d e r o f the mantle. F r o m the g e n i t a l pore, the g e n i t a l g r o o v e
r u n s t h r o u g h the a n t e r i o r o p e n i n g o f the mantle s l i t , a l o n g the back, to the
r i g h t side o f the head, w h e r e i t enters the p e n i s - o p e n i n g . T h e free r i g h t
p a r t o f the mantle, u n d e r the shell, i s v e r y g l a n d u l a r a n d o f a d a r k b r o w n
c o l o u r . T h e f l o o r o f the m a n t l e c a v i t y , r i g h t o f the grooves to the g e n i t a l
pore, is also v e r y g l a n d u l a r
(Glandula Bohadschi). Near
the
anterior
entrance o f the mantle c a v i t y lies the o s p h r a d i u m , v i s i b l e as a s m a l l cleft.
T h e o t h e r specimens here e x a m i n e d w i l l be c o m p a r e d w i t h each other
a n d w i t h the a n i m a l s f r o m the l i t e r a t u r e i n the g e o g r a p h i c a l section (see
b e l o w ) , but a p r e l i m i n a r y c o n s i d e r a t i o n o f the v a r i a b i l i t y o f the e x t e r n a l
features m a y take its place here. A c c o r d i n g to me, the e x t e r n a l
characters
s h o w s u c h w i d e v a r i a t i o n , that i t i s i m p o s s i b l e to d i s t i n g u i s h species o r
even v a r i e t i e s a m o n g these a n i m a l s . A c c o r d i n g to l i t e r a t u r e the c o l o u r is
v e r y v a r i a b l e a n d m a y change i n the .same a n i m a l f r o m b l a c k o r b r o w n to
green ( R i s b e c , 1928, p. 4 5 . See f u r t h e r u n d e r the h e a d i n g : F i g u r e s and
descriptions o f l i v i n g a n i m a l s ) . T h e e x t r a o r d i n a r y t h i c k a n d musculous
s k i n a l l o w s o f a n endless v a r i a t i o n i n f o r m a n d m e a s u r e m e n t s i n the
same a n i m a l .
F i g u r e 7 represents a section t h r o u g h the w h o l e s k i n , not
far
from
the d i s c , the section m e a s u r i n g 9 m m f r o m the u p p e r to the u n d e r surface.
T h e upper surface, w h i c h is covered b y a n e p i d e r m i s o f c y l i n d r i c a l cells,
shows large r o u n d glands, i m b e d d e d i n the u n d e r l y i n g tissue, r e s e m b l i n g
the glands described a n d f i g u r e d b y F r o u t i n (1937, p. 78 seq. a n d p i . I l l
f i g . 31) a n d s h o w i n g the same p a r a l l e l fissures. T h e connective tissue is
r a t h e r compact near the surface, b e c o m i n g m o r e loosely t e x t u r e d i n the
deeper layers. T h e r e i s no r e g u l a r i t y whatever^ but the u s u a l elements are
present (cf. H o f f m a n n , 1939, p. 527 seq. a n d F r o u t i n , 1937). I m b e d d e d i n
the e n o r m o u s mass o f collagene f i b r e s , the muscle fibres, o f t e n connected
i n bundles, are c o n s p i c u o u s . A s the figures s h o w , they m a y pass across the
w h o l e section, but this is not necessarily the case.
T h e u n d e r s u r f a c e has no c y l i n d r i c a l e p i t h e l i u m , the cells seem to be
r a t h e r o f the flat type. N e a r i t heavy transverse a n d l o n g i t u d i n a l muscle
THE
bundles are present.
figured
GENUS
211
DOLABELLA
T h e lacunae s h o w n i n f i g . 7 c o r r e s p o n d to
i n f i g . 6 ( o n the
cross
section o f
the
base o f
the
those
removed
parapodia).
I t is clear, that s u c h a t h i c k l a y e r (4-12 m m a n d m o r e ) o f criss-cross
f i b r e s m u s t be v e r y elastic. T h e m a n y
muscles w i l l a l l o w f o r c o n t r a c t i o n i n
all
directions.
M o r e o v e r the
water
c o n t a i n e d i n the s p o n g y t e x t u r e m u s t
c o n t r i b u t e to the e l a s t i c i t y a n d
ex-
t e n s i b i l i t y o f the w h o l e ( c f . J o r d a n ,
1901).
I t w i l l be necessary to k n o w w h e t h e r
the presence o r absence o f the
villi
o n the s k i n i s a s p e c i f i c o r s u b s p e c i f i c
character o r so v a r i a b l e that i t is o f
no
importance.
There
described "species" o f
have
been
Dolabella
with
a s m o o t h s k i n o r w i t h some s m a l l a n d
l o w t u b e r c u l a t i o n s as
character.
their
specific
many
ani-
mals at m y d i s p o s a l , I d i d not
find
a n y w i t h o u t v i l l i , tubercles o r
their
remnants,
S t u d y i n g the
which
last
both
gave the i m p r e s s i o n to be
retracted
v i l l i . O f three a n i m a l s f r o m
near
Lorenzo
clear
and
Marques,
distinct
villi,
always
Inhaca,
two
while
have
the
t h i r d has a s m o o t h s k i n , o n w h i c h
o n l y i n some places rests o f the v i l l i
can be f o u n d , e s p e c i a l l y o n protected
o r less c o n t r a c t e d p a r t s o f the s k i n .
Fig. 7 . Dolabella scapula (Martyn) from Kerkhof Island, Bay of Batavia. L e f t : section
through the whole skin, not far from the disc ( 9 mm high); right: muscles in this part
of skin (diagrammatical), at the right a villus-top, near the under surface lacunae
and heavier transverse and longitudinal muscles.
212
H.
ENGEL
Fig. 8 . Dolabella scapula (Martyn). Same piece of skin
as fig. 7, section through a partly retracted villus.
F i g . io. Dolabella scapula (Martyn).
Same piece of skin as fig. 7 , section
through the skin in a place where
obviously a villus was totally retracted.
F i g . 9 . Dolabella scapula (Martyn). Same piece of
skin as fig. 7, section through a partly retracted villus.
THE
GENUS
213
DOLABELLA
It m u s t be supposed, that some a u t h o r s r e p o r t i n g a w h o l l y s m o o t h s k i n ,
h a v e o v e r l o o k e d tfhe i n d i s t i n c t a n d s m a l l r e s t s o f the
tuberculations.
I have the i m p r e s s i o n that, i n the l i v i n g a n i m a l , v i l l i are a l w a y s present,
be i t i n greater q u a n t i t y i n one t h a n i n another a n i m a l (see b e l o w u n d e r
the h e a d i n g : T h e l i v i n g a n i m a l ; cf. R a n g a n d A n o n y m u s e s p e c i a l l y ) . T h e s e
v i l l i are v e r y retractile, as has been r e p o r t e d a l r e a d y b y the e a r l i e r a u t h o r s
( c f . « V a n H a s s e l t , E h r e n b e r g ) . O n p r e s e r v a t i o n they w i l l be retracted m o r e
or
less s t r o n g l y , but, as s a i d , even w h e n v e r y s t r o n g c o n t r a c t i o n takes
place, remains are a l w a y s v i s i b l e . T h e q u e s t i o n arises, h o w e v e r ,
whether
these v i l l i can be retracted so completely, that they disappear t o t a l l y i n the
s k i n , w h i c h then l o o k s q u i t e s m o o t h a n d even, i n the places w h e r e they
disappeared. L a r g e r stretches o f s m o o t h s k i n i n some a n i m a l s suggest this
question. S e c t i o n s w e r e made to f i n d the answer. I n f i g . 9 a v i l l u s i s s h o w n
i n a p a r t l y retracted state. W e see that it contains the c o m m o n elements,
some muscles e x t e n d t h r o u g h its w h o l e l e n g t h . T h e base o f the v i l l u s ,
h o w e v e r , l o o k s so n a r r o w , that one doubts w h e t h e r this " n e c k " r e a l l y can
e x t e n d i n s u c h a measure as to m a k e r o o m f o r the w h o l e s k i n o f the v i l l u s ,
w h i c h a c c o r d i n g l y m u s t s m o o t h e n out, l e a v i n g n o w r i n k l e s o r folds i n the
e p i t h e l i u m . J o r d a n (1901) figures ( f i g . 3 ) a bubble o f the s k i n w h i c h is
suggestive o f this p o s s i b i l i t y . O u r f i g . 8 gives another v i l l u s , m o r e retracted, w h i l e i n f i g . 7. J a less retracted v i l l u s has been cross sectioned. T h e
a u t h o r a w a i t e d to see i n the m a n y sections t h r o u g h the s k i n the d i f f e r e n t
s t a d i a i n the r e t r a c t i o n o f the v i l l i , but he w a s d i s a p p o i n t e d i n f i n d i n g
but one place i n the s k i n , w h e r e one c o u l d suppose, that a v i l l u s
had
disappeared because o f the compressed e p i d e r m i s a n d the m a n y muscles
concentrated i n that one place ( f i g . 10). I n f i g . 7 a the d i r e c t i o n o f the
collagene fibres a l o n g the muscles i n the upper p a r t suggests t h e i r connect i o n w i t h the muscle i n its i n w a r d a n d o u t w a r d movement. I n f i g . 7 h,
h o w e v e r , n o r the i n v a g i n a t e d p a r t o n the left, n o r the v i l l u s o n the r i g h t
(in
the
sections w h e r e
i t meets the
s k i n ) are a c c o m p a n i e d b y
strong
muscles. C o n s i d e r i n g the v e r y loose a n d changeable t e x t u r e o f the c o n n e c t i v e tissue, it seems possible, that the total disappearance o f the v i l l i
really
takes p l a c e : the elements
o f the tissue s h i f t i n g u n t i l a s o r t o f
e q u i l i b r i u m o r r e g u l a r dispersement o f the f i b r e s i s attained (cf. J o r d a n ,
1901,
f i g . 3 ) . S o m e h o w , h o w e v e r , the f o r m o f the ( o f t e n b r a n c h e d ) v i l l u s
must be p r e s e r v e d i n the a r r a n g e m e n t o f the tissue, so that o n r e l a x a t i o n
of
the muscles, o r some other
m e c h a n i c a l cause,
the v i l l u s is
formed
again. T h i s total r e t r a c t i o n is yet a p r o b l e m a n d the o n l y w a y to solve i t
completely, i s a n i n v e s t i g a t i o n i n the l i v i n g a n i m a l .
We
can o n l y say, that w e have no reason to suppose that the
presence
214
H.
ENGEL
or absence o f v i l l i i s a s p e c i f i c character, tubercles are to be r e g a r d e d as
retracted v i l l i . I n the g e o g r a p h i c a l s e c t i o n there a r e g i v e n m o r e details
about the m a n y a n i m a l s w e c o u l d compare, there the v a r i a b i l i t y o f the v i l l i
i n a r r a n g e m e n t a n d state o f p r e s e r v a t i o n becomes v e r y o b v i o u s .
T h e shell ( f i g . n )
i s o f the c o m m o n f o r m . I n the 153 m m l o n g a n i m a l
f r o m N i a s i t is 4 9 m m l o n g a n d 38 m m b r o a d . T h e c u t i c u l a r p o r t i o n i s
o n l y some m i l l i m e t e r s l a r g e r at the m a r g i n t h a n the calcareous l a y e r . T h e
s p i r e as u s u a l w i t h a h e a v i l y calloused s m o o t h m a r g i n .
I n f i g u r e 12 I have g i v e n the shells o f most a n i m a l s here e x a m i n e d , to
Fig. 11. Dolabella scapula (Martyn). T h e shell of the animal
of fig. 6. a, from above, b, from below. Natural size.
s h o w the v a r i a t i o n . F i g u r e 13 g i v e s f o r the same reason the shells f i g u r e d
in
literature
under
different
s p e c i f i c names, but, a c c o r d i n g to me, a l l
belonging to the one species D. scapula.
Internal
characters.
