Commemorative volume for the 80th birthday of Willem Vervoort in 1997
The find of a whale barnacle, Cetopirus complanatus (Mörch,
1853), in 10th century deposits in the Netherlands
L.B. Holthuis, C . Smeenk & F.J. Laarman
Holthuis, L.B., C. Smeenk & F.J. Laarman. The find of a whale barnacle, Cetopirus complanatus (Mörch,
1853), in 10th century deposits in the Netherlands.
Zool. Verh. Leiden 323, 31.xii.1998: 349­363, figs 1­4. ­ ISSN 0024­1652/I SBN90­73239­68­0.
L.B. Holthuis & C. Smeenk, National Museum of Natural History, P.O. Box 9517, Leiden, The Nether­
lands.
F.J. Laarman, Rijksdienst voor Oudheidkundig Bodemonderzoek, P.O. Box 1600, 3800 BP Amersfoort,
The Netherlands.
Key words: Cetopiruscomplanatus;Cirripedia; whale barnacles; The Netherlands; archaeological find;
history; distribution; host species; Eubalaena; right whales; Eubalaena glacialis; northern right whale;
North Atlantic; North Sea; whaling.
A specimen of Cetopirus complanatus dating from the 10th century A.D. is described from archaeologi­
cal excavations at Tiel, the Netherlands. Two vertebral parts of northern right whales Eubalaena
glacialis: a vertebral arch and an epiphysis, were also found, possibly dating from the same period.
The disc­like epiphysis had been used as a cutting board. The specimens probably had reached Tiel
through early trade in whale products. Cetopirus complanatus is only known from right whales of the
genus Eubalaena. I t has not been found in the Northern Hemisphere since the late 19th century. I ts
host species in the North Atlantic and North Pacific, E. glacialis, is now very rare as a result of wha­
ling.
Introduction
D u r i n g a r c h a e o l o g i c a l excavations i n the t o w n centre of T i e l , p r o v i n c e of G e l d e r ­
l a n d , the N e t h e r l a n d s , s o m e a n i m a l r e m a i n s w e r e f o u n d d a t i n g f r o m the 10th c e n t u r y
A . D . T h e y consist of t w o parts of the v e r t e b r a l c o l u m n of a w h a l e : a n e p i p h y s i s a n d a
v e r t e b r a l arch, a n d of a w h a l e b a r n a c l e Cetopirus complanatus, a rather p o o r l y k n o w n
species, w h i c h is dealt w i t h b e l o w . T h e d i s c u s s i o n of that species is f o l l o w e d b y a
d e s c r i p t i o n of the a r c h a e o l o g i c a l context of the f i n d s a n d s o m e notes o n the occur­
rence of the host species, the n o r t h e r n r i g h t w h a l e Eubalaena glacialis,
i n the N o r t h
A t l a n t i c a n d N o r t h Sea.
Notes on Cetopirus complanatus ( M ö r c h ,
1853)
(L.B. Holthuis)
Material.— The specimen from Tiel (accession number 5­2­12, site code number 39D­256N; fig. la­b)
is the property of the National Service for Archaeological I nvestigations (Rijksdienst voor Oudheid­
kundig Bodemonderzoek), Amersfoort, the Netherlands. It was found in two pieces, but is surprising­
ly well preserved and consists of a complete set of the six compartments of the barnacle, total length
39 mm, width 33 mm, height 14 mm, dorsal opening 15 x 17 mm, lower opening 14 χ 15 mm.
T a x o n o m y . — Cetopirus complanatus is a n easily r e c o g n i z a b l e species, w h i c h has
b e e n k n o w n for a l o n g t i m e ; a n excellent f i g u r e of it w a s p u b l i s h e d as e a r l y as
1705.
350
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
Fig. la. Cetopirus complanatus from Tiel, the Netherlands, dating from the 10
ure: dorsal view; lower figure, oblique view.
th
century A . D . Upper fig-
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
Fig. lb. Cetopirus complanatus from Tiel, the Netherlands, dating from the 10
ure: ventral view; lower figure, lateral view.
th
351
century A . D . Upper fig-
352
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
There are n o great taxonomie p r o b l e m s . O l d e r authors, s u c h as D a r w i n a n d P i l s b r y ,
assigned the species to the genus Coronula L a m a r c k , 1802, b u t m o d e r n c i r r i p e d o l o gists agree that it s h o u l d be p l a c e d i n a separate genus Cetopirus R a n z a n i , 1817, of
w h i c h it is the o n l y species.
N o m e n c l a t u r e . — There are some n o m e n c l a t u r a l p r o b l e m s . F o r a l o n g time, the
specific n a m e balaenaris w a s u s e d f o r this species (as, e.g., b y D a r w i n , 1854), b u t Stebb i n g (1910: 572) m a d e clear that Lepas balaenaris O . F . Müller, 1776, is a j u n i o r s y n o n y m
of Lepas diadema L i n n a e u s , 1758 (= Coronula diadema). Stebbing p r o p o s e d a n e w name,
Coronula darwini, for the present species. P i l s b r y (1916: 277) f o u n d that the n a m e
Cetopirus complanatus Mörch, 1853 ( P i l s b r y gave the year as 1852, b u t M ö r c h ' s descript i o n w a s p u b l i s h e d i n 1853, i n the second part of h i s 1852-1853 " C a t a l o g u s " ) w a s the
first v a l i d n a m e u s e d for the species. Mörch (1853: 67) u n d e r "Ceteopirus [sic!] complanatus" referred to several p r e v i o u s authors, b u t clearly based the n a m e complanatus
o n Lepas complanata polythalamia described a n d f i g u r e d b y C h e m n i t z (1785: 325, p i . 99
figs 845, 846). P i l s b r y is f o l l o w e d b y m o s t zoologists a n d the n a m e Cetopirus complanatus is n o w generally accepted.
In the o r i g i n a l account of Cetopirus complanatus, Mörch d i d not indicate a h o l o type. Therefore, a l l the specimens o n w h i c h h i s o r i g i n a l p u b l i c a t i o n of C. complanatus
is based are syntypes. Tarasov & Z e v i n a (1957: 246) r e m a r k e d that the type s p e c i m e n
of the species w a s f i g u r e d b y C h e m n i t z (1785). U n d e r A r t . 74 (a) of the International
C o d e of Z o o l o g i c a l N o m e n c l a t u r e this c a n be considered the d e s i g n a t i o n of a lectotype, a n d thus the s p e c i m e n f i g u r e d b y C h e m n i t z (1785, p i . 99 figs 845, 846) is n o w
definitely accepted as the lectotype of Cetopirus complanatus.
N o t so s t r a i g h t f o r w a r d , h o w e v e r , is the nomenclature of the generic n a m e
Cetopirus. A c c o r d i n g to P i l s b r y (1916: 271, footnote 2), R a n z a n i (1818) i n d i c a t e d
Cetopirus balaenaris (recte Lepas balaenaris) as the type of Cetopirus. A c c o r d i n g to A r t .
