HELGOL.~NDER MEERESUNTERSUCHUNGEN Helgol~inder Meeresunters. 52, 197-207 (1998)
T w e l v e i n v e r t e b r a t e a n d e i g h t fish s p e c i e s n e w
to t h e m a r i n e f a u n a of M a d e i r a , a n d a d i s c u s s i o n
of t h e z o o g e o g r a p h y of t h e a r e a
P e t e r Wirtz
Centro de Ci~ncias Biol6gicas e Geol6gicas, Universidade da Madeira, Largo do Col~gio, P - 9000 Funchal, Portugal, Madeira
ABSTRACT: The benthic ctenophore Vallicula multiformis, a large undescribed flatworm species of the genus Pseudoceros, the prosobranch gastropod Tonna maculosa, the opisthobranch gastropods Placida cf. dendritica, Calona elegans, Aeolidiella sanguinea, Janolus cristatus, the decapod Balssia gasti, the sea urchin Schizaster canaliferus and the tunicates Cla velina lepa diformis, Cla velina della- vallei and Pycnoclavella taureanensis are recorded from Madeira for the first time. This is the first record of a platyctenid ctenophore in the eastern Atlantic. The teleost fishes Pomatoschistus pictus, Vaneaugobius canariensis, Chromogobius sp., Nerophis ophidion, Hippocampus hippocampus, Acanthocybium solandri, Sphyraena viridensis and Sphyraena barracuda are recorded from Ma- deira for the first time. The presence of the sea-hare Aplysia dactylomela at Madeira is confirmed; the species has increased tremendously in abundance in the last four years. The crocodile fish Gram- moplites gruveli can occasionally be found in the mantle cavity of cuttlefish (Sepia officinalis) sold at the fish market of Funchal, but does not originate from Madeiran waters. An analysis of 100 new records from the coastal fauna of Madeira shows that, while predominantly of lusitanian, medi- terranean and mauritanian affinity, Madeira's shallow water fauna contains a large component of tropical species.
I N T R O D U C T I O N
D u r i n g a n o n g o i n g s u r v e y of t h e l a r g e r m a r i n e v e r t e b r a t e s a n d i n v e r t e b r a t e s of t h e c o a s t s of M a d e i r a {Biscoito & Wirtz, 1994; Wirtz, 1994, 1995a, 1995b; W i t t m a n n & Wirtz, in press), s e v e r a l s p e c i e s w e r e e n c o u n t e r e d t h a t a p p a r e n t l y h a d n o t y e t b e e n r e c o r d e d f r o m M a d e i r a . T w e n t y of t h e s e a r e d e s c r i b e d in this p a p e r . A l a r g e n u m b e r of H y d r o z o a , N u d i b r a n c h i a a n d D e c a p o d a , all n e w for t h e m a r i n e f a u n a of M a d e i r a a n d i n c l u d i n g s e v e r a l u n d e s c r i b e d s p e c i e s , w i l l b e t r e a t e d in s e p a r a t e p u b l i c a t i o n s b y e x p e r t s for t h e s e g r o u p s .
M A T E R I A L A N D M E T H O D S
T h e s u r v e y is p e r f o r m e d b y S C U B A - d i v i n g a n d t h e r e f o r e l i m i t e d to a d e p t h r a n g e d o w n to a b o u t 60 m. A n i m a l s a r e p h o t o g r a p h e d in t h e field. W h e n it a p p e a r s n e c e s s a r y , s p e c i m e n s a r e t h e n c o l l e c t e d for l a t e r i d e n t i f i c a t i o n . In m o s t c a s e s , p r e s e r v e d s p e c i m e n s a r e s e n t to s p e c i a l i s t s for i d e n t i f i c a t i o n or c o n f i r m a t i o n of i d e n t i f i c a t i o n (see " A c k n o w - l e d g e m e n t s " ) .
198 R Wirtz
nicipal (Hist6ria N a t u r a l ) of F u n c h a l , e x c e p t for the following: the s i n g l e s p e c i m e n of Aeolidiella sanguinea is in the collection of the Z o o l o g y L a b o r a t o r y of t h e U n i v e r s i t y of Oviedo, Spain; Hippocampus hippocampus a n d Sphyraena viridensis w e r e d o c u m e n t e d b y u n d e r w a t e r p h o t o g r a p h y (slides d e p o s i t e d in t h e M u s e u M u n i c i p a l of Funchal); Acanthocybium solandri w a s d o c u m e n t e d b y n u m e r o u s p h o t o s of t h e fish p l u s t h e p e r s o n h a v i n g c a p t u r e d it (one s u c h p h o t o w a s d e p o s i t e d in the M u s e u M u n i c i p a l of Funchal). A p p a r e n t l y , n e i t h e r p h o t o s nor p r e s e r v e d s p e c i m e n s exist, as yet, of Sphyraena barracuda from M a d e i r a n w a t e r s .