L i t e r a t u r e . R u m p h i u s (1705, p. 1 2 3 ) : s h e l l lies i n the f l e s h ; C u v i e r
(1804, pp. 4 3 7 - 4 4 0 ) : s t r o n g r e t r a c t o r muscles o f the head r e g i o n ; A m a u d r u t (1886, p. 6 8 ) : n e r v o u s s y s t e m ; M a z z a r e l l i & Z u c c a r d i ( 1 8 9 2 ) : f i r s t
detailed a n a t o m y ; G i l c h r i s t (1895, p. 2 6 4 ) : p a l l i a l c o m p l e x ; P i l s b r y (1896,
p.
150): radula o f
D. californica;
E l i o t (1900, p. 5 1 6 ) : some
remarks;
T h i e l e (1900, p. 251) : r a d u l a ; F a r r a n (1905, p. 3 5 4 ) : r a d u l a ; B e r g h (1905,
pp. 13-20) : d e t a i l e d a n a t o m y ; B e r g h (1907, pp. 1 5 - 1 9 ) : d e t a i l e d a n a t o m y ;
Vayssiere
(1906,
p.
71):
radula;
MacFarland (1918): very
detailed
d e s c r i p t i o n ; R i s b e c (1928, p. 4 5 ) : i m p o r t a n t details, f i g u r e s s i t u s v i s c e rum;
O ' D o n o g h u e (1929, pp. 33-34) : r a d u l a a n d m a n d i b l e s ; H i r a s e (1929,
pp. 1 0 6 - 1 0 8 ) : g e n i t a l o r g a n s ; P r u v o t (1934, p . 4 5 ) : b u c c a l a r m a t u r e o f
THE
GENUS
215
DOLABELLA
Fig. 12. Dolabella scapula (Martyn). Shells of the animals here examined, all seen
from above, to show the variation, a, b, Telok Dalam (Nias); c, Sabang; d, Java
(leg. V a n Hasselt); e, H o o r n Island; /, Bay of Batavia (leg. Sluiter); g, Kerkhof
Island; h, Pulu Panggang; i, j, k, Sailus Ketjil; /, Bay of B i m a ; m, Cyrus B a y ;
*J 0, p, q Pepela B a y ; r, s, Sanana Bay; t, Moluccas (Van der Hucht coll.); u, v,
w, x, y, Moluccas (De Seriere coll.); z, a', Moluccas; b' Ambon (Bleeker) ; c , Ambon
(Forsten); d', Ambon (Ludeking); e', Banda; f, New Guinea; g', Waigeu; h' Biak
Island;
Kaimana; f, k', m', Pulu Sanguisiapo; ri, *'Japan"; q' r', T i o p ; t', New
Caledonia; u', w', x\ y', Inhaca; z', a", b", c", Mauritius; d", e", f", g", without
locality. X
y.
t
f
t
t
t
2
2l6
H. ENGEL
the A s t r o l a b e a n i m a l i s i d e n t i c a l w i t h t h a t o f M a c F a r l a n d ' s a n i m a l ; B a b a
(1936, p . 17) : s o m e d e t a i l s .
Internal anatomy of our
largest specimen
from
Nias.
T h e s k i n o f the foot is v e r y t h i c k , i n f r o n t 9 m m , i n the m i d d l e 6 m m
Fig. 13. Dolabella scapula (Martyn). T h e shells copied from literature, a, Rumphius,
1705, pl- X I fig. N ; b, Martyn, 1784, III, fig. 99; c, Cuvier, 1804, pl. 29 fig. 2;
d, Sowerby, i820- 25, pl. 156; e, f, h, Quoy & Gaimard, 1832, pl. 23 Moll. figs. 5, 7, 3
(D. rumphii, tongana, hasseltii); i, /, k, Rang, 1828, pis. I l l , I, II (D. teremidi,
rumphii, ecaudata); nv, Deshayes, 18153, pl- Q fig- 45 > °» P> Q> Sowerby, 1868,
pl. II figs. 3c, 3b, 6, 4 (D. rumphii 2 X guayaquilensis, ecaudata) ; r, s, t, Mazzarelli
& Zuccardi, 1892, pl. 1 figs. 1, 2, 3 (JD. hasselti, teremidi, tongana); u, w, D. californica (Pilsbry, 1896, pl. 66 fig. 17, and Stearns, 1879, pl. V I I fig. 1); x, Clessin,
1899, pl. 10 fig. 1 (D. neira) ; y, Bergh, 1907, pl. X I V fig. 12; z, Thiele, 1929, I,
P- 398, fig. 501; a , O'Donoghue, 1929, pl. I l l fig. 33; V> Baba, 1936, p. 16, fig. 6 B ;
c' Allan, 1932, p. 422 (D. andersoni). X about fa
,
2
t
1
t
n
THE
GENUS
217
DOLABELLA
a n d b e h i n d 20 m m . T h e p h a r y n g e a l b u l b is 21 m m l o n g a n d 16 m m b r o a d .
T h e oesophagus is o n l y 5 m m w i d e . T h e mandibles ( f i g u r e 14 m) are about
10 m m l o n g a n d 4 m m b r o a d , the p o s t e r i o r p o r t i o n is c o v e r e d b y s k i n .
In
the t w o o t h e r specimens f r o m N i a s the measurements are n e a r l y the
same, perhaps the m a n d i b l e s are somewhat b r o a d e r a n d
shorter.
R a d u l a . T h e l i t e r a t u r e o n t h i s o r g a n is i n t e r e s t i n g because the m i d d l e
tooth f o r m s a point o f d i s c u s s i o n : M a z z a r e l l i & Z u c c a r d i (1892, p. 5, p i . I
figs. 11, 18, 19, 16, 2 0 ) : m i d d l e tooth w i t h a " c a p p u c c i o d i tre c a v i t a " ,
( i b i d . , p. 8, p i . I f i g . 12) : i d e m , 33-40 r o w s , ( i b i d . , p. 10, p i . I f i g . 14) :
i d e m ; P i l s b r y (1896, p. 150, p i . 67 f i g s . 17, 18) : m e d i a n tooth a r u d i m e n t a l
p l a t e ; E l i o t (1900, p. 5 1 6 ) : no c e n t r a l tooth o r c e n t r a l s p a c e ; T h i e l e (1900,
p. 251, f i g . 6 a, b ) : no r h a c h i d i a n to6th, but the m i d d l e tooth i s " o b v i o u s l y "
a coalescence o f the f i r s t side teeth o f each s i d e ; F a r r a n (1905, p. 354, p i .
V figs. 16, 17) : 44 r o w s o f about 120-1-120 teeth, m i d d l e t o o t h i s f i g u r e d
w i t h a cusp a n d a b i f i d plate l i k e T h i e l e ' s " c o a l e s c e d " o n e ; B e r g h (1905,
P-
J
S)
:
58-60 r o w s , t i l l 200 teeth each r o w , ( i b i d . , p i . V I I I
f i g s . 3-5) :
figures one r h a c h i d i a n tooth w i t h r u d i m e n t a l cusp a n d b i f i d plate
and
another w i t h a b e a u t i f u l 5-lobed cusp, ( i b i d . , p. 18) : 6 0 r o w s w i t h about
200 teeth each r o w , ( i b i d . , p i . V I I I f i g . 9 ) : a b e a u t i f u l m i d d l e tooth w i t h a
d e n t i c u l a t e d cusp, l i k e f i g . 4 ; ( i b i d . , p. 1 9 ) : 61 r o w s , ( i b i d . , p i . I X figs.
1-3): o n l y side teeth f i g u r e d , but the text says, that the r h a c h i d i a n teeth
are l i k e the p r e c e d i n g ; B e r g h (1907, p. 17, p i . X I V figs. 14-17)-: m e d i a n
tooth w i t h b e a u t i f u l l y d e n t i c u l a t e d cusp, 86 r o w s w i t h about 2 0 0 teeth
e a c h ; V a y s s i e r e (1906, p. 71, p i . I f i g . 3 ) : 40-50 r o w s , 120-160 teeth o n
each side, f i g u r e s a m e d i a n tooth w i t h a s m a l l s i m p l e c u s p ; M a c F a r l a n d
(1918, p. 3 1 6 ) : 62 r o w s w i t h 198-230 side teeth o n each side, ( i b i d . , p i . I V
figs. 3-4, 6 - 9 ) : m e d i a n tooth w i t h a s m a l l i r r e g u l a r k n o b o n the top o r w i t h out t h i s ; O ' D o n o g h u e (1929, p. 34) : 87 r o w s , 240-250 side teeth o n each
side, m e d i a n tooth s m a l l r e c u r v e d t r i a n g u l a r blade p r o v i d e d w i t h a n u m b e r
o f denticles, but somewhat v a r i a b l e ; P r u v o t (1934, p. 45, f i g . n ) : m e d i a n
tooth as i n M a c F a r l a n d ; B a b a (1936, p. 17, f i g . 6 E ) : 70 r o w s , 275-1-275,
m e d i a n t o o t h w i t h a v e r y s m a l l t r i a n g u l a r cusp.
It is clear that the m e d i a n tooth has
a tendency
towards
reduction,
t h o u g h one m a y doubt T h i e l e ' s i n t e r p r e t a t i o n o f h i s m i d d l e t o o t h as the
coalesced f i r s t side teeth. It has been i m p o s s i b l e to m e to f i n d a s p e c i m e n
w i t h s u c h b e a u t i f u l cusps as B e r g h figures them, a n d here also
doubt
arises, w h e t h e r denticles are ever present o n the cusp.
I n the largest N i a s - s p e c i m e n the r a d u l a is 13.5 m m l o n g , 17.5 m m b r o a d .
I counted t i l l 213 teeth i n one r o w o n one side (about the 30th r o w ) a n d
218
H.
ENGEL
t i l l 71 r o w s . H e r e the m i d d l e t o o t h (see f i g u r e 14) does not d i f f e r v e r y
m u c h f r o m the side teeth i n the f o r m a n d the l e n g t h o f the cusp. It looks
l i k e the m i d d l e teeth f i g u r e d b y T h i e l e , F a r r a n , V a y s s i e r e (see
above).
T h e basal plate i s b i f i d as i f is f i g u r e d b y the n a m e d a u t h o r s , t h o u g h , as
my
f i g u r e shows, not i n every m e d i a n tooth. T h e side teeth are o f
the
c o m m o n f o r m . I n the radulae o f the other specimens f r o m N i a s I counted
68 r o w s ( r a d u l a 11.5 m m l o n g , 15.5 m m b r o a d , t i l l 235 side teeth o n one
Fig. 14. Dolabella scapula (Martyn). Radula and mandible. The numbers indicate
the rows and the place of the side teeth counted from the rhachidian tooth (nr. o).
Only the teeth indicated by a are from the second Nias animal (radula length
11.5 mm), the others from the larger specimen. T h e mandible is indicated by m,
the broken line shows the limit of the half that is covered by skin.
side at about the 2 5 t h r o w ) a n d 59 r o w s ( r a d u l a 18 m m l o n g a n d 16 m m
b r o a d , t i l l 199 teeth o n one side i n the 2 0 t h r o w ) . I n figure 14 I have g i v e n
some teeth o f the largest specimen, the m i d d l e p a r t s o f the 37th, 38th,
a n d 39th r o w , also the r h a c h i d i a n tooth o f the 6 8 t h r o w , seen f r o m the
side. O f the 38th r o w some m o r e side teeth are
figured in
different
p o s i t i o n s . M o r e o v e r I have g i v e n the m i d d l e t o o t h o f the 41st r o w o f the
second s p e c i m e n ( l e n g t h o f r a d u l a 11.5 m m ) to s h o w the v a r i a t i o n o f the
cusp. W h i l e the last side teeth w e r e somewhat d i f f e r e n t i n this s p e c i m e n
I have f i g u r e d some o f the last teeth o f its 3 3 r d r o w . I f I a m a s k e d to
g i v e the d i f f e r e n c e between these radulae a n d that o f D. gigas, I cannot
tell i t . T w o specimens f r o m I n h a c a
(near L o r e n c p M a r q u e s ) s h o w
the
same f o r m o f teeth.