70 of the International C o d e , the species selected b y a n author as the type species of a
genus is a s s u m e d to be correctly i d e n t i f i e d . A s Lepas balaenaris is a j u n i o r subjective
s y n o n y m of Lepas diadema L i n n a e u s , a n d the latter is the type species of Coronula, the
generic n a m e Cetopirus becomes a j u n i o r subjective s y n o n y m of Coronula. I n order to
save Cetopirus for C. complanatus, a n a p p l i c a t i o n has been s u b m i t t e d to the Internat i o n a l C o m m i s s i o n o n Z o o l o g i c a l N o m e n c l a t u r e requesting the use of their p l e n a r y
p o w e r s to designate the species Lepas complanata Mörch, 1853, as the type species of
the genus Cetopirus R a n z a n i , 1817. T h e suggestion to replace the n a m e Cetopirus b y
Ceteopirus cannot be accepted, as Ceteopirus is a m i s s p e l l i n g b y M ö r c h of Cetopirus a n d
thus has n o n o m e n c l a t u r a l status.
G e o g r a p h i c d i s t r i b u t i o n . — U n t i l recently, the m a t e r i a l of Cetopirus complanatus
present i n m a n y z o o l o g i c a l collections consisted of v e r y f e w fresh or w e l l - d o c u m e n t e d specimens. P i l s b r y (1916: 277), i n h i s m o n o g r a p h i c treatment of the g r o u p , s a i d o n
this account: "I h a v e seen a considerable series of this species, b u t o n l y three lots bear
locality data. O n e i n the collection of the A c a d e m y of N a t u r a l Sciences, P h i l a d e l p h i a ,
is labeled C a l i f o r n i a , b u t as it w a s p u r c h a s e d i n L o n d o n m a n y years ago, the label is
p r o b a b l y a p o c r y p h a l . O n e of the lots i n the U n i t e d States N a t i o n a l M u s e u m is labeled
W e s t A f r i c a , a n d another is i n a b o x w i t h the label V a l p a r a i s o , b u t these specimens
h a v e been i n the collection m a n y years, a n d the collector is n o t stated. I c a n not f i n d
that a n y definite records h a v e been p u b l i s h e d since D a r w i n ' s M o n o g r a p h [1854],
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
353
except those g i v e n b y G r u v e l [(1903: 152)] for specimens i n the Paris M u s e u m , f r o m
collectors of the first half of the last c e n t u r y . "
The collection of the L e i d e n M u s e u m c o n f i r m s P i l s b r y ' s observations: of the f o u r
d r y lots of C. complanatus, t w o (nos. 517 a n d 518) are w i t h o u t a n y i n d i c a t i o n of l o c a l i ty or date; n o . 518 bears the name E . A . Forsten, b u t it is n o t clear w h e t h e r this is the
collector or the d o n o r (Forsten collected i n the East Indies, a n d h i s material is a l w a y s
p r o p e r l y labeled). A t h i r d s p e c i m e n (no. 1321), also d r y , w a s collected a r o u n d 1900 o n
the beach of the N o r t h Sea near T h e H a g u e , the N e t h e r l a n d s . T h e s p e c i m e n is i n c o m plete, rather w o r n a n d d i s c o l o u r e d ; it m a y have been w a s h e d ashore after h a v i n g
been i n the water for a v e r y l o n g time. A n o t h e r d r y lot (no. 519) has the label St. Vaast
l a - H o u g u e , N o r m a n d y , France, w i t h o u t further details. T h e o n l y a l c o h o l l o t (no. 51)
contains t w o specimens f r o m C a p e of G o o d H o p e , S o u t h A f r i c a , collected there
between 1825 a n d 1838 b y H . B . v a n H o r s t o k , a D u t c h p h y s i c i a n i n C a p e t o w n , w h o
r e g u l a r l y sent material f r o m there to the L e i d e n M u s e u m . T h e specimens m u s t have
been taken f r o m a s o u t h e r n right w h a l e Eubalaena australis ( D e s m o u l i n s , 1822), as that
w a s the o n l y species of large w h a l e w h i c h V a n H o r s t o k obtained a n d of w h i c h he
sent one complete skeleton a n d one s k u l l to the L e i d e n M u s e u m . T h e w h a l e barnacles
e v i d e n t l y w e r e cut b y V a n H o r s t o k f r o m one o r b o t h of these whales, as part of the
w h a l e ' s s k i n is still attached to either specimen.
Occurrence i n the S o u t h e r n H e m i s p h e r e . — A l t h o u g h there are records f r o m the
N o r t h A t l a n t i c for Cetopirus complanatus, most of these are incomplete o r unreliable
a n d for a l o n g time it has been a s s u m e d that the species w a s restricted to the S o u t h e r n H e m i s p h e r e . D a r w i n (1854: 415) gave the d i s t r i b u t i o n as " A t t a c h e d to w h a l e s i n
the S o u t h e r n O c e a n " , a n d 132 years later Scarff (1986a: 130) d e f i n e d the range as
" S o u t h e r n H e m i s p h e r e " . B o t h P i l s b r y (1916: 277) a n d N e w m a n & Ross (1976: 45) liste d seven localities, of w h i c h o n l y one (coast of N o r w a y , based o n G r u v e l , 1903: 152),
w a s f r o m the N o r t h e r n H e m i s p h e r e (see b e l o w ) . T h e southern records so far k n o w n
are C h i l e ( C o q u i m b o , V a l p a r a i s o ) , B r a z i l , W e s t A f r i c a , S o u t h A f r i c a , K e r g u e l e n
Island,
Australia ( N e w South
Wales,
Tasmania),
a n d Indonesia
( K e i Islands,
A m b o i n a ) . M o s t of the k n o w n specimens of C. complanatus w i t h reliable locality data
were f o u n d i n the area i n h a b i t e d b y the southern right w h a l e Eubalaena australis.
H o w e v e r , there are some records f r o m the Southern H e m i s p h e r e that d o n o t fit
this picture. O n e of those is " A m b o i n a " , m e n t i o n e d b y P i l s b r y (1916) a n d cited b y
later authors. P i l s b r y ' s record is based o n the figure p u b l i s h e d i n G . E . R u m p h i u s ' s
1705 " D ' A m b o i n s c h e R a r i t e i t k a m e r " . I n this book, R u m p h i u s described a n d f i g u r e d
n u m e r o u s m a r i n e invertebrates ( M o l l u s c a , Crustacea, E c h i n o d e r m a t a , etc.) f o u n d i n
the r e g i o n of A m b o i n a , M o l u c c a s , N e t h e r l a n d s East Indies, n o w Indonesia. O n p i . 14
f i g . H , a n u n m i s t a k a b l e figure of C. complanatus is s h o w n (fig. 2). P i l s b r y ' s record
therefore
is perfectly
understandable.