T h e p a l a t a l folds a n d spines are the same as described b y M a c F a r l a n d
for D. agassizi.
THE
The
GENUS
219
DOLABELLA
p e n i s i s u n a r m e d i n t h i s species, g i v i n g a n i m p o r t a n t
difference
f r o m D. gigas, w h e r e i t bears spines. A s f i g u r e 15 shows, the glans i s also
m u c h s h o r t e r here. T h e p r a e p u t i u m has a r a t h e r t h i c k s k i n ( f i g . 15 a )
a n d three r e t r a c t o r muscles, the h i n d e r m o s t inserts i n the base o f the glans,
the other t w o m o r e f o r e w a r d . T h e glans i s v e r y s h o r t , m u c h s h o r t e r t h a n
i n D. gigas, i t i s r a t h e r f o l i a t e , w i t h a n a r r o w e r a n d a b r o a d e r f o l d o f
s k i n , each at a s i d e o f the s p e r m a t i c g r o o v e . T h e p r a e p u t i u m i s w r i n k l e d ,
here the s p e r m a t i c g r o o v e i s v e r y d i s t i n c t , as i t i s b o r d e r e d b y t w o t h i c k
r o u n d e d f o l d s o f s k i n . T h e l e n g t h o f the p r a e p u t i u m i n t h i s p r e s e r v e d
a n i m a l i s 50 m m , that o f the glans 23 m m ( i n D. gigas the glans i s 40 m m
l o n g ) . I n the t w o o t h e r specimens f r o m N i a s the relations a r e the same,
the l e n g t h i s 40 a n d 30 m m respectively f o r the p r a e p u t i u m a n d 18 a n d 13
m m r e s p e c t i v e l y f o r the glans. T h e penes o f a l l t h e specimens at h a n d w e r e
Fig.
body
15. Dolabella
cavity, with
scapula
its t h r e e
(Martyn).
P e n i s , a, u n o p e n e d as it is t a k e n o u t o f
retractor muscles;
b, p r a e p u t i u m o p e n e d ,
showing
the
the
s h o r t f o l i a t e u n a r m e d g l a n s t o the l e f t a n d the c o n s p i c u o u s g e n i t a l g r o o v e , c o n t i n u i n g
a l o n g the
praeputium.
e x a m i n e d a n d they a l l a r e b u i l t i d e n t i c a l l y , none s h o w i n g a n y v e s t i g e o f
a r m a t u r e , a n d a l l p r o v i d e d w i t h the s h o r t c o n i c a l glans.
A
beautiful
f i g u r e o f the e x t e n d e d
penis was given b y Q u o y and
G a i m a r d (1832, M o l l . , p i . 23 f i g . i ) , . f r o m w h i c h i t m a y be seen, to w h a t
l e n g t h the p r a e p u t i u m m a y stretch. H i rase (1929, p . 108) g i v e s f o r the
l e n g t h o f the e x t e n d e d p e n i s (i.e., the glans o n l y ) 42 m m , i t s b r e a d t h w a s
11 m m . T h e not e x t e n d e d p r a e p u t i u m w a s 90 m m l o n g . O n p . 109 the
same a u t h o r describes the m a t i n g ; the penis reached t i l l the base o f the
spermatocyst. T h i s a u t h o r figures the p e n i s ( p i . V f i g . 4). O t h e r d e s c r i p tions a n d f i g u r e s a r e g i v e n b y M a z z a r e l l i & Z u c c a r d i (1892, p p . 7, 9, 11);
B e r g h (1905, p p . 17, 19, p i . V I I f i g s . 38, 39, 4 0 ) ; B e r g h (1907, p . 18);
M a c F a r l a n d (1918, p p . 328-9); R i s b e c (1928, p . 48, f i g . 26); A d a m ( i n
l i t t e r i s ) tells m e , that h i s a n i m a l ( c f . A d a m & L e l o u p , 1938, p . 199) h a d
a n u n a r m e d penis.
220
H.
ENGEL
The living animal.
T h e c o n i c a l b o d y f o r m suggests a d i g g i n g m o d e o f l i f e . T h e head r e g i o n
w i t h its s t r o n g r e t r a c t o r muscles p r o b a b l y has a b o r i n g f u n c t i o n . I t can be
r e t r a c t e d to i n v a g i n a t i o n b y these muscles a n d t h e n p r o b a b l y
protrudes
a g a i n b y force o f the b o d y f l u i d , w h i c h b y the heavy m u s c u l a t u r e o f the
s k i n is pressed f o r w a r d . T h e d i s c p r o b a b l y functions as a resistance a n d
as a s u p p o r t f o r this
Figures
or
movement.
descriptions
of
the
living animals
were given b y :
Van
H a s s e l t (1824, p. 54) : l o n g branched v i l l i are r e t r a c t i l e ; R a n g (1828, p. 46
seq., pis. I - I I I , X X I V
f i g . 1 ) ; L e s s o n (1830, p. 2 9 3 ) ; E h r e n b e r g (1831) :
areae posticae r o t u n d a e l i m b u s membranaceus crispus e x p a n d i et c o n t r a h i
p o t e s t ; £ ) u o y & G a i m a r d (1832, pp. 303, 305, 306, p i . M o l l . 23 figs.
A d a m s & Reeve
(1848, p. 65, p i . X V I I I
1-7);
f i g . 4 ) ; Pease ( i 8 6 0 , p. 2 2 ) ;
A n g a s (1867, p. 2 2 8 ) ; S t e a r n s (1879, p. 3 9 5 ) ; M a r t e n s
(1879, P- 7 3 ) J
8
E l i o t (1900, p. 515, p i . X I X f i g . 3 ) : a d m i r a b l y i m i t a t i n g a m a s s o f o l d
s e a w e e d ; P i l s b r y (1896, p. 150 s e q . ) : copies m a n y o l d f i g u r e s ;
Farran
( ^ S . P- 3 5 4 ) ; B e r g h (1905, p i . I l l f i g . 1) ; M a c F a r l a n d (1918, pp. 306-7,
p i . I ) ; R i s b e c (1928, pp. 45-6) : o n c o l o u r change, v e r y i m p o r t a n t ; Iredale
(1929, p. 2 9 2 ) ; H i r a s e (1929, pp. 105, 1 0 9 ) : c o p u l a t i o n , egg
depositing,
( i b i d . , p i . V f i g . 1) ; A l l a n (1932, p. 4 2 2 ) : f i g u r e , ( i b i d . , p . 4 2 3 ) : description,
a n i m a l , e g g s ; H i r a s e (1934, p i . 120 f i g . 17) ; A n o n y m u s (1935, I I , p i . X C I
fig.
1 1 ) : b e a u t i f u l f i g u r e ; B a b a (1936, p. 17, f i g . 6 ) .
D r . J . V e r w e y ( i n l i t t e r i s ) was so k i n d to g i v e me h i s a n n o t a t i o n s o n a n
a n i m a l he observed
Purmerend
i n the B a y o f B a t a v i a , n e a r the east o f the
on January 23rd,
1929; " L a r g e violet animal. T h e
island
animal
creeps w i t h a v e r y regular movement, suggesting the use o f a c i l i a t e d s k i n .
T h e r e are no waves o n the w a t e r " .
H a b i t a t : R u m p h i u s (1705, p. 1 2 3 ) : deep i n the m u d ; C u v i e r (1804, p.
439):
t h i n c o v e r i n g o f m u d ; R a n g (1828, p. 4 6 s e q . ) : o n s a n d but m o s t
o f t e n i n the m u d ; Q u o y & G a i m a r d (1832, p. 305) : o n the r o c k s a m o n g
seaweeds a n d zoophytes, can l a y d r y d u r i n g l o w tide, ( i b i d . , 3 0 7 ) : a m o n g
r o c k s ; D u f o (1840, p. 2 0 2 ) : o h a n d i n the r o c k s ; A d a m s & R e e v e (1848,
p. 6 5 ) : f e e d i n g i n g r o u p s i n a s m a l l m u d d y inlet o f the s e a ; A n g a s (1867,
p. 2 2 8 ) : o n sandy m u d ; S t e a r n s (1879, p. 395 a n d 1894, p. 3 4 2 ) : d a r k
places i n pools left b y the t i d e ; M a r t e n s (1880, p. 306) : i n sand a n d m u d ;
E l i o t (1900, p. 5 1 5 ) : a m o n g seaweed g r o w i n g o n the s a n d ; H i r a s e (1929,
p. 105) : o n sand w i t h s e a g r a s s : Hyponia; A l l a n (1932, p. 4 2 3 ) : c r a w l o v e r
the m u d flats o r b u r y themselves i n the soft
mud.
P u r p l e f l u i d : C u v i e r (1804, p. 4 3 8 ) : glands are present, but P e r o n never
s a w a n a n i m a l r e p a n d i n g some f l u i d ; V a n H a s s e l t
(1824, p. 5 5 ) : v i o l e t
THE
coloured f l u i d ; E h r e n b e r g
221
GENUS DOLABELLA
( 1 8 3 1 ) : saw no f l u i d ; Q u o y & G a i m a r d (1832,
p. 307) : p u r p l e f l u i d ; A n g a s (1867, p. 2 2 8 ) : p u r p l e f l u i d ; M a r t e n s
(1879,
P- 7 3 8 ) : p u r p l e f l u i d ; M a r t e n s (1880, p. 3 0 6 ) : p u r p l e s l i m e ; E l i o t (1900,
p. 515) : copious p u r p l e f l u i d .
E m b r y o l o g y : H i r a s e (1929, p. i n ) .
Geographical
section
(the n u m b e r s i n h e a v y type refer to the map, f i g . 16)
Malayan Archipelago.
Limax marina tertia R u m p h i u s , 1705, p. 38, p l . X f i g . 5 : A m b o n ( 1 ) .
Derde soort der Opercula callorum (Limax marina), derde van de vijfde
Fig.
16. Geographical distribution of the genus Dolabella. A a, b, c, D. gigas;
O 1-50, D. scapula. F o r numbers see text: Geographical section.
soort KliMeevers
Rumphius,
1705, p. 123, p l . X L f i g . N : A m b o n
T h e a n i m a l o f R u m p h w a s called
D. rumphii
D. c alios a
by L a m a r c k
(1).
(1801, p. 6 2 ) ,
b y C u v i e r (1817, p. 3 9 8 ) , w h o i d e n t i f i e d i t w i t h h i s a n i m a l
c llected b y P e r o n at M a u r i t i u s . B l a i n v i l l e (1819, p. 3 9 5 ) , h o w e v e r , n a m e d
the last s p e c i m e n
D. peronii
shell by S o l a n d e r
D. rumphii f o r R u m p h ' s a n i m a l .
Patella auricularia was g i v e n to R u m p h ' s
and preserved
A c c o r d i n g to Iredale the name
(1786, p. 154). O f the same y e a r is the f o l l o w i n g name
w h i c h is better k n o w n a n d therefore p r e f e r a b l e ; let us hope, that M a r t y n
p u b l i s h e d his paper earlier i n the y e a r 1786 t h a n
(Patella) scapula M a r t y n , 1786, v o l . I l l ,
Dolabella spec. Q u o y & G a i m a r d , 1824,
(2), T i m o r (3).
Solander.
fig. 9 9 : A m b o n (1).
p. 4 2 2 : l i e s P a p o u s
(Rawak)
222
H.
Dolabella rumphii
(coloured
here)
figure
ENGEL
1824, p . 5 4 : P e p e r b a a i ,
V a n Hasselt,
in Rang,
preserved
animal
West
i n L e i d e n and
Java
described
(4).
Aplysia hasseltii R a n g ,
V a n Hasselt reproduced
Aplysia rumphii
1828, p. 4 9 , p i . X X I V f i g . 1 : c o l o u r e d f i g u r e o f
a n d described
(4).
R a n g , 1828, p. 4 6 , p i . I : T i m o r ( P e r o n leg.),
studied
also the a n i m a l s o f Q u o y & G a i m a r d (1824) f r o m W a i g e u a n d R a w a k ( 3 ) .
Aplysia ecaudata
R a n g , 1828, p. 47, p i . I I : a n i m a l s o f Q u o y & G a i m a r d
(1824) f r o m W a i g e u a n d R a w a k
Aplysia truncata R a n g ,
(2).
1828, p. 4 7 : a n i m a l o f Q u o y & G a i m a r d .(1824)
from W a i g e u and R a w a k (2).
Dolabella
spec. M . E . G r a y , 1850-4, p i . 270 f i g s . 1, 2 : C e y l o n (cf. P i l s b r y ,
1896, p. 160)
(5).
Dolabella rumphii
Ceylon)
T h u r s t o n , 1887, p. 2 2 : R a m e s w a r a m
Dolabella hasselti M a z z a r e l l i
ippines)
(Phil-
& Z u c c a r d i , 1892, p. 7 : S a n J a c i n t o
(Phil-
(6).
Dolabella tongana
ippines)
& Z u c c a r d i , 1892, p . 3 : S a n J a c i n t o
(6).
Dolabella teremidi M a z z a r e l l i
ippines)
M a z z a r e l l i & Z u c c a r d i , 1892, p . 9 : S a n J a c i n t o
(Phil-
(6).
Dolabella rumphii V o n M a r t e n s , 1894, p.
Dolabella neira C l e s s i n , 1899, P- 7 » pi2
only)
(Adam's Bridge,
(5).
9 3 : A m b o n (1).
X f i g . 1: B a n d a n e i r a
(shell
(1).
Dolabella rumphii
B e r g h , 1905, p. 1 3 : reef o f f A t j a t u n i n g
(W.
Coast
N e w G u i n e a ) ( 7 ) ; reef i n B a y o f B i m a (8) ; reef o f H a i n g s i s i ( 3 ) ; reef
o f P e p e l a B a y ( E a s t coast o f R o t t i I s l a n d )
Dolabella rumphii
var.
(3).
maculosa (= Hasseltii)
B e r g h , 1905, p. 18, p i . I l l
f i g . 1 : reef o f S a i l u s K e t j i l ( P a t e r n o s t e r I s l a n d s ) ( 8 ) ; reef o f Sasana-bay
( E a s t coast o f S u l a B e s i ) (9) ; reef o f the B o e k a o r C y r u s - b a y
coast o f the i s l a n d R o t t i )
P u l u Sanguisiapo
Dolabella scapula
Dolabella scapula
(12).
(5).
W i n c k w o r t h , 1927, p. 1 0 1 : K r u s a d a i - i s l a n d i n the G u l f
(5).
Dolabella tongana
gala
(10);
(11)-
F a r r a n , 1905, p. 3 5 4 ; South-east M o d r a g a m a n d south
of A d a m ' s B r i d g e (Ceylon)
of M a n a a r
( 3 ) ; anchorage o f T u a l ( K e i - i s l a n d s )
(Tawi-tawi-islands, Sulu-archipelago)
(South
A d a m & L e l o u p , 1938, p. 1 9 9 : P a l u - b a y near D o n -
THE
GENUS
223
DOLABELLA
R u m p h i u s notes (p. 3 8 ) that h i s Umax marina tertia has a s m a l l bone
o n the back, n o t l a r g e r t h a n a " d u b b e l t j e " (i.e. about a threepence) w i t h a
p r o j e c t i n g bent hook, i n the f o r m o f a h a m , a n d a g a i n ( p . 122-3) notes,
Limax marina
that p l . X I I f i g . N is the s h e l l o f
w h i c h l i v e s r a t h e r deep
i n the sand near C a s t l e V i c t o r i a a n d that it l o o k s l i k e the s h o u l d e r blade
o f a p i g . F r o m this a n d h i s d r a w i n g w e m a y i n f e r w i t h c e r t a i n t y that h i s
animal was a
Dolabella.
I n the H i s t o r y o f the genus at the b e g i n n i n g o f
this a r t i c l e , I have g i v e n the names, w h i c h succeeding a u t h o r s gave to t h i s
animal o f R u m p h i u s and its shell.
M a r t y n also f i g u r e d a s h e l l f r o m A m b o n .
T h e second d e s c r i p t i o n f r o m the E a s t I n d i e s w a s g i v e n b y V a n H a s s e l t ,
w h o notes, that the l o n g b r a n c h e d appendages o f the s k i n c a n s h o r t e n
a n d lengthen themselves a n d a r e v e r y s e n s i t i v e l i k e so m a n y tentacles. H i s
b e a u t i f u l d r a w i n g o f a l i v i n g specimen w a s g i v e n to R a n g f o r r e p r o d u c t i o n
b y the D i r e c t o r o f the L e i d e n M u s e u m . I t is s t i l l there, P r o f . B o s c h m a
was so k i n d to send i t to m e , R a n g ' s r e p r o d u c t i o n i s a d m i r a b l y g o o d a n d
exact. I n t h i s same M u s e u m a
Dolabella
is preserved b e a r i n g the l a b e l :
" D o l a b e l l a , K u h l et V a n H a s s e l t , J a v a " . Its length i n a l c o h o l i s 140 m m ,
its breadth 62 m m , i t s h e i g h t 45 m m . T h e s h e l l i s 4 4 X 32 m m . A c c o r d i n g to R a n g the l e n g t h o f the l i v i n g a n i m a l w a s 190 m m , V a n H a s s e l t
(1928, p . 55) gives 0.69 (this is i n " A m s t e r d a m - f e e t " )
the breadth he gives 0.38, i.e., ±
t e r i o r tentacles 0.07, i.e., ±
= ± 195 m m . F o r
107 m m a n d f o r the distance o f the a n -
19 m m . I n the o r i g i n a l d r a w i n g the a n i m a l ' s
l e n g t h is 182 m m , i t s breadth
is 97 m m , the distance
o f the
anterior
tentacles 17 m m . I n the preserved specimen, as s a i d 140 m m , 6 2 m m a n d
the
distance
between
the tentacula
18.5 m m . A l l o w i n g
f o r the
strong
c o n t r a c t i o n i n the p r e s e r v i n g f l u i d w e m a y c o n s i d e r i t v e r y probable, that
t h i s preserved a n i m a l i s the one V a n H a s s e l t h a d d r a w n b y the a r t i s t
J . T h . B i k , w h o h a d been sent to h i m after V a n R a a l t e h a d f a l l e n i l l (see
S i r k s , 1915, p . 106). O n M a y 25th V a n H a s s e l t w r o t e about t h i s a n i m a l
to P r o f e s s o r V a n S w i n d e r e n . T h i s letter w a s p u b l i s h e d i n the A l g . K o n s t en L e t t e r b o d e . W i t h the d r a w i n g w a s f o u n d a note i n about the same
w o r d i n g s . V a n H a s s e l t does not say, w h e t h e r
he f o u n d one o r m o r e
a n i m a l s , b u t he speaks o f a r a r a species, w h i l e the f o l l o w i n g a n i m a l , Bullaea alba is c a l l e d r a r i s s i m a . S o perhaps there were m o r e than one, w h e n
V a n H a s s e l t collected i n the P e p e r b a a i . H e w a s then t r a v e l l i n g i n the
W e s t o f J a v a s t a y i n g o n l y a short time o n the sea coast. O n Sept. 8 t h
o f that year h e d i e d . S o , I t h i n k , there is some p r o b a b i l i t y , that
w r i t i n g h i s letter to P r o f . V a n S w i n d e r e n he collected no more
a n d that the a n i m a l , that w a s f i g u r e d , has been p r e s e r v e d
after
Dolabella's
a n d sent to
224
H.
ENGEL
L e i d e n . W e m a y say w i t h m o r e c e r t a i n t y , that i t m u s t have been
from
the same l o c a l i t y , a n d f o r that reason suppose, that this p r e s e r v e d a n i m a l ,
w h e n l i v i n g , resembled the d r a w i n g . T h e d r a w i n g (see R a n g , 1828, p i .
XXIV,
f i g . 1) shows a n a n i m a l , w i t h m a n y a n d l o n g appendages. I n the
preserved specimen, h o w e v e r , w e f i n d o n l y some non-contracted b r a n c h e d
v i l l i ( d i f f i c u l t to f i n d , as they are l y i n g flat o n the s o m e w h a t ragged s k i n ) ,
some fine tops o f r e t r a c t e d appendages, a n d m a n y places, w h e r e the v i l l i
m u s t have been w h o l l y retracted. O u r s p e c i m e n h a d the p e n i s e x p a n d e d to
a length o f 52 m m , i t is u n a r m e d a n d c o n f o r m s to the d e s c r i p t i o n g i v e n
above.
W i t h t h i s k n o w l e d g e a n d that f u r n i s h e d b y o u r i n v e s t i g a t i o n o f the v i l l i
(see a b o v e ) , as also w i t h the i n f o r m a t i o n s o n the v a r i a b i l i t y g i v e n b y E l i o t
(1900) a n d R i s b e c ( 1 9 2 8 ) , w e m u s t come to the c o n c l u s i o n , that w e c a n
r e g a r d the d i f f e r e n t species o f R a n g , v i z . ,
cata,
living
imens,
ned"
A. rumphii, ecaudata
and
trun-
as d i f f e r e n t states o f p r e s e r v a t i o n o f one species. R a n g o n l y s a w the
A. rumphii at M a u r i t i u s . F o r the rest he s t u d i e d p r e s e r v e d
w h i l e o f A. hasseltii he o n l y h a d a d r a w i n g . E v e n the " s m o o t h
specskin-
A. ecaudata has s m a l l t u b e r c l e s ! A large c o l l e c t i o n o f Dolabella's,
as I have at present at h a n d , c l e a r l y s h o w s a l l degrees o f c o n t r a c t i o n o f the
tubercles, f r o m l o n g b r a n c h e d v i l l i to s m a l l p o i n t - l i k e holes i n the s k i n .
L i k e w i s e the t w o a n i m a l s f i g u r e d b y M r s . G r a y ( a c c o r d i n g t o P i l s b r y ,
1896,
p . 160 f r o m C e y l o n a n d b y h i m r e g a r d e d as D. ecaudata a n d D.
scapula) o n l y represent the c o m m o n f o r m . T h e fact, that P i l s b r y regards
her f i g . 1 as
D. ecaudata,
w h i l e i t c l e a r l y s h o w s m a n y p o i n t e d appendages,
indicates, h o w l i t t l e this a u t h o r r e a l l y h o l d s to the n e a r l y s m o o t h s k i n as a
separating character o f
D. ecaudata.
M a z z a r e l l i & Z u c c a r d i h a d s i x p r e s e r v e d a n i m a l s f r o m the P h i l i p p i n e
Islands a n d b r o u g h t t h e m under three d i f f e r e n t species: three u n d e r D.
tongana, t w o under D. teremidi a n d one under D. hasselti. T h e y ingeniousl y f o u n d as the d i f f e r e n c e between
D. rumphii
m a r k i n g s o f the last a n d the absence
and
D. hasseltii, the
dark
o f p a p i l l a e o n its tentacles, b u t
c u r i o u s l y a l l o w b l a c k m a r k i n g s a n d v e r y d i s t i n c t tubercles i n tonganal
I n D. teremidi a g a i n s m a l l papillae w e r e present o n the tentacles. S t u d y i n g
a l a r g e r m a t e r i a l w e cannot f o l l o w the authors i n these subtle d i s c r i m i n a tions, w h i c h o n nearer i n v e s t i g a t i o n seem u n i m p o r t a n t a n d p a r t l y
p r i n c i p a l l y f r o m the characters g i v e n b y the o r i g i n a l
Dolabella neira w a s
differ
authors.
founded o n l y o n a s h e l l , w h i c h p r o b a b l y i s a v a r i e t y
or a n a b n o r m a l i t y o f the c o m m o n f o r m .