What
he d i d not k n o w is that,
when
R u m p h i u s ' s m a n u s c r i p t a r r i v e d i n H o l l a n d to be p u b l i s h e d , the p u b l i s h e r w a n t e d it
to be m o r e c o p i o u s l y illustrated, i n order to f i n d a better market for it. I n the m e a n time, R u m p h i u s h a d d i e d i n 1702, a n d the e d i t i n g of h i s b o o k w a s entrusted to S i m o n
Schijnvoet, a n A m s t e r d a m amateur naturalist, w h o h a d figures m a d e of m a n y of the
u n i l l u s t r a t e d species m e n t i o n e d i n the " R a r i t e i t k a m e r " . F o r this p u r p o s e h e u s e d
specimens f r o m the n u m e r o u s private n a t u r a l h i s t o r y collections w h i c h at that time
a b o u n d e d i n the N e t h e r l a n d s . Schijnvoet d i d h i s best, b u t several times the figures
354
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
w h i c h he h a d m a d e d i d not represent the
species that they w e r e s a i d to illustrate.
Therefore, there are m a n y discrepancies
b e t w e e n text a n d figures. Fortunately,
Schijnvoet i n d i c a t e d w h i c h figures w e r e
not p r o v i d e d b y R u m p h i u s a n d m e n ­
t i o n e d the o w n e r of the s p e c i m e n after
w h i c h the f i g u r e w a s m a d e . P L 4 f i g . H is
s a i d i n the text to illustrate "Steentjes"
(small stones), w h i c h w e r e i d e n t i f i e d b y
v o n M a r t e n s (1902: 135) a n d E n g e l (1959:
217) as corals g r o w i n g a r o u n d a s i p u n ­
c u l i d o r a m o l l u s c . Schijnvoet's e x p l a n a ­
t i o n of p i . 14 f i g . H s h o w i n g C. complanaFig. 2. Cetopirus complanatus, pl. 14 fig. H in
tus reads ( i n translation): " F r o m the c o l ­
Rumphius: "D'Amboinsche Rariteitkamer",
lection of D r . D ' A c q u ê t [burgomaster of
1705.
Delft, a n d the o w n e r of a w e l l ­ k n o w n a n d
v e r y extensive cabinet of n a t u r a l curiosities] w e h a v e a d d e d t w o m o r e figures, one of
w h i c h is s h o w n w i t h the letter H , b e i n g stony a n d s t r o n g " . A s already r e m a r k e d b y
Stebbing (1910: 572), there is n o i n d i c a t i o n of the provenance of D ' A c q u e t ' s s p e c i m e n
of C. complanatus, a n d the locality A m b o i n a c a n therefore safely be deleted for this
species.
T h e r e c o r d " K e i I s l a n d s " b y N e w m a n & Ross (1976: 45) is p u z z l i n g . I h a v e not
been able to f i n d the source of it. I n h i s report o n the C i r r i p e d i a collected d u r i n g T h .
M o r t e n s e n ' s 1922 e x p e d i t i o n to the K e i Islands, B r o c h (1931) d i d n o t m e n t i o n the
species, neither i n the r u n n i n g text, n o r i n h i s table of the C i r r i p e d i a k n o w n f r o m
I n d o m a l a y a n waters. I h a v e n o t been able to f i n d a n y later reference to these islands
as a locality for C. complanatus. A s to P i l s b r y ' s (1916: 277) r e c o r d of the species f r o m
W e s t A f r i c a , this also falls largely outside the range of the r i g h t w h a l e s (see b e l o w ) . I n
h i s account of the c i r r i p e d f a u n a of t r o p i c a l W e s t A f r i c a , Stubbings (1967: 300) gave
strong arguments against the correctness of P i l s b r y ' s r e c o r d (Pilsbry h i m s e l f already
s o u n d e d d o u b t f u l about it) a n d c o n c l u d e d that " i t is u n l i k e l y that c o n f i r m a t o r y mate­
r i a l w i l l ever be f o r t h c o m i n g . " G r u v e l ' s (1903:152) r e c o r d of the species f r o m " A s t r o ­
labe: mers d u Brésil" also is rather d o u b t f u l . T h e A s t r o l a b e , at least o n h e r 1837­1840
e x p e d i t i o n , d i d not encounter w h a l e s i n B r a z i l i a n waters, a n d o n the o u t b o u n d trip i n
1826 d i d n o t t o u c h B r a z i l . I k n o w of n o later B r a z i l i a n records of C. complanatus. I n h i s
r e v i s i o n of the C o r o n u l o i d e a of the B r a z i l i a n coast, Y o u n g (1991: 190­212) d i d not
i n c l u d e it as a B r a z i l i a n species.
T h u s , the d e f i n i t e l y k n o w n range of C. complanatus i n the S o u t h e r n H e m i s p h e r e at
present comprises C h i l e , S o u t h A f r i c a , K e r g u e l e n Island, a n d A u s t r a l i a , c o i n c i d i n g
w e l l w i t h the range of Eubalaena australis. Recently, Pastorino & G r i f f i n (1996: 770)
reported H o l o c e n e fossil m a t e r i a l of C. complanatus f r o m A r g e n t i n a .
Occurrence i n the N o r t h e r n H e m i s p h e r e . — Pacific: P i l s b r y ' s (1916) d o u b t f u l
record of the species f r o m C a l i f o r n i a has n o t been c o n f i r m e d . Scarff (1986) sighted
barnacles o n a n o r t h e r n r i g h t w h a l e Eubalaena glacialis at 1.5 k m S W of P i l l a r Point,
near H a l f M o o n B a y , C a l i f o r n i a (37°30' Ν 122°03' W ) ; these barnacles, w h i c h w e r e
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
p h o t o g r a p h e d , s h o w e d to be " s i m i l a r to Coronula diadema, C. reginae, a n d Cetopirus
complanatus" (Scarff, 1986a: 131). O n p . 133, h o w e v e r , Scarff c o n c l u d e d "that the barnacles o n the right w h a l e w e r e a l l Coronula s p p . " , the p h o t o g r a p h s not b e i n g sufficiently detailed to a l l o w a definite identification. N o other records of C. complanatus
f r o m the N o r t h Pacific are k n o w n .
A t l a n t i c : The type locality of C. complanatus is i n the N o r t h A t l a n t i c . C h e m n i t z
(1785: 325-326) r e m a r k e d of the type material: "Derjenige Schiffer, w e l c h e r m i r e i n
paar Stücke v o n diesen [p. 326] raren flachen M e e r e i c h e l n überreichte, hatte d e n
N o r d c a p e r [= Eubalaena glacialis] auf der H ö h e z w i s c h e n N e u f u n d l a n d u n d Island
angetroffen." This type locality actually is rather precise, a n d it is d i f f i c u l t to see w h y
it w a s so completely i g n o r e d b y a l l subsequent cirripedologists. The
cetologists
Eschricht & R e i n h a r d t (1866: 35) p r o v i d e d some details about the capture of the type
material. T h e y stated that the vessel, " c a l l e d the ' C h r i s t i a n s h a v n ' , d i d really succeed,
o n one of the voyages (we suppose o n that of 1779), i n catching a ' N o r d k a p e r '
b e t w e e n N e w f o u n d l a n d a n d Iceland, the h e a d of w h i c h w a s infested w i t h s u c h a
m u l t i t u d e of C i r r i p e d s that it w o u l d have been easy, a c c o r d i n g to the statement of the
captain, to gather a w h o l e s a c k f u l of these ' w h i t e patches,' as he called t h e m . O n the
r e t u r n of the vessel to C o p e n h a g e n , C h e m n i t z , the d i s t i n g u i s h e d conchologist,
obtained a f e w specimens w h i c h the captain h a d b r o u g h t w i t h h i m , a n d recognised i n
t h e m the Balanus polythalamius
complanatus,
described b y W a l c h , ..." T h e y
also
r e m a r k e d that " t h i s a n i m a l exists o n l y o n r i g h t - w h a l e s " .