F a r r a n h a d three specimens, w h i c h he r e g a r d e d as D. scapula, t h o u g h
"the p a p i l l a e w e r e m o s t numerous i n the largest s p e c i m e n . . . a n d a l m o s t
THE
GENUS
225
DOLABELLA
absent f r o m the s m a l l e s t " a n d t h o u g h the l i v i n g a n i m a l s w e r e "chestnut
b r o w n i n g e n e r a l effect, y e l l o w i s h basis m o t t l e d w i t h r e d a n d b r o w n , m a n y
y e l l o w tags o r spines a l l o v e r " .
B e r g h , s t u d y i n g the r i c h S i b o g a m a t e r i a l , came to the c o n c l u s i o n (I.e.,
p. 1 3 ) : " E i n i g e ( A r t e n ) s i n d n u r nach der F o r m - v a r i a b e l e n S c h a l e aufgestellt, u n d die anderen s i n d v i e l l e i c h t n u r V a r i e t a t e n einer w e i t v e r b r e i teten
A r t " . Accordingly,
maculosa. A b o u t
D. hasselti
he des/cribes
the p a p i l l a e he r e m a r k s ,
as:
D. rumphii
var.
that they are " z u m T h e i l i n
Griibchen retraktil".
B e r g h h a d the b a d c u s t o m , to dissect h i s a n i m a l s to pieces, so that o n l y
part
of
study
4
his
one
from
material
specimen
Station
is
from
301
preserved.
Station
(reef
of
Oi;
47
his
(reef
Pepela
Bay,
D. rumphii
B a y of
East
Bima)
Coast
I
could
(8)
of
and
island
R o t t i ( 3 ) . T h e s p e c i m e n f r o m S t a t i o n 4 7 measures 75 m m i n length, 41
i n breadth, 27 m m i n height, the s h e l l measures 28 X 2 4 m m . T h e
mm
penis i s o f the c o m m o n short a n d u n a r m e d f o r m . T h i s specimen has
its
v i l l i p a r t l y w h o l l y retracted, but o n the r i g h t d o r s a l p a r t o f the s k i n i s a
field
with
many
beautifully
preserved
compound
papillae.
The
four
specimens o f S t a t i o n 301 measure i n l e n g t h 7 0 , 70, 67, 41 m m respectively,
i n b r e a d t h 50, 4 8 , 53, 28 m m , i n h e i g h t 28, 28, 2 3 , 15 m m , the s h e l l measures
27 X 2 1 , 26 X 21, 2 8 X 2 0 , 17 X
i2y
2
m m respectively. T h e p e n i s i s o f
the c o m m o n f o r m . T h e s e specimens are a l l d i s t i n c t l y m a c u l a t e d a n d it is
not
quite
clear
to
me,
w h y B e r g h d i d not count these to his v a r i e t y
t h o u g h u n d e r that f o r m he r e m a r k s : " D i e H a u t k e g e l n aber k l e i n e r , m e h r
e i n f a c h u n d w e n i g e r z a h l r e i c h . D e r R a n d der S c h e i b e w e n i g e r
hervortre-
tend u n d w e n i g e r z a c k i g " . I cannot emphasize these differences. T h e v i l l i
are d i s t i n c t a n d o f t e n c o m p o u n d , but p a r t l y contracted, so that large p a r t s
of the s k i n , especially i n the. t w o s m a l l e r specimens, m a y be smooth, as it
is figured for D. ecaudata a n d D. tongana. T h e smallest specimen, h o w ever, has v e r y d i s t i n c t c o n i c a l v i l l i o n its d i s c . T h e tentacles a l l bear
p a p i l l a e . T h e penis i s o f the c o m m o n f o r m .
Of
B e r g h ' s v a r i e t y maculosa I c o u l d s t u d y 3 specimens f r o m Stat. 37
(reef o f S a i l u s K e t j i l , P a t e r n o s t e r Islands) ( 8 ) , 2 f r o m Stat. 193 ( S a n a n a
Bay,
E . coast o f S u l a B e s i ( 9 ) , 1 f r o m Stat. 299 (reef o f C y r u s B a y , S.
coast o f i s l a n d R o t t i ) ( 3 ) . A s stated above, it is not clear to me, w h y these
specimens s h o u l d f o r m the v a r .
maculosa, they
do not d i f f e r essentially
f r o m the others, e.g., the a n i m a l o f Stat. 299 is not m a c u l a t e d , w h i l e those
of
Stat. 301 are. I can n e i t h e r v e r i f y B e r g h ' s r e m a r k o n the s m a l l e r a n d
less n u m e r o u s v i l l a e . T h e here n a m e d specimens m e a s u r e : Stat. 37 length
118, 100, 53 m m , Stat. 193 l e n g t h 160, 70, Stat. 299 length 79 m m respectZoologische Mededeelingen X X I V
z
5
226
H.
ENGEL
i v e l y , the b r e a d t h is 6 5 , 4 0 , 3 3 , 8 0 , 4 0 , 4 5 , the h e i g h t 32, 34, 16, 30, 26, 33,
w h i l e the s h e l l measures 3 8 X 29, 34 X 25, 19 X 15, 4 6 X 34, 26 X 2 0 ,
28 X 21 m m . T h e a n i m a l o f Stat. 37 h a d been d r y f o r some time a n d
therefore
had a peculiar outlook.
F o u r animals f r o m Stat. 93 m e a s u r e 8 8 , 87, 64, 62 m m i n length, 54,
57* 39, 4
2
m m i n breadth, 34, ?, 2 3 , 18 m m i n height. T h e second is o n l y
a n e m p t y flat s k i n , the s h e l l o f the f i r s t a n i m a l measures 36 X 29, that o f
the t h i r d 26 X 21 m m . ' t h e p e n i s i s o f the c o m m o n f o r m .
T h e three a n i m a l s f r o m N i a s i s l a n d ( 1 3 ) , c o l l . K l e i w e g de Z w a a n , w e r e
described
above.
S a b a n g ( 1 4 ) , 1 specimen, leg. G . H e r m a n . L e n g t h 130, b r e a d t h 9 0 , height
60 m m . A flesh c o l o u r e d a n i m a l w i t h d a r k m a r k i n g s , w h i c h has the head
so s t r o n g l y retracted, that i t invaginates i n t o the body. T h e v i l l i are a l l
retracted, o n l y l e a v i n g here a n d there a tubercle, w i t h a s m a l l h o l e , w h i c h
sometimes yet shows the top o f the retracted v i l l u s . L i k e i n m o s t specimens
v i l l i o n the tentacles are present. T h e penis is large, u n a r m e d , o f the c o m f o r m . T h e s h e l l 41 X 27 m m .
mon
B a y o f B a t a v i a ( 1 5 ) , leg. C . P h . S l u i t e r , 1 specimen. A l s o flesh c o l o u r e d
w i t h d a r k m a r k i n g s . V i l l i here, h o w e v e r , n u m e r o u s , especially o n the d i s c ,
m a n y o f t h e m p a r t l y retracted. L e n g t h 160 m m , breadth 8 0 m m , height
65 m m , s h e l l 4 9 X 38 m m . P e n i s o f c o m m o n f o r m .
B a y o f B a t a v i a ( 1 5 ) , reef o f H o o r n i s l a n d , M a r c h 1st, 1930, l e g . D r . J .
V e r w e y et M i s s W . S. S. v a n B e n t h e m J u t t i n g . A b e a u t i f u l a n i m a l w i t h
distinct v i l l i , partly branching, partly half invaginated, and dark markings.
L e n g t h 130, b r e a d t h 79, h e i g h t 50 m m , shell 45 X 36. P e n i s o f the c o m m o n
form.
Bay
van
o f B a t a v i a ( 1 5 ) , K e r k h o f i s l a n d , J u l y 31st, 1927, leg. J h r . W . C .
H e u r n . L e n g t h 117, b r e a d t h 74, h e i g h t 3 0 m m , shell 38 X 35 m m .
T h i s flesh c o l o u r e d a n i m a l is a g a i n i n t e r e s t i n g , as its back shows o n l y
some d i s t i n c t v i l l i a n d f o r the rest is tuberculated. T h e s e tubercles
have
no clear holes, but they are m o r e o r less t u b e r c u l a t e d o r bear v e r y s m a l l
c o n i c a l elevations. T h e disc s h o w s m a n y branched v i l l i i n a w e l l p r e s e r v e d
state. P e n i s as u s u a l .
B a y o f B a t a v i a ( 1 5 ) , P u l u P a n g g a n g , D u i z e n d - ( T h o u s a n d ) islands, l e g .
Dr.
J . D . F . H a r d e n b e r g . L e n g t h 125, b r e a d t h
8 5 , h e i g h t 50 m m , s h e l l
40 X 26. T h i s a n i m a l preserved i n f o r m a l i n e remembers one s t r o n g l y o f
Van
H a s s e l t ' s d r a w i n g . U n d e r m a g n i f i c a t i o n the g r e e n i s h g r o u n d c o l o u r
shows s m a l l w h i t e p o i n t s . T h i s a n i m a l is i n t e r e s t i n g because i t shows the
v i l l i o f a blue g r e e n l i g h t e r c o l o u r . S o m e o f these are not yet
wholly
THE
GENUS
227
DOLABELLA
retracted, others s h o w themselves as a blue g r e e n b u t t o n i n the m i d d l e o f
the tubercle, o t h e r tubercles s h o w a s m a l l blue g r e e n hole w h e r e the v i l l u m
disappeared.
F r o m a l l these a n i m a l s f r o m the B a y o f B a t a v i a , a l l i n a d i f f e r e n t state
of
p r e s e r v a t i o n , most o f them s h o w i n g a c o m b i n a t i o n o f the
common
(D. rumphii a n d D. hasselti auct.) a n d the t u b e r c u l a t e d
(D. ecaudata, tongana, agassizi a u c t . ) , w e m a y a g a i n i n f e r , that the
v i l l o u s appearance
form
presence o f the v i l l i is no s p e c i f i c character, as these appendages are a l l
r e t r a c t i l e i n t o the s k i n .
I n the A m s t e r d a m G e o l o g i c a l I n s t i t u t i o n a recent shell f r o m T j i l a o e t e u r e u n (51) ( S o u t h coast o f J a v a ) i s preserved 4 8 m m l o n g , 33 m m b r o a d .
We
first
n o w come to a g r o u p o f a n i m a l s f r o m A m b o n a n d v i c i n i t y . T h e
t w o are o l d specimens
from
the L e i d e n
M u s e u m and show
the
characteristics o f a l o n g stay i n a l c o h o l , the smallest c u r i o u s l y l o o k s l i k e
R u m p h i u s * d r a w i n g , t h o u g h the v i l l i are m o r e c o n i c a l a n d p o i n t e d , w h i l e
m a n y o n the d i s c are r i c h l y branched.
A m b o n ( 1 ) , l e g . F o r s t e n . L e n g t h 138, b r e a d t h 6 3 , height 38, s h e l l 41 X
32 m m , v i l l i n u m e r o u s , m o r e s i m p l e o n the back, composed o n the d i s c ;
d a r k m a r k i n g s are v i s i b l e . P e n i s o f the c o m m o n f o r m .
A m b o n ( 1 ) , l e g . L u d e k i n g . L e n g t h 38, b r e a d t h 4 6 , height 35, s h e l l 31 X
24 m m . M a n y p o i n t e d c o n i c a l v i l l i , some b r a n c h i n g . P e n i s as u s u a l .
A m b o n ( 1 ) , l e g . B l e e k e r , L e n g t h 106, b r e a d t h 50, height 4 2 , s h e l l 33 X
22 m m . T h i s i s a flabby, faded specimen, large p a r t s o f the s k i n are quite
s m o o t h , o n l y some v i l l i a r e not contracted, there are some tubercles w i t h a
s m a l l pore.
M o l u c c a s ( 9 ) , 1858, l e g . V a n der H u c h t . L e n g t h 75, b r e a d t h 3 8 , height
30, shell 2 4 X 18 m m . A f l a b b y a n i m a l w i t h some rests o f d a r k m a r k i n g s
a n d some v i l l i . P e n i s v e r y fine, but o f the c o m m o n f o r m .
B a n d a ( 1 ) , l e g . V a n de V e l d e . L e n g t h 78, b r e a d t h 47, h e i g h t 3 4 , s h e l l
34 X 25 m m . T h i s is a s t r o n g l y contracted specimen, flesh c o l o u r e d , w i t h
a somewhat tuberculated s k i n , w h i c h o n m a g n i f i c a t i o n shows m a n y v i l l i
o r the rests o f these.
The
f o l l o w i n g specimens
from
N e w G u i n e a cannot be d i s t i n g u i s h e d
f r o m the p r e c e d i n g .
An
old animal from
the L e i d e n
M u s e u m w a s collected b y
Macklot
(1828) o n the coast o f N e w G u i n e a ( 1 6 ) . It is b a d l y p r e s e r v e d , b l a c k i s h
b r o w n , but c l e a r l y s h o w s the v i l l i a n d tubercles. L e n g t h 8 5 , b r e a d t h 4 8 ,
height 32, shell 32 X 25 m m .
228
H.
ENGEL
B i a k I s l a n d ( 1 7 ) , near N e w G u i n e a , 1915, leg. W . K . H . F e u i l l e t e a u de
B r u y n . L e n g t h 50, b r e a d t h 4 5 , height 32, shell 23 X 25 m m . T h e head
r e g i o n is s t r o n g l y retracted,
rugose,
i n v a g i n a t e d . T h e b o d y is s w o l l e n ,
s h o r t a n d b r o a d , g i v i n g the a n i m a l a c u r i o u s f o r m . T h e foot is s t r o n g l y
contracted,
rugose. T h e d i s c i s c l e a r l y tuberculated,
but the back o n l y
shows some i n d i s t i n c t tubercles. R e s t s o f d a r k m a r k i n g s are present.
K a i m a n a (16), S . W . N e w Guinea, i33°44' E , 3 ° 4 i
/
S., 2 8 - V I I I - 1 9 1 0 ,
on the reef, leg. L . H e n d r i k s . T w o specimens m e a s u r i n g i n length 102, 87,
b r e a d t h 52, 50, h e i g h t 4 2 , 4 0 m m respectively, s h e l l o f the l a r g e r specimen
31
X 22 m m . S o m e s i m p l e o r composed v i l l i o n the back a n d m a n y o n the
disc. R e s t s o f d a r k e r m a r k i n g s are present. P e n i s as u s u a l .
A l l these a n i m a l s f r o m the M a l a y a n R e g i o n o n l y strengthen me i n m y
o p i n i o n , that they a l l f o r m but one v a r i a b l e species, v a r i a b l e e s p e c i a l l y
i n its b o d y f o r m a n d i n its r e a c t i o n o n the p r e s e r v i n g f l u i d . C o m p o s e d
v i l l i are present, but they can be w h o l l y retracted i n the s k i n , l e a v i n g o n l y
indistinct
Red
traces.
Sea.
Dolabella hemprichii
1869,
E h r e n b e r g , 1 8 3 1 : C o s s e i r (18)
(quoted b y Issel,
P- 165).
Dolabella rumphii V a y s s i e r e , 1906, p. 69 (51) : G u l f o f A d e n ( 1 9 ) , t w o
f r o m D j i b o u t i a n d one f r o m a reef between D j i b o u t i a n d O b o c k .
T h e o n l y a n i m a l o f E h r e n b e r g w a s pale green w i t h d a r k e r m a r k i n g s .
T h e " r u g u l o s e b a n d s " o n the back a n d the fact, that E h r e n b e r g gives no
figures o f the l i v i n g a n i m a l , suggest a not quite fresh specimen. I m p o r tant for o u r k n o w l e d g e o f the v a r i a b i l i t y is E h r e n b e r g ' s r e m a r k : " A r e a e
posticae
rotundae
limbus
membranaceus
crispus
expandi
et
contrahi
potest". V a y s s i e r e got no notes o n the l i v i n g a n i m a l , he considers h i s three
a n i m a l s a n d the one o f E h r e n b e r g as i d e n t i c a l w i t h
D. rumphii
The villi
w e r e d i s t i n c t , o f d i f f e r e n t length a n d the c o l o u r v a r i e g a t e d i n l i g h t e r a n d
d a r k e r green.
S. E . A f r i c a .
Dolabella...
coll. P e r o n ( =
C u v i e r , 1804, p. 4 3 6 , p i . 2 9 f i g s . 1-3: H e de F r a n c e
Dolabella rumphii
C u v i e r , 1817, p. 398
(20),
= Dolabella peronii
B l a i n v i l l e , 1819, p. 3 9 5 ) .
Dolabella spec. Q u o y & G a i m a r d , 1824, p. 4 2 2 : He de F r a n c e ( 2 0 ) .
Aplysia rumphii R a n g , 1828, p. 4 7 : He de F r a n c e (20) ( c o l l . R a n g ) .
Aplysia hasseltii v a r . Q u o y & G a i m a r d , 1832, p. 307, p i . 23 f i g . 1: i l o t s
a u x cerfs ( H e de F r a n c e )
(20).
THE
GENUS
229
DOLABELLA
Aplysia dolabella D u f o , 1840, p. 2 0 2 : Seychelles a n d A m i r a n t e s ( 2 1 ) .
Dolabella rumphii S g a n z i n , 1843, P- 4 M a d a g a s c a r , H e de F r a n c e ( 2 0 ) .
Dolabella rumphii K r a u s s , 1848, p. 7 2 : N a t a l B a y ( 2 2 ) .
Dolabella rumphii A d a m s & R e e v e , 1848, p . 6 5 : M a u r i t i u s ( 2 0 ) .
Dolabella rumphii Deshayes, 1863, p . 5 3 : R e u n i o n ( i l e de B o u r b o n ) ( 2 0 ) .
Dolabella teremidi Deshayes, 1863, p. 5 3 : R e u n i o n (ile de B o u r b o n ) ( 2 0 ) .
Dolabella ecaudata M a r t e n s , 1869, p . 6 5 : Z a n z i b a r ( 2 3 ) .
Dolabella eiongata S o w e r b y ( R e e v e ) , 1868, spec. 2 : Seychelles ( t h i s is
I
:
probably o n l y a m o n s t r o u s shell, never f o u n d a g a i n ) ( 2 1 ) .
Dolabella rumphii
Dolabella rumphii
L i e n a r d , 1877, p . 5 3 : M a u r i t i u s ( n o t seen) ( 2 0 ) .
Martens,
1879, P-
73&: M o z a m b i q u e
(24)
and
rumphii M a r t e n s , 1880, p. 3 0 6 : M a u r i t i u s ( M o b i u s coll.)
rumphii A m a u d r u t , 1886, p. 6 8 : Seychelles ( 2 1 ) .
scapula T h i e l e , 1900, p . 2 5 1 : Z a n z i b a r ( 2 3 ) .
rumphii B e r g h , 1907, p. 1 5 : E a s t L o n d o n ( 2 6 ) .
scapula O ' D o n o g h u e , 1929, p. 3 1 : I n h a c a (near L o r e n c p
(20).
Q u e r i m b a islands ( 2 5 ) .
Dolabella
Dolabella
Dolabella
Dolabella
Dolabella
Mar-
ques) ( 2 7 ) .
O u r data o n the S o u t h A f r i c a n Dolabella's a r e m a n y , b u t w e have o n l y
t w o c o l o u r e d f i g u r e s , v i z . : Q u o y & G a i m a r d (I.e.) ( c o p i e d b y P i l s b r y , p l .
28 f i g . 3 4 ) a n d A d a m s & R e e v e (I.e.) ( c o p i e d i b i d e m , p l . 2 6 f i g . 2 8 ) . T h e
b e a u t i f u l f i g u r e a n d d e s c r i p t i o n o f Q u o y & G a i m a r d seem to be the best.
S o m e a u t h o r s report, l i k e thd last, a d a r k green
to d i r t y y e l l o w b o d y
c o l o u r , r e s e m b l i n g R a n g ' s f i g u r e o f D. rumphii. A c r i t i c a l s u r v e y o f the
l i t e r a t u r e s h o w s a w i d e v a r i a t i o n , as w i d e as i n the M a l a y a n specimens.
E s p e c i a l l y the Japanese authors (see b e l o w ) h a v e g i v e n us a g o o d i d e a
o f the c o l o u r v a r i a t i o n i n the l i v i n g a n i m a l a n d lead us to the c o n c l u s i o n
that Q u o y & G a i m a r d ' s
D. rumphii
and
D. hasseltii
a r e b u t one species.
I c o u l d e x a m i n e f o u r a n i m a l s collected b y P r o f . V a n d e r H o r s t , J u l y
x
934> at I n h a c a
(27), a small island near L o r e n z o Marques. These are
b e a u t i f u l l y preserved,
s h o w i n g the d a r k b r o w n a n d y e l l o w patches o n a
g r e e n i s h g r o u n d color. W h e r e the s k i n i s w e l l preserved, a n d the v i l l i o n l y
little contracted,
they resemble Q u o y & G a i m a r d ' s f i g u r e quoted
above,
but w h e r e i n one s p e c i m e n a l l appendages are retracted, i t q u i t e resembles
Q u o y & G a i m a r d ' s D. tongana. T h e a n i m a l s m e a s u r e i n l e n g t h 125, 94, 94,
93, i n b r e a d t h 8 2 , 55, 50, 4 8 , i n height 4 8 , 42, 39, 3 0 respectively, the s h e l l
o f the f i r s t a n i m a l 45 X 37, o f the last 32 X 26 m m . T h e penis has the
c o m m o n f o r m . T h e r a d u l a i s l i k e that described above.
230
H.
ENGEL
Japan
Dolabella
nov. s p e c ? H i r a s e , 1929, pp. 105-114, p i . V
f i g . 1:
Misaki
( S a g a m i B a y ) ( 2 8 ) ; O k i n a w a - i s l a n d (29) ; N a g a s a k i ( K i u s u ) (30) ; A m a kusa ( K i u s u ) ( 3 0 ) ; Nabuto
(Bosu)
(31) ; K o m i n a t o ( B o s u )
(31).
Dolabella scapula H i r a s e , 1934, p i . 120 f i g . 17 ( w i t h o u t l o c a l i t y ) .
Dolabella spec. A n o n y m u s , 1935, p i . X C I f i g . 1 1 : c o m m o n i n C e n t r a l
Southern
and
Japan.
Dolabella scapula
B a b a , 1936, pp. 15-17, t e x t f i g . 6 : O k i n a w a
islands
( 2 9 ) ; Nagasaki (30), T o m i o k a (on Amakusa-island) (30), M i s a k i (Sagami
b a y ) ( 2 8 ) , K o m i n a t o ( P a c i f i c coast, E. o f T o k i o ) (31) a n d N a b u t o
(31).
B e a u t i f u l f i g u r e s , t w o c o l o u r e d a n d t w o i n b l a c k a n d w h i t e , are g i v e n
by these a u t h o r s , the best w e possess t i l l n o w o f the l i v i n g a n i m a l . T h e
c o l o u r v a r i a t i o n is v e r y large, the d a r k patches m a y be conspicuous
or
absent. T h e r e is no reason to d i s t i n g u i s h subspecies, based either o n the
c o l o u r o r o n the v i l l i . B a b a mentions no t r a n s v e r s e r o w s o f v i l l i ,
though
H i r a s e as w e l l as A n o n y m u s f i g u r e them, a r r a n g e d i n that w a y .
I c o u l d e x a m i n e one s p e c i m e n f r o m the B e r l i n M u s e u m " a n g e b l i c h v o n
J a p a n " , that s h o w e d no c o l o u r . T h e l e n g t h is 77, the b r e a d t h 38, the height
32 m m , the s h e l l 31 X 21 m m . T h e p e n i s o f the c o m m o n f o r m . T h e v i l l i
are r e t r a c t e d i n the p o s t e r i o r p a r t o f the back, but d i s t i n c t o n the a n t e r i o r
p a r t a n d o n the head.