The other N o r t h A t l a n t i c records of C. complanatus are, as s a i d , f e w a n d i n c o m plete. A p a r t f r o m the t w o D u t c h specimens f r o m T i e l a n d The H a g u e m e n t i o n e d
above, there is a t h i r d reported f r o m the N e t h e r l a n d s b y N i l s s o n - C a n t e l l (1931: 116),
w h o listed a d r y s p e c i m e n i n the N a t u r a l H i s t o r y M u s e u m i n Basel, S w i t z e r l a n d , collected i n 1844 i n the d u n e s near K a t w i j k , a t o w n o n the D u t c h N o r t h Sea coast near
L e i d e n . The three D u t c h localities are a l l secondary: the specimens w e r e b r o u g h t
there after their death, a n d they most l i k e l y o r i g i n a t e d f r o m w h a l i n g or f r o m strande d w h a l e s . Since at the time w h e n the specimens w e r e f o u n d , there w a s n o D u t c h
w h a l i n g i n the S o u t h e r n H e m i s p h e r e , it can be a s s u m e d that a l l three came f r o m the
N o r t h A t l a n t i c , m a y b e e v e n f r o m the N o r t h Sea. The u n d a t e d s p e c i m e n f r o m St.
Vaast l a - H o u g u e i n the L e i d e n collection p o s s i b l y also resulted f r o m w h a l i n g . D e
Smet (1976: 2) r e m a r k e d that " i n the v i l l a g e of S a i n t - V a a s t - l a - H o u g u e (in N o r m a n d y ) ,
w h a l e bones w e r e so n u m e r o u s that they w e r e u s e d for several p u r p o s e s " (though he
e v i d e n t l y w a s referring to the situation i n the M i d d l e A g e s ) . A n o t h e r d r y s p e c i m e n i n
the Basel M u s e u m ( N i l s s o n - C a n t e l l , 1931) w a s s i m p l y labeled " N o r t h A t l a n t i c , 1871".
G r u v e l (1903: 152) listed a s p e c i m e n f r o m the N o r w e g i a n coast collected i n 1883 b y
the Pouchet e x p e d i t i o n . Scarff (1986a: 139-145) discussed several o l d records of b a r n a cles o n right w h a l e s i n c l u d i n g N o r t h A t l a n t i c specimens, b u t he f o u n d the available
data insufficient for a certain identification; the more, as the p o s s i b i l i t y exists that the
w h i t e callosities f o u n d o n right w h a l e s m a y h a v e been m i s t a k e n for barnacles.
The present f i n d of C. complanatus at T i e l s h o w s that the species m u s t have
o c c u r r e d i n the N o r t h A t l a n t i c i n the 10th century A . D . D e Smet (1976: 1) presented
evidence that w h a l i n g " w a s practised i n the N o r t h Sea a n d E n g l i s h C h a n n e l d u r i n g
the M i d d l e A g e s , surely f r o m the 9th century o n w a r d s . " The w h a l e s taken i n those
days w e r e almost e x c l u s i v e l y right w h a l e s . M u c h later, w h a l i n g e x p a n d e d a l l over the
355
356
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
N o r t h A t l a n t i c a n d w a s carried o u t b y different nations, w i t h right w h a l e s as the
p r i m e target w i t h i n that species' range; for further details, see b e l o w . Q u i t e l i k e l y , C.
complanatus w a s f o u n d o n the n o r t h e r n right w h a l e , a n d w h a l e r s took specimens
h o m e . I n this w a y , the species e n d e d u p i n c u r i o s i t y cabinets i n the N e t h e r l a n d s a n d
elsewhere. A t the time, the o w n e r s of those cabinets w e r e p r i m a r i l y interested i n the
b e a u t i f u l shape of their specimens a n d i n their scientific a n d v e r n a c u l a r names. Inform a t i o n about geographic o r i g i n a n d habitat w e r e h a r d l y ever n o t e d . O n l y i n the 19th
century, the interest i n the d i s t r i b u t i o n a n d b i o l o g y of the a n i m a l s b e g a n to increase,
b u t b y that time the n o r t h e r n right w h a l e , the source of the w h a l e barnacles, h a d
become rare (see b e l o w ) . T h i s explains w h y the greater part of N o r t h A t l a n t i c specimens of C. complanatus i n collections are d r y , o l d a n d w i t h o u t data.
The i d e n t i t y of the host species.— O n l y f e w p u b l i s h e d records of C. complanatus
are a c c o m p a n i e d b y a n i n d i c a t i o n of its host. T h e most detailed i n this respect is the
o r i g i n a l d e s c r i p t i o n of the type material. C h e m n i t z (1785: 325) gave the c i r r i p e d the
vernacular name " D i e L a u s des N o r d c a p e r W a l l f i s c h e s " , a n d o n the same page
r e m a r k e d : " m a n findet sie g e m e i n l i c h n u r auf solchen W a l l f i s c h e n , w e l c h e d e n
N a m e n der N o r d c a p e r führen"; furthermore, o n p . 326 he repeated that he received
the material f r o m a w h a l e r w h o h a d collected it f r o m a " N o r d c a p e r " b e t w e e n Iceland
a n d N e w f o u n d l a n d . A l t h o u g h the G e r m a n n a m e " N o r d c a p e r " or " N o r d k a p e r " a n d
the D u t c h " n o o r d k a p e r " have i n p o p u l a r accounts been u s e d for several cetacean
species, i n m o r e scientific literature it has a l w a y s been the name for the n o r t h e r n right
w h a l e Eubalaena glacialis. T h e correctness of C h e m n i t z ' s record seems c o n f i r m e d b y
o u r material f r o m T i e l , even t h o u g h the f i n d s d o not a l l o w a definite association of
the w h a l e barnacle w i t h the skeletal remains of E. glacialis f o u n d nearby.
The second host record for C. complanatus is that b y D a r w i n (1854: 417), w h o cited
the s o u t h e r n right w h a l e Eubalaena australis as its host species i n the S o u t h e r n H e m i sphere. Scarff (1986: 130) listed C. complanatus as the o n l y c o r o n u l i d u n t i l then reporte d f r o m right w h a l e s . H e h i m s e l f observed w h a t he thought to be Coronula o n Eubalaena glacialis; b u t h i s i n situ p h o t o g r a p h s of those barnacles d o not a l l o w a definite
identification a n d n o material w a s collected (see above). Scarff (1986a) reported the
right w h a l e as the o n l y host k n o w n w i t h certainty for C. complanatus (he r e g a r d e d the
n o r t h e r n a n d southern right w h a l e s as conspecific). T h e specimens of C. complanatus
f r o m S o u t h A f r i c a i n the collection of the L e i d e n M u s e u m , collected b y V a n H o r s t o k ,
c o n f i r m that E. australis is a host for this species. G u i l e r (1956: 3) reported C. complanatus (as Coronula balaenaris) f r o m the " f i n s a n d l i p s " of the h u m p b a c k w h a l e Megaptera
novaeangliae ( B o r o w s k i , 1781) f r o m M a r i o n B a y , T a s m a n i a . H o w e v e r , as this is the
o n l y r e c o r d of Megaptera as a host for Cetopirus, this observation s h o u l d be r e g a r d e d
w i t h s u s p i c i o n , as c o n f u s i o n w i t h Coronula reginae D a r w i n , 1854, seems l i k e l y .
The present data m a k e it clear that C. complanatus still occurs i n the S o u t h e r n
H e m i s p h e r e , a n d at least f o r m e r l y w a s also present i n the N o r t h e r n H e m i s p h e r e , at
least u n t i l the e n d of the 19th century. B o t h species of right w h a l e s b e l o n g i n g to the
genus Eubalaena are its host. It w o u l d be interesting to compare the material f r o m the
N o r t h e r n a n d S o u t h e r n H e m i s p h e r e , to s t u d y w h e t h e r or n o t there are a n y (sub)specific differences. Cetopirus complanatus has not been r e c o r d e d f r o m the other species of
the f a m i l y Balaenidae: the G r e e n l a n d right w h a l e o r b o w h e a d w h a l e Balaena mysticetus L i n n a e u s , 1758, a n d the p y g m y right w h a l e Caperea marginata (Gray, 1846).
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
357
T h e a r c h a e o l o g i c a l context o f the f i n d s at T i e l
(F.J. L a a r m a n )
The s m a l l city of T i e l is situated o n the r i v e r W a a l , w h i c h is n o w the m a i n a r m of
the r i v e r R h i n e i n the N e t h e r l a n d s . I n f o r m e r times, at T i e l the W a a l f o r k e d into the
W a a l p r o p e r a n d the r i v e r L i n g e (fig. 3). T i e l is first m e n t i o n e d i n A . D . 896 a n d i n its
h e y d a y , d u r i n g the second half of the 10th a n d the 11th century, it d e v e l o p e d into a
major trade settlement a n d as s u c h is thought to h a v e succeeded D o r e s t a d , situated
N W of T i e l o n the N e d e r r i j n , another c h a n n e l of the R h i n e delta. T h e t o w n ' s e c o n o m ic success i n this p e r i o d w a s d u e to its p o s i t i o n o n the trade route l i n k i n g the
R h i n e l a n d , the t o w n s o n the r i v e r M e u s e , a n d E n g l a n d . Its mintage rights also contributed to its economic rise. D u r i n g the 12th-14th centuries, h o w e v e r , Tiel's e c o n o m ic i m p o r t a n c e w a n e d , one of the reasons b e i n g that the riverbeds of the W a a l a n d
L i n g e shifted their p o s i t i o n . B y the e n d of the 14th century, the i m p o r t a n c e of this
once f l o u r i s h i n g port h a d d i m i n i s h e d to that of a s m a l l market t o w n (Sarfatij, 1997).
Fig. 3. Cadastral map of Tiel, showing the sites mentioned in the text.
358
H o l t h u i s et al. T h e f i n d of a w h a l e barnacle, Cetopirus complanatus. Z o o l . V e r h . L e i d e n 323 (1998)
The N a t i o n a l Service for A r c h a e o l o g i c a l Investigation (Rijksdienst v o o r O u d h e i d k u n d i g B o d e m o n d e r z o e k : R O B ) has since 1968 been r u n n i n g the project " U r b a n i s a t i o n i n the C e n t r a l R i v e r A r e a d u r i n g the M i d d l e A g e s " . T h i s project encompasses
inner-city archaeology i n the cities of Deventer, D o r d r e c h t , N i j m e g e n a n d T i e l . W i t h i n
the f r a m e w o r k of this project, i n 1996 a n d 1997 excavations w e r e p e r f o r m e d at v a r i ous locations i n T i e l . T h e w h a l e barnacle Cetopirus complanatus w a s f o u n d at the site
T o l - Z u i d (fig. 3). D u r i n g the excavations at the K o o r n m a r k t site, the e p i p h y s i s of a
w h a l e vertebra emerged, a n d the B l e e k v e l d site p r o d u c e d a vertebral arch of a large
w h a l e . B o t h specimens w e r e i d e n t i f i e d b y C . Smeenk a n d E.J.O. K o m p a n j e as b e l o n g i n g to the n o r t h e r n right w h a l e Eubalaena glacialis.
The w h a l e barnacle (find n o . 5-2-12) comes f r o m a f i l l i n g - i n deposit i n the h a r b o u r
zone of Tiel. A s the h a r b o u r silted u p w i t h s a n d f r o m the r i v e r W a a l , the q u a y s h a d to
be replaced o n several occasions. D u r i n g those constructions, the area b e t w e e n the
o l d a n d n e w quays w a s a l w a y s f i l l e d i n w i t h soil. B y means of d e n d r o c h r o n o l o g y the
f i l l i n g - i n deposit i n question w a s dated as b e i n g f r o m a r o u n d A . D . 986. T h e R O B site
name is T i e l T o l - Z u i d ; site code: 3 9 D - 2 5 6 N . T h e R O B m u n i c i p a l code is TL96-5; coordinates: 158.520/433.020.
The vertebral e p i p h y s i s of E. glacialis (find n o . 3-5-115) w a s f o u n d i n a w i d e d i t c h
i n the merchants' quarter of Tiel. T h e d i t c h dates f r o m the late 12th century, b u t it
cuts across the remains of a w a r e h o u s e that w a s b u i l t a r o u n d A . D . 965. H e n c e , the
bone m a y w e l l have been a m o n g d u g - o v e r debris f r o m the latter half of the 10th century. T h e b e a u t i f u l l y r o u n d , d i s c - l i k e bone s h o w s m a n y cuts caused b y k n i v e s a n d
h a d clearly been u s e d as a c u t t i n g b o a r d (fig. 4). T h e R O B site n a m e is T i e l K o o r n m a r k t ; site code: 3 9 D - 2 2 3 N . T h e R O B m u n i c i p a l code is TL96-4; coordinates: 158.360/
433.100.
The vertebral arch of the same species (find n o . 1-7-97) w a s l y i n g i n a d i t c h i n the
p e r i p h e r y of the settlement of T i e l , near the c h u r c h , w h i c h itself also dates f r o m the
late 10th century. A s yet, this excavation has n o t been f u l l y w o r k e d out. T h e R O B site
n a m e is T i e l B l e e k v e l d ; site code: 3 9 D - 2 7 0 N . T h e R O B m u n i c i p a l code is TL97-3; coordinates: 158.350/432.950.