Australia.
Dolabella scapula A n g a s , 1867, p. 2 2 7 : P a r a m a t t a
Dolabella rumphii B r a z i e r , 1878, p. 8 8 : D u n g e n e s s
river (32).
a n d D a r n l e y islands,
T o r r e s S t r a i t s ( 3 3 ) ; L o w I s l a n d , T r i n i t y B a y , H o m e Islands, N . E . A u s t r a l i a , o n the reefs u n d e r c o r a l i n s m a l l pools ( 3 4 ) ; P o r t J a c k s o n (32)
and
Bellinger river (35).
Dolabella ecaudata
B r a z i e r , 1878, p. 8 9 : H o m e Islands,
N.E. Australia
(34).
Dolabella rumphii S m i t h , 1884, p. 8 9 : N . E . A u s t r a l i a
Dolabella rumphii M a r t e n s , 1894, p. 9 3 : T h u r s d a y
Straits)
(34).
island
(Torres
(33).
Dolabella scapula H e d l e y , 1910, p. 3 7 1 : Q u e e n s l a n d coast ( 3 6 ) .
Dolabella ecaudata H e d l e y , 1910, p. 3 7 1 : Q u e e n s l a n d coast ( 3 6 ) .
Dolabella auricularia Iredale, 1929, p. 2 9 2 : Q u e e n s l a n d coast
(36)
( I r e d a l e c u r i o u s l y considers this Q u e e n s l a n d a n i m a l i d e n t i c a l w i t h R u m p h i u s ' a n i m a l f r o m as far as A m b o n , a n d calls i t w i t h the o l d s y n o n y m o f
Solander:
new n a m e :
D. auricularia,
D. andersoni.
w h i l e he g i v e s the N e w S o u t h W a l e s a n i m a l s a
W h y then do the Q u e e n s l a n d a n i m a l s not deserve
THE
ria, but preferable
231
DOLABELLA
D. scapula is
a new name ? A s s a i d above,
designation
GENUS
o f the same year as
D. auricula-
w h i l e better k n o w n a n d m o r e c o n f o r m i n g
Rumph's
"schonkje".
Dolabella andersoni
Iredale,
1929, p. 2 9 2 : N e w
South
Wales
Coast
(Gunnamatta B a y , P o r t H a c k i n g ) (32).
Dolabella ecaudata
39' S . , 1 1 7
0
T h i e l e , 1930, p. 5 8 6 : C o s s a c k ( N . W . A u s t r a l i a , 2 0
0
1 3 ' E . ) (37).
Dolabella andersoni A l l a n ,
near S y d n e y
1932, pp. 422 ( f i g . ) , 4 3 3 - G u n n a m a t t a B a y ,
(32).
I f w e compare the b e a u t i f u l f i g u r e g i v e n b y M i s s A l l a n (I.e.) w i t h the
f i g u r e a n d descriptions o f the Japanese f o r m , it i s not possible to see a n y
d i f f e r e n c e s . T h e characters noted b y I r e d a l e : " p o s t e r i o r not so a b r u p t l y
cut o f f , n o r i s the r i m edged w i t h b r a n c h i n g papillae, n o r are these so
n u m e r o u s o r so l o n g o n the rest o f the b o d y " are o f no value. O f course
A p l y s i i d a e are sometimes easily d i s t i n g u i s h e d , w h i l e l i v i n g , b y characters,
w h i c h they loose i n a l c o h o l . S o w e m a y hope, that the f u l l d e s c r i p t i o n ,
p r o m i s e d b y Iredale, w i l l g i v e these a n d w i l l be accompanied b y a c r i t i c a l
comparative
however,
study
the
of
the
figures
and
descriptions
N e w S o u t h W a l e s a n d the
already
Queensland
extant.
specimens
If,
really
d i f f e r , w e m a y presume, that the last also d i f f e r f r o m the A m b o n specimens, a n d these a g a i n f r o m the Japanese a n i m a l s . B u t w e m u s t a d d , that
these differences w i l l most p r o b a b l y be o f s u b s p e c i f i c value. A s f a r as o u r
present k n o w l e d g e goes, I cannot
distinguish geographical varieties. A s
l o n g as no better characters are k n o w n ,
D. andersoni
falls i n the s y n o n y m y
of the c o m m o n species D. scapula.
Pacific
Islands.
Dolabella
Islands
spec. Q u o y & G a i m a r d , 1824, p. 4 2 2 : M a r i a n n e o r L a d r o n e
(38).
Dolabella teremidi L e s s o n ,
(Aplysia teremidi R a n g , 1828, p.
D. temnida b y G r a y , 1857, p. 198,
f i g . 1 ( R a n g ' s f i g u r e ) a n d D. temnida
1830, p. 293
48, p i . I l l f i g . 1. E r r o n e o u s l y called
D. termida b y
M . E . G r a y , p i . 136
by M . E . G r a y , p i . 270 figs. 1, 2 ( n e w v e r y g o o d figures o f t w o w e l l preserved specimens
o f the c o m m o n f o r m ) ) :
S o c i e t y islands
(O-Taiti
=
T a h i t i , B o r a b o r a , ilot de T u b a i dans l a baie de B o r a b o r a ) ( 3 9 ) , C a r o l i n e
islands ( i s l a n d O u a l a n - K u s a i e )
Aplysia rumphii
I
(40).
Q u o y & G a i m a r d , 1832, p. 303, p i . 23 f i g s . 4-5 ( P r u v o t ,
9 3 4 , P- 4 4 ) • i s l a n d P a n g a i - M o d o u ( T o n g a - T a b o u o r T o n g a t a b u )
Aplysia tongana
(41).
Q u o y & G a i m a r d , 1832, p. 305, p i . 23 figs. 6-7 ( P r u v o t ,
232
H.
1934,
ENGEL
p . 4 4 ; called D. tongensis b y M . E . G r a y , p l . 137 f i g . 1) : i s l a n d
Pangai-Modou (Tonga-Tabou) (41).
Dolabella variegata Pease, i 8 6 0 , p . 2 2 : S a n d w i c h i s l a n d s
Dolabella hasseltii E l i o t , 1900, p . 515, p l . X I X f i g . 3 :
(43).
(42).
Apia
(Samoa)
O n e o f these specimens w a s used f o r c o m p a r i s o n b y M a c F a r l a n d
(1918, p. 3 0 1 ) .
Dolabella rumphii
C o u t u r i e r , 1907, p . 174: L a g o n de H a o ( P a u m o t u
island, Society Islands) (44).
Dolabella agassizf M a c F a r l a n d , 1918, p . 306, p l . I : E a s t e r Island ( 4 5 ) .
Dolabella rumphi
R i s b e c , 1928, p. 4 5 : P e n i n s u l a N o u m e a ( N e w C a l e -
donia) (46).
The
with
q u e s t i o n w h e t h e r the n o n - v i l l o u s species are o n l y c o m m o n a n i m a l s
retracted v i l l i , i s v e r y i m p o r t a n t f o r this r e g i o n .
D. tongana
and
D. agassizi b o t h are destitute o f v i l l i , b u t the figures s h o w a n i m a l s , that
look as i f they w e r e dead a n d somewhat c o r r u g a t e d , w h e n p i c t u r e d . B o t h
a u t h o r s m e n t i o n tubercles a n d i n D. agassizi they a r e o f the c o m m o n f o r m ,
as the f i g u r e shows. Q u o y & G a i m a r d say, that they disappear i n alcohol,
a n d c o n t i n u e : " B i e n qu* elle habite avec l a precedente
(D. rumphii),
on
ne peut p o i n t l a c o n f o n d r e avec elle". T h i s o f course i s a r e m a r k , that m u s t
be t a k e n i n t o c o n s i d e r a t i o n a n d the o n l y possible w a y o f r e a l l y s o l v i n g t h i s
p r o b l e m w o u l d be a n e x a m i n a t i o n i n loco. W e have, h o w e v e r , t w o authors,
themselves specialists i n t h i s f i e l d o f malacology, w h o collected o n P a c i f i c
i s l a n d s : E l i o t (1900) r e m a r k s : " I a m i n c l i n e d to t h i n k that
D. variegata
and
D. teremidi
D. hasselti,
a r e a l l one species" a n d R i s b e c ( 1 9 2 8 ) :
" B e r g h d e c r i t p l u s i e u r s varietes de
D. rumphi,
. . . m a i s j e c r o i s , ...que le
m e m e i n d i v i d u change de c o u l e u r s u i v a n t les saisons. A N o u m e a , . . . a
certaines epoques, tous les e x e m p l a i r e s sont v e r t c l a i r ; a d'autres, i l s sont
tous bruns, presque n o i r s . D e plus, ayant songe m o i - m e m e a. une v a r i e t e
d i f f e r e n t e , j ' a i constate q u ' e n faisant couler de l'eau s u r T a n i m a l , l a teinte
n o i r d i s p a r a i s s a i t a l a m a n i e r e d ' u n m u c u s . " U n h a p p i l y the a u t h o r g i v e s
no observations about the v i l l i o r tubercles, b u t i t is i m p o r t a n t , that b o t h
i n v e s t i g a t o r s stress the great v a r i a b i l i t y .
I c o u l d s t u d y t w o a n i m a l s f r o m the M u s e u m i n B e r l i n , collected at T i o p ,
B o u g a i n v i l l e (47) (one o f the S o l o m o n I s l a n d s ) , l e g . H . Schoede, o n the
3 r d a n d the 4 t h o f N o v e m b e r , 1909. T h e s e b e a u t i f u l l y p r e s e r v e d a n i m a l s
measure 93 a n d 110 m m respectively i n l e n g t h , b o t h 6 0 m m i n b r e a d t h , 4 0
a n d 4 3 m m respectively i n height, the shell measures 36 X 31 a n d 34 X 26
mm
respectively. S h e l l a n d penis o f the c o m m o n f o r m . T h e s m a l l e r a n i m a l
is g r e e n i s h , b e a u t i f u l l y m a c u l a t e d w i t h b r o w n , the l a r g e r a n i m a l has lost
its c o l o u r . T h e y b o t h s h o w d i s t i n c t v i l l i , t h o u g h i n some parts o f the b o d y
THE
GENUS
233
DOLABELLA
these have altogether disappeared. O n the disc o f the l a r g e r a n i m a l i t w a s
i n t e r e s t i n g to note a g a i n , h o w a branched v i l l u s c a n be w h o l l y retracted,
f i r s t a p p e a r i n g as a tubercle w i t h a c e n t r a l c o n i c a l c i r r u s a n d some s m a l l e r
appendages a r o u n d i t , u n t i l these are a l l retracted, c o n f i r m i n g V a n H a s selt's r e m a r k : " T h e apendices cutanei w h i c h i n R u m p h ' s figure are s i m p l e
w a r t s , are l o n g branched, p o i n t e d appendages
o f the s k i n , w h i c h
shorten
a n d lengthen themselves".
I n o u r collections there is a shell, l a b e l l e d :
"Dolabella sulcata B e r n a r d i .
N . C a l e d . . ab a u c t o r e " . A s f a r as I k n o w , B e r n a r d i never p u b l i s h e d t h i s
name. T h e shell has n o t h i n g that m i g h t d i s t i n g u i s h i t f r o m D. scapula.
W e s t Coast o f A m e r i c a .
Dolabella guayaquilensis
S o w e r b y ( R e e v e ) 1868, p i . I I f i g . 6 ( s h e l l o n l y :
G u a y q u i l (48) ( C o a s t o f E c u a d o r ) . M a r t i n i & C h e m n i t z e r r o n e o u s l y g i v e
quayaquilensis.