A t present, T i e l is situated at about 100 k m f r o m the N o r t h Sea coast (about 80 k m
as the c r o w flies). It seems i m p o s s i b l e that the w h a l e s a n d attached c i r r i p e d a r r i v e d
here i n a n a t u r a l w a y . N e i t h e r does it seem l i k e l y that w h a l e s w e r e t o w e d to T i e l a n d
d i s m e m b e r e d there. It c a n o n l y be a s s u m e d that the large w h a l e s h a d been caught i n
the N o r t h Sea close to, o r h a d been w a s h e d ashore o n , the D u t c h coast. T r a d e i n
w h a l e p r o d u c t s m u s t have b r o u g h t the bones a n d w h a l e barnacle to T i e l .
T h e status a n d d i s t r i b u t i o n of Eubalaena glacialis i n the N o r t h A t l a n t i c a n d
N o r t h Sea
(C. S m e e n k )
The w h a l e genus Eubalaena G r a y , 1864, has a n antitropical distribution. M o s t recent
authors d i s t i n g u i s h t w o species: the n o r t h e r n r i g h t w h a l e E. glacialis ( O . F . Müller,
1776) a n d the s o u t h e r n right w h a l e E. australis ( D e s m o u l i n s , 1822). T h e southern
species has a c i r c u m p o l a r d i s t r i b u t i o n , r a n g i n g f r o m about 20° to 50° S. T h e o r i g i n a l
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
359
Fig. 4. Vertebral epiphysis of Eubalaena glacialis from Tiel, the Netherlands, possibly dating from the
10th century A . D . The disc­like bone has been used as a cutting board. The light­coloured scratches
were caused during the excavation.
range (i.e. before the p e r i o d of w h a l i n g ) of the n o r t h e r n right w h a l e p r o b a b l y extend­
e d f r o m about 25° to 60° Ν i n the N o r t h Pacific, f r o m about 25° to 75° Ν i n the N o r t h
A t l a n t i c ( C u m m i n g s , 1985; A g u i l a r , 1995). The A t l a n t i c a n d Pacific p o p u l a t i o n s of E.
glacialis are sometimes regarded as subspecifically or e v e n specifically distinct, the
Pacific f o r m b e i n g d i s t i n g u i s h e d as E. (glacialis) japonica (L acepède, 1818). R i g h t
w h a l e s s h o w regular n o r t h ­ s o u t h migrations, generally o c c u r r i n g at h i g h e r latitudes
(feeding grounds) i n s u m m e r , at l o w e r latitudes i n w i n t e r . Since they n o r m a l l y a v o i d
w a r m t r o p i c a l waters, the t w o species are n o w effectively isolated.
A l l right w h a l e s have been h e a v i l y e x p l o i t e d b y m a n . The h i s t o r y of w h a l i n g for
these species has been r e v i e w e d b y B r o w n e l l et a l . (1986) a n d K l i n o w s k a (1991). The
present n u m b e r s of right w h a l e s are o n l y a fraction of w h a t they m u s t h a v e been
before w h a l i n g started, a n d several stocks h a v e been h u n t e d to the verge of extinc­
360
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
tion. The southern right w h a l e E. australis s h o w s clear signs of recovery i n some areas
a n d , p r o v i d e d w h a l i n g is not r e s u m e d , appears n o longer i n danger of extinction.
The situation for the n o r t h e r n right w h a l e E. glacialis, h o w e v e r , is critical. It is n o w
the most endangered of the large w h a l e s a n d there are no signs of recovery. The
Pacific p o p u l a t i o n w a s e x p l o i t e d d u r i n g the p e r i o d 1840-1968 a n d is n o w thought to
n u m b e r o n l y a f e w h u n d r e d a n i m a l s (Scarff, 1986b; K l i n o w s k a , 1991). W h a l i n g i n the
N o r t h A t l a n t i c goes back to v e r y early times. The Basques w e r e the first to h u n t
w h a l e s systematically i n the B a y of Biscay, the first d o c u m e n t a t i o n d a t i n g f r o m the
11th century A . D . ( A g u i l a r , 1986). D e Smet (1976) presented evidence that w h a l i n g
w a s practised as early as the 9th a n d 10th centuries i n the E n g l i s h C h a n n e l a n d N o r t h
Sea. In a l l cases the n o r t h e r n right w h a l e w a s the p r i n c i p a l target of those early
w h a l e r s . The species often occurs v e r y close to the coast, i n s h a l l o w bays or lagoons,
p a r t i c u l a r l y d u r i n g the p e r i o d of c a l v i n g . T h e y are s l o w - s w i m m i n g a n d c o u l d therefore be h u n t e d w i t h h a n d h a r p o o n s a n d lances t h r o w n f r o m r o w i n g boats. D u e to
their thick layer of blubber, right w h a l e s stay afloat w h e n d e a d . Struck a n d lost a n i mals often d i e d later a n d were then w a s h e d ashore. A l l this, as w e l l as the large
a m o u n t of meat, o i l a n d baleen one a n i m a l p r o v i d e d , m a d e that this species w a s w i t h
reason called the " r i g h t " w h a l e , the o n l y one w i t h i n easy reach of the first w h a l e r s .
In later centuries, Basque w h a l i n g extended to other parts of the N o r t h A t l a n t i c .
In the early 15th century, the Basques reached Iceland a n d i n the 16th century they
established w h a l i n g settlements o n the coasts of N e w f o u n d l a n d a n d L a b r a d o r , w h i c h
w e r e o c c u p i e d u n t i l the early 18th century ( A g u i l a r , 1986). In the meantime, other
nations h a d j o i n e d i n , partly trained b y Basque whalers, a n d e v e r y w h e r e i n the N o r t h
A t l a n t i c the right w h a l e w a s b e i n g p u r s u e d relentlessly. D u r i n g the 19th century, the
p o p u l a t i o n s h a d d w i n d l e d to the extent that the N o r t h A t l a n t i c right w h a l e w a s b y
some considered extinct or nearly so (Reeves & M i t c h e l l , 1986b). In the N o r t h w e s t
A t l a n t i c , h o w e v e r , h u n t i n g c o n t i n u e d off L o n g Island, N e w Y o r k , till 1924 (Reeves &
M i t c h e l l , 1986a). In the Northeast A t l a n t i c too, the species w a s still caught d u r i n g the
first decades of the 20th century, m a i n l y f r o m the H e b r i d e s , w h e r e the last right
w h a l e s w e r e obtained i n 1923; the last one off N o r w a y w a s c a p t u r e d i n 1926 (see
B r o w n , 1986, w h o gives a total of 134-137 a n i m a l s taken i n the Northeast A t l a n t i c
b e t w e e n 1900 a n d 1926). The last N o r t h A t l a n t i c right w h a l e s k n o w n to have been
k i l l e d w e r e a mother a n d calf near M a d e i r a i n 1967 ( M a u l & Sergeant, 1977; B r o w n ,
1986).
A t present, there are a s s u m e d to be t w o p o p u l a t i o n s of E. glacialis i n the N o r t h
A t l a n t i c : a western a n d a n eastern stock (Reeves & M i t c h e l l , 1986b), t h o u g h it is not
quite clear w h e t h e r these have a l w a y s been m o r e or less separate, or w h e t h e r this
fragmentation is the result of w h a l i n g . The western stock is n o w estimated to n u m b e r
about 300 a n i m a l s a n d s h o w s no signs of recovery ( B r o w n et al., 1994; K r a u s , 1997).
The n u m b e r s of the eastern stock are u n k n o w n , b u t it appears to be o n the b r i n k of
extinction. V e r y f e w reliable sightings of right w h a l e s a n d n o strandings h a v e been
reported for the Northeast A t l a n t i c since 1900.
The occurrence of right w h a l e s i n the N o r t h Sea has been r e v i e w e d b y K o m p a n j e
& Smeenk (1996). T h e y f o u n d records of o n l y f o u r sightings/captures off the Scottish
a n d E n g l i s h N o r t h Sea coast i n the 17th, 18th a n d 19th century (Southwell, 1881;
E v a n s & Scanlan, 1989) a n d t w o strandings reported for the F l e m i s h coast: one i n
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
361
1178 w h i c h w a s tentatively i d e n t i f i e d as a r i g h t w h a l e , a n d o n e w e l l ­ d o c u m e n t e d case
i n 1751 ( M o l , 1962; D e Smet, 1974). F i n a l l y , K o m p a n j e & S m e e n k describe the f i n d s of
a f e w bones d r e d g e d b y f i s h i n g vessels i n the s o u t h e r n N o r t h Sea i n 1994 a n d 1995
( p r o b a b l y b e l o n g i n g to the same specimen), w h i c h they assume to be f r o m the second
h a l f of the 20th century. O b v i o u s l y , the n o r t h e r n r i g h t w h a l e h a s b e e n v e r y rare i n the
N o r t h Sea d u r i n g the last f e w centuries.
D e Smet (1974, 1981) s u m m a r i z e d the p r e h i s t o r i c f i n d s of r i g h t w h a l e b o n e s i n the
coastal districts of B e l g i u m a n d N W France; E v a n s & S c a n l a n (1989) refer to s i m i l a r
m a t e r i a l f o u n d near the E n g l i s h coast of the N o r t h Sea. S o m e i d e n t i f i c a t i o n s of this
o l d a n d often i n c o m p l e t e m a t e r i a l are tentative, b u t c o n f u s i o n w i t h the G r e e n l a n d
r i g h t w h a l e Balaena mysticetus c a n be r u l e d out, as that species h a s a t r u l y arctic a n d
subarctic d i s t r i b u t i o n a n d has never o c c u r r e d i n the N o r t h Sea. T h e h u n t f o r G r e e n ­
l a n d r i g h t w h a l e s started o n l y i n the late 16th century, so the earlier m a t e r i a l cannot
h a v e o r i g i n a t e d f r o m that source. T h e e m e r g i n g p i c t u r e is that there has b e e n a p o p u ­
l a t i o n of n o r t h e r n r i g h t w h a l e s i n the N o r t h Sea at least t i l l the M i d d l e A g e s ; o u r f i n d s
f r o m T i e l fit i n w i t h this. W e d o n o t k n o w w h e t h e r this p o p u l a t i o n w a s a l r e a d y o v e r ­
e x p l o i t e d b y early w h a l e r s , or w h e t h e r there h a v e b e e n other causes f o r its d e c l i n e . A t
present, the n o r t h e r n r i g h t w h a l e is at most a n e x t r e m e l y rare v a g r a n t i n the N o r t h
Sea.
References
Aguilar, Α., 1986. A review of old Basque whaling and its effect on the right whales (Eubalaena
glacialis) of the North Atlantic. In: R.L. Brownell, Jr., P.B. Best & J.H. Prescott (eds). Right whales:
Past and present status.— Reports of the International Whaling Commission, Special Issue 10:
191­199.
Aguilar, Α., 1995. Eubalaena glacialis (Müller, 1776) ­ Nordkaper (auch Nördlicher Glattwal). I n: D.
Robineau, R. Duguy & M . Klima (eds). Handbuch der Säugetiere Europas. Band 6: Meeressäuger.
Teil I : Wale und Delphine ­ Cetácea. Teil I B: Ziphiidae, Kogiidae, Physeteridae, Balaenidae, Ba­
laenopteridae: 650­668.— Aula­Verlag, Wiesbaden.
Broch, H . , 1931. I ndomalayan Cirripedia. Papers from Dr. Th. Mortensen's Pacific expedition 1914­
1916.— Videnskabelige Meddelelser fra Dansk naturhistorisk Forening 91:1­146.
Brown, M.W., S.D. Kraus, D.E. Gaskin & B.N. White, 1994. Sexual composition and analysis of repro­
ductive females in the North Atlantic right whale, Eubalaena glacialis, population.— Marine Mam­
mal Science 10: 253­265.
Brown, S.G., 1986. Twentieth­century records of right whales (Eubalaena glacialis) in the Northeast
Atlantic Ocean. In: R.L. Brownell, Jr., P.B. Best & J.H. Prescott (eds). Right whales: Past and pre­
sent status.— Reports of the International Whaling Commission, Special I ssue 10:121­127.
Brownell, Jr., R.L., P.B. Best & J.H. Prescott (eds), 1986. Right whales: Past and present status. Pro­
ceedings of the workshop on the status of right whales. New England Aquarium, Boston, Massa­
chusetts 15­23 June 1983.— Reports of the International Whaling Commission, Special Issue 10:1­
289.
Chemnitz, J.H., 1785. Neues systematisches Conchylien­Cabinet 8: (i­xviii), 1­372, pis.— Raspe, Nürn­
berg.
Cummings, W.C., 1985. Right whales Eubalaena glacialis (Müller, 1776) and Eubalaena australis
(Desmoulins, 1822). In: S.H. Ridgway & R. Harrison (eds). Handbook of marine mammals. Vol­
ume 3. The sirenians and baleen whales: 275­304.— Academic Press, London/Orlando.
Darwin, C , 1854. The Balanidae, (or sessile cirripedes); the Verrucidae, etc., etc., etc. A monograph of
the sub­class Cirripedia, with figures of all the species: i­viii, 1­684, pis 1­30.— Ray Society, Lon­
don.
362
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
Engel, H., 1959. The echinoderms of Rumphius. In: H.C.D. de Wit (ed.). Rumphius Memorial Volume:
209-223, pis 23-26.— Hollandia, Baarn.
Eschricht, D.F. & J. Reinhardt, 1866. O n the Greenland right-whale (Balaena mysticetus, Linn.), with
especial reference to its geographical distribution and migrations in times past and present, and
to its external and internal characteristics. In: W . H . Flower (ed.). Recent memoirs on the Cetácea
by professors Eschricht, Reinhardt and Lilljeborg: 1-150.— Ray Society, London. (Translation of
an article in Danish, published in 1861 by Eschricht & Reinhardt).
Evans, P.G.H. & G . M . Scanlan, 1989. Historical review of cetaceans in British and Irish waters: 1-95.—
U K Cetacean Group, c/o Department of Zoology, University of Oxford.