Dolabella californica
California
(=
Stearns, 1879, P- 3 9 5 M u l e g e B a y ( 4 9 ) , G u l f o f
:
Stearns, 1894, p . 341, 1895, p . 1 5 8 ) . P r e s e r v e d specimens
were e x a m i n e d b y P i l s b r y (1895, p . 7 3 ) a n d a g a i n P i l s b r y (1896, p . 159,
pi. 66 figs. 14-18).
Dolabella californica
P i l s b r y , 1895, p. 73 ( a n d a g a i n 1896, p . 159, p i . 66
figs. 14-18) e x a m i n e d the p r e s e r v e d type specimens f r o m M u l e g e B a y a n d
also one f r o m the W e s t C o a s t o f M e x i c o ( 5 0 ) .
Pilsbry
tends
to r e g a r d h i s specimens as i d e n t i c a l w i t h the s m o o t h
D. ecaudata a n d D. tongana, yet he mentions "traces o f sparcely scattered
w a r t l i k e p a p i l l a e " , w h i l e S t e a r n s o n the a u t h o r i t y o f the collector F i s h e r
says, that the surface is c o v e r e d w i t h w a r t l i k e papillae. W h i l e D. ecaudata
and
D. tongana w e r e
described as one-coloured green, P i l s b r y ' s a n i m a l s
s h o w e d a m o r e o r less b l a c k m a c u l a t i o n . W e see, that a l l these d i s t i n c t i o n s
are not r e g a r d e d as v e r y i m p o r t a n t b y most authors, none o f t h e m is used
as a constant character. T h e tuberculate specimens a r e c o n s i d e r e d i d e n t i c a l
w i t h the s m o o t h ones. T h e tuberculate specimens, h o w e v e r , m a y , as o u r
i n v e s t i g a t i o n s s h o w , be r e g a r d e d as v i l l o u s a n i m a l s w i t h retracted
villi;
v i l l i , tubercles a n d a s m o o t h s k i n most o f t e n o c c u r a l l at a t i m e i n the
same a n i m a l .
Our
c o n c l u s i o n m u s t be, that a l l these
D. scapula,
Dolabella,
f o r m b u t one s p e c i e s :
w h i c h c a n be e a s i l y a n d c l e a r l y d i s t i n g u i s h e d f r o m
D. gigas
by
the shell a n d the p e n i s .
F o s s i l species. T w o f o s s i l shells have been described u n d e r the genus
Dolabella.
B o t h w e r e f i g u r e d . N e i t h e r seems to b e l o n g to t h i s genus. E v e n
234
H.
ENGEL
not to the f a m i l y A p l y s i i d a e . A . Issel ( A l c u n i f o s s i l i n u o v i d e l Savonese.
A t t i S o c . l i g u s t i c a G e n o v a , S c . nat. geogr., 1922, 1, p. 123) d e s c r i b e d as
Dolabella (Sabaziella) texturata a
f o s s i l (pliocene) shell f r o m n e a r G e n u a .
T h e f i g u r e s h o w s a shell ( f r a g m e n t
?) w h i c h t h o u g h its general f o r m is
l i k e that o f Dolabella, has the s p i r e o n the r i g h t side, w h i l e Dolabella has
it o n the left side. M o r e o v e r the s h e l l o f
Sabaziella
s h o w s a series o f r i b s ,
a s t r u c t u r e w h i c h does not belong to the shells o f the A p l y s i i d a e , t h e i r
d i r e c t i o n a n d h a b i t u s is f o r e i g n to the s t r u c t u r e o f the A p l y s i i d shell. W e
can say w i t h c e r t a i n t y that
D a l l (where
Sabaziella texturata Issel
Dolabella aldrichi f r o m
?) d e s c r i b e d a
is no
Dolabella.
the t e r t i a r y o f F l o -
r i d a . T h e s h e l l w a s r e f i g u r e d b y C o s m a n n ( E s s a i de P a l e c o n c h o l o g i e c o m pared, I, P a r i s , 1895, p l . V I I f i g . 14). T h e s h e l l f r a g m e n t o f this f i g u r e
c e r t a i n l y s h o w s no
Dolabella.
Z o o g e o g r a p h y o f the G e n u s
T h e species
D. gigas
o c c u r s i n the R e d Sea, M a u r i t i u s , a n d the M a l a y a n
A r c h i p e l a g o , so is a c o m m o n i n h a b i t a n t o f the I n d o - W e s t - P a c i f i c R e g i o n .
But
f r o m a z o o g e o g r a p h i c a l p o i n t o f v i e w the species
D. scapula
is re-
m a r k a b l e , because it i n h a b i t s the w h o l e t r o p i c a l I n d i a n a n d P a c i f i c Oceans,
even the E a s t e r I s l a n d a n d the W e s t C o a s t o f t r o p i c a l A m e r i c a , w h i l e i t
fails i n the W e s t Indies.
A s E k m a n (1935, p. 105) r e m a r k s , the t i m e a n a n i m a l has to spend to
c i o s s the E a s t e r n P a c i f i c B a r r i e r f o r m s a n i m p o r t a n t factor i n i t s p a r t i c u l a r d i s t r i b u t i o n . W e k n o w n o t h i n g about the t i m e a
Dolabella
l i v e s as
a free s w i m m i n g l a r v a . H i r a s e (1921, p. 113) gave us some d a t a about the
development. B u t he o n l y c u l t i v a t e d the eggs d u r i n g the 8 days o f t h e i r
f i r s t development, t i l l they left the eggs as free V e l i g e r s . T h i s t i m e i s
t w i c e that a
Philine
w a n t s f o r that stage. P e r h a p s t h i s is a n i n d i c a t i o n ,
that the free s w i m m i n g stage also lasts r a t h e r l o n g here. Y e t it m a y be
doubted, w h e t h e r i t lasts l o n g e n o u g h to help the a n i m a l across the b r o a d
stretch o f O c e a n between M i d d l e a n d E a s t P a c i f i c .
T h e most probable e x p l i c a t i o n , therefore,
bella scapula seems
o f the d i s t r i b u t i o n o f
Dola-
to be, that o r i g i n a l l y the species o c c u r r e d i n the W e s t
I n d i e s a n d the w h o l e T e t h y s - S e a , w h i l e a f t e r w a r d s , i n the colder p e r i o d o f
the M i o c e n e , the species became e x t i n c t i n the W e s t Indies, but r e m a i n e d
on
the W e s t C o a s t o f M i d d l e A m e r i c a a n d i n the w h o l e t r o p i c a l I n d i a n
West-Pacific
Region. T h e question
rises w h e t h e r
the a n i m a l
remained
unchanged d u r i n g that l o n g t i m e o r o n l y f o r m e d local v a r i e t i e s , w h i c h w e
cannot d i s t i n g u i s h t i l l n o w .
THE
GENUS
235
DOLABELLA
Species erroneously placed i n the genus
Dolabella dolabrifera
Dolabella
Dolabrifera dolabri-
C u v i e r , 1817, p. 398, t h i s is
fera (cf. E n g e l et H u m m e l i n c k , 1936, p. 3 0 ) .
Dolabella rondeletii
C u v i e r , 1817, p. 398 ( f o u n d e d o n R o n d e l e t , 1554, p.
Aplysia de pilaris (cf. P i l s b r y , 1896, p. 160).
Dolabella laevis B l a i n v i l l e , 1819, p. 395, i.e., shell o f Aplysia depilans
520, 1558, p. 3 7 6 ) , i.e.,
(cf. P i l s b r y , 1896, p. 160).
Dolabella fragilis
Lamarck,
1822, p. 42, i.e., s h e l l o f
Aplysia depilans
(cf. P i l s b r y , 1896, p. 160).
Dolabella lepus R i s s o ,
1826, p. 44, p i . I figs. 1, 2, i.e.,
Aplysia fasciata
(cf. P i l s b r y , 1896, p. 160).
Dolabella unguifera
Lamarck,
1836, p. 701, i.e.,
Petalifera petalifera
(cf. E n g e l et H u m m e l i n c k , 1936, p. 4 3 ) .
Dolabella petalifera
1836, p. 702, i.e.,
Lamarck,
Petalifera petalifera
(cf. E n g e l et H u m m e l i n c k , 1936, p. 4 3 ) .
Dolabella ascifera
Lamarck,
1836, p. 702, i.e.,
Dolabrifera dolabrifera
(cf. E n g e l et H u m m e l i n c k , 1936, p. 3 0 ) .
Dolabella ornata Deshayes, 1853, p. 57, p i . 89 f i g . 5, i.e., p r o b a b l y Phyllaplysia lafonti (cf. E n g e l , 1936, p. 2 0 4 ) .
Dolabella oahouensis M a z z a r e l l i , 1893, P- 4°> *--> Dolabrifera dolabrifera
e
(cf. E n g e l et H u m m e l i n c k , 1936, p. 3 0 ) .
Dolabella maillardi M a z z a r e l l i , 1893, p. 40, i.e., Dolabrifera dolabrifera
(cf. E n g e l et H u m m e l i n c k , 1936, p. 3 0 ) .
Summary
O u r i n c r e a s i n g k n o w l e d g e o f the genus
v a r i a b i l i t y . R i s b e c (1928) pointed
Hasselt
Dolabella
tends to s h o w the great
to the v a r i a b i l i t y o f the colour. V a n
(1824) h a d a l r e a d y made t w o i m p o r t a n t r e m a r k s , one
concerning
the r e t r a c t i b i l i t y o f the v i l l i , that i n the l i v i n g a n i m a l cover the s k i n o f
the back, a n d the o t h e r h i s o p i n i o n , that R u m p h ' s a n i m a l , w i t h its tuberculate s k i n , is the same as h i s o w n
Dolabella
w i t h the b r a n c h i n g a n d l o n g
v i l l i . W e t r i e d to s h o w , that the v i l l i are v e r y retractile, even so that w e
came to the conclusion, that a l l species o f
Dolabella
but one, f o r m o n l y
one species, f i r s t d e s c r i b e d a n d f i g u r e d b y R u m p h i u s a n d f i r s t
named
b y M a r t y n : D. scapula, i n h a b i t i n g the I n d i e a n d the P a c i f i c f r o m the R e d
S e a a n d S . E . A f r i c a to J a p a n , A u s t r a l i a , H a w a i i a n d E a s t e r - I s l a n d ,
to the W e s t C o a s t o f A m e r i c a . O n l y the shell described as
even
Dolabella gigas
b y R a n g (1828) is s p e c i f i c a l l y d i f f e r e n t . It p r o v e d to belong to a n a n i m a l
that f u r t h e r d i f f e r s f r o m
D. scapida
i n h a v i n g the p e n i s l o n g , slender a n d
236
H.
a r m e d w i t h s m a l l spines, w h i l e i n
ENGEL
D. scapula
i t is s h o r t a n d u n a r m e d . I t
w a s recorded t i l l n o w f r o m M a u r i t i u s , the R e d S e a a n d the M o l u c c a s (see
map).
LITERATURE
ADAM,
W.,
et
E. LELOUP,
1938.
Prosobranchia
et
Opisthobranchia. Mem.
Mus.
R.
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de Paris (7), vol. io, p. 68.
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55, PP. 123-178.
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de mollusques a coquille cachee, nommee Parmacelle. A n n , d. Museum Nat.
d'Hist. Nat. vol. 5, p. 435.
, 1817. Reghe Animal, ie ed., Paris, vol. 2, p. 398.
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E H R E N B E R G , C. G., 1831. Symbolae physicae seu Icones et Descriptiones corporum
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THE
GENUS
DOLABELLA
237
E L I O T , C , 1900. Notes on Tectibranchs and Naked Mollusks from Samoa. Proc. Acad.
Nat. Sc. Philadelphia for 1899, vol. 51, p. 515.
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ENGEL,
H . , et P. W A G E N A A R
HUMMELINCK,
1936.
Uber westindische Aplysiidae
und
Verwandten anderer Gebiete. Capita Zoologica, vol. 8 pt. 1, pp. 1-76.
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et experimentale, vol. 30.
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2 8
H.
3
ENGEL
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&
CHEMNITZ,
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See
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_
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239
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