Gruvel, A , 1903. Révision des cirrhipèdes appartenenant à la collection du Muséum d'Histoire
Naturelle [pt. 2].— Nouvelles Archives du Muséum d'Histoire Naturelle Paris, (4) 5: 95-170, pis
1-4.
Guiler, E.R., 1956. Supplement to a list of the Crustacea of Tasmania.— Records of the Queen Victoria
Museum, Launceston, Tasmania (n. ser.) 5:1-8.
International Code of Zoological Nomenclature, third edition, 1985: i-xx, 1-338.— International Trust
for Zoological Nomenclature, London.
Klinowska, M., 1991. Dolphins, porpoises and whales of the world. The IUCN red data book: i-viii, 1429.— International Union for the Conservation of Nature and Natural Resources, Gland/Cambridge.
Kompanje, E.J.O. & C. Smeenk, 1996. Recent bones of right whales Eubalaena glacialis from the southern North Sea.— Lutra 39: 66-75.
Kraus, S.D., 1997. North Atlantic right whales: Can we stop extinction? In: P.G.H. Evans (ed.). European Research on Cetaceans - 10.— Proceedings of the tenth annual conference of the European
Cetacean Society, Lisbon, Portugal 11-13 March 1996: 9.
Martens, E. von, 1902. Die Mollusken (Conchylien) und die übrigen wirbellosen Thiere im
Rumpf'schen Raritätkammer. In: M . Greshoff (ed.). Rumphius gedenkboek 1702-1902: 109-136.—
Koloniaal Museum, Haarlem.
Maul, G.E. & D.E. Sergeant, 1977. New cetacean records from Madeira.— Bocagiana 43:1-8.
Mol, Th., 1962. Afbeeldingen van walvisachtigen (Cetacea) in een catalogus van een oud kabinet in
Brugge.— Lutra 4: 21-22, pis II-III.
Mörch, O.A.L., 1853. Acephala. Annulata. Cirripedia. Echinodermata. Catalogus Conchyliorum quae
reliquit D. Alphonso d'Aguirra & Gadea Comes de Yoldi: 1-74.— L. Klein, Copenhagen.
Newman, W . A . & A . Ross, 1976. Revision of the balanomorph barnacles; including a catalog of the
species.— Memoir San Diego Society of Natural History 9:1-108.
Nilsson-Cantell, C.A., 1931. Revision der Sammlung recenter Cirripedien des Naturhistorischen
Museums in Basel.— Verhandlungen der Naturforschenden Gesellschaft in Basel 42:103-137, pl.
2.
Pastorino, G . & M . Griffin, 1996. A n extant whale barnacle (Cirripedia, Coronulidae) from Holocene
deposits of Buenos Aires (Argentina).— Crustaceana 69: 769-772.
Pilsbry, H.A., 1916. The sessile barnacles (Cirripedia) contained in the collections of the U.S. National
Museum; including a monograph of the American species.— Bulletin of the United States
National Museum 93: i-xi, 1-366, pis 1-76.
Ranzani, C , 1818. Osservazioni su i Balanidi.— Opuscoli Scientifici 2 (2): 63-93.
Reeves, R.R. & E. Mitchell, 1986a. The Long Island, New York, right whale fishery: 1650-1924. In: R.L.
Brownell, Jr., P.B. Best & J.H. Prescott (eds). Right whales: Past and present status.— Reports of
the International Whaling Commission, Special Issue 10: 201-220.
Reeves, R.R. & E. Mitchell, 1986b. American pelagic whaling for right whales in the North Atlantic. In:
R.L. Brownell, Jr., P.B. Best & J.H. Prescott (eds). Right whales: Past and present status.— Reports
of the International Whaling Commission, Special Issue 10: 221-259.
Rumphius, G.E., 1705. D'Amboinsche Rariteitkamer, behelzende eene beschryvinge van allerhande
zoo weeke als harde schaalvisschen, te weeten raare krabben, kreeften, en diergelyke zeedieren,
als mede allerhande hoorntjes en schulpen, die men in d'Amboinsche zee vindt: daar beneven
zommige mineraalen, gesteenten, en soorten van aarde, die in d'Amboinsche, en zommige
Holthuis et al. The find of a whale barnacle, Cetopirus complanatus. Zool. Verh. Leiden 323 (1998)
363
omleggende eilanden gevonden worden: 1­340, pis 1­60.— F. Halma, Amsterdam.
Sarfatij, H . , 1997. Tiel in succession to Dorestad. I n: G . de Boe & F. Verhaeghe (eds). Exchange and
trade in Medieval Europe.— Papers of the 'Medieval Europe Brugge 1997' conference, Volume 3:
193­201.
Scarff, J.E., 1986a. Occurrence of the barnacles Coronula diadema, C. reginae and Cetopirus complanatus
(Cirripedia) on right whales.— The Scientific Reports of the Whales Research I nstitute 37: 129­
153.
Scarff, J.E., 1986b. Historic and present distribution of the right whale (Eubalaena glacialis) in the east­
ern North Pacific south of 50° Ν and east of 180° W. In: R.L. Brownell, Jr., P.B. Best & J.H. Prescott
(eds). Right whales: Past and present status.— Reports of the I nternational Whaling Commission,
Special Issue 10: 43­63.
Smet, W . M . A . De, 1974. I nventaris van de walvisachtigen (Cetácea) van de Vlaamse kust en de
Schelde.— Bulletin Koninklijk Belgisch I nstituut voor Natuurwetenschappen, Biologie 50 (1): 1­
156, pis I­IV.
Smet, W . M . A . De, 1976. Evidences of whaling in the North Sea and the English Channel during the
Middle Ages.— F A O Advisory Committee on Marine Resources, Scientific Consultation on
Marine Mammals Bergen, Norway, 31 August ­ 9 September 1976. ACMRR/MM/SC/102:1­8.
Smet, W . M . A . De, 1981. Gegevens over de walvisachtigen (Cetácea) van de Vlaamse kust en de
Schelde uit de periode 1969­1975.— Bulletin Koninklijk Belgisch I nstituut voor Natuurweten­
schappen, Biologie 53 (4): 1­34.
Stebbing, T.R.R., 1910. General catalogue of South African Crustacea (Part V of S.A. Crustacea, for the
Marine Investigations in South Africa).— Annals of the South African Museum 8: 281­593, pis 15­
22.
Stubbings, H.G., 1967. The cirriped fauna of tropical West Africa.— Bulletin of the British Museum
(Natural History), Zoology 15: 229­319, pi. 1.
Tarasov, N.J. & G.B. Zevina, 1957. Usonogie raki (Cirripedia thoracica) morei SSSR.— Fauna SSSR (η.
ser.) 69 (= Crust. 6 no. 1): 1­268, pis 1­3,1 col. frontisp.
Young, P.S., 1991. The superfamily Coronuloidea Leach (Cirripedia, Balanomorpha) from the Brazil­
ian coast, with redescription of Stomatolepas species.— Crustaceana 61:190­212.