RESULTS
C t e n o p h o r a P l a t y c t e n i d a
Vallicula multiformis Rankin, 1956
A c r e e p i n g c t e n o p h o r e c a n c o m m o n l y b e e n c o u n t e r e d at n i g h t on a l g a e such as Zonaria tourne[ortii in d e p t h s of 5 to 15 m d u r i n g the s u m m e r months. T h e s p e c i e s ob- viously b e l o n g s to the g e n u s Vallicula R a n k i n 1956. Vallicula multiformis is the only k n o w n s p e c i e s in this g e n u s a n d the s p e c i m e n s from M a d e i r a p e r f e c t l y c o r r e s p o n d to the d e s c r i p t i o n of the s p e c i e s as g i v e n b y R a n k i n (1956).
A n i m a l s w e r e s e e n c l i n g i n g to the tips of the a l g a e only u n d e r c o n d i t i o n s of ( m o d e - rate) w a t e r m o v e m e n t . V. multiforrnis p r o b a b l y n e e d s w a t e r flow for p a s s i v e e x t e n s i o n of its t e n t a c l e s (Emson & Whitfield, 1991). V o u c h e r s p e c i m e n s h a v e b e e n d e p o s i t e d not only in the M u s e u M u n i c i p a l (Hist6ria N a t u r a l ) in F u n c h a l but also in the N a t i o n a a l N a t u u r - historisch M u s e u m in L e i d e n a n d t h e N a t u r a l History M u s e u m in London.
T h e s p e c i e s h a s so far only b e e n r e c o r d e d from the W e s t e r n Atlantic ( F r e e m a n , 1967; Rankin, 1956; Sterrer, 1986). T h e f i n d i n g of V. multiformis in M a d e i r a n w a t e r s r e p r e s e n t s the first r e c o r d of a p l a t y c t e n i d c t e n o p h o r e from the e a s t e r n Atlantic.
P l a t h e l m i n t h e s P o l y c l a d i d a
Pseudoceros sp.
A l a r g e , colourful, a n d a p p a r e n t l y u n d e s c r i b e d f l a t w o r m of the g e n u s Pseudoceros c a n o c c a s i o n a l l y b e f o u n d w h e n t u r n i n g o v e r stones in a d e p t h r a n g e of (at least) 10 to 20 m a l o n g t h e s o u t h c o a s t of M a d e i r a . I n d i v i d u a l s u s u a l l y are t h r e e to six c m long. A p p a r e n t l y , no o t h e r s p e c i e s in t h e g e n u s Pseudoceros h a s the colour p a t t e r n of g o l d e n a n d w h i t e lines on a p i t c h b l a c k body. A colour p h o t o of a n i n d i v i d u a l of this s p e c i e s c o l l e c t e d n e a r Cani~o, s o u t h coast of M a d e i r a , in 12 m d e p t h is g i v e n in Wirtz (1995a, p a g e 69 bottom).
T h e s p e c i e s also occurs in the C a n a r y Islands: a c o l o u r p h o t o of the s a m e s p e c i e s is g i v e n in P~rez S 6 n c h e z & M o r e n o Batet (1991, p a g e 101).
M o l l u s c a G a s t r o p o d a
Tonna maculosa (Dillwyn, 1810)
Twelve i n v e r t e b r a t e a n d eight fish species n e w to the m a r i n e f a u n a of M a d e i r a 199 borehole n e a r the tip, otherwise the shell is in perfect condition. While the f i n d i n g of a m o l l u s c a n shell c a n n o t b e definitive proof of the existence of hve a n i m a l s i n the area, it seems u n h k e l y t h a t a n y b o d y w o u l d h a v e t r a n s p o r t e d the shell to M a d e i r a a n d t h e n dis- c a r d e d it.
T h e species lives in the W e s t e r n Atlantic ( C a r i b b e a n to Bermuda) a n d has occasion- ally b e e n f o u n d a l o n g the west African coast a n d in the C a n a r y Islands (Sterrer, 1986; Poppe & Goto, 1991}.
Placida cf. dendritica (Alder & Hancock, 1843}
T h e species was c o m m o n on the g r e e n alga Codium decorticatum in a b o u t 2 - 6 m d e p t h in the shallow a n d very p r o t e c t e d b a y of Reis Magos, s o u t h e a s t e r n coast of Madeira, b e g i n n i n g of M a y 1995. N i n e w e e k s later most of the C. decorticatum h a d dis- a p p e a r e d a n d no P1acida could be f o u n d a n y more. In M a y a n d J u n e 1996, the species was a g a i n c o m m o n o n C. decorticatum at Reis Magos a n d $8o Pedro, s o u t h e a s t e r n coast of Madeira. In M a y 1996 it was also f o u n d to b e c o m m o n on Codium tomentosum at Porto Novo a n d in A u g u s t 1996 on Codium adhaerens at Caniqo.
Placida dendritica is a c o s m o p o l i t a n species living on several species of the g r e e n alga Codium; in the e a s t e r n Atlantic it has previously b e e n recorded from N o r w a y to the M e d i t e r r a n e a n (Millen, 1980; Gosliner, 1987; T h o m p s o n , 1988; Trowbridge, 1995}.
Caloria elegans (Alder & Hancock, 1845)
Large n u m b e r s of this species w e r e s e e n c r a w l i n g over rocks in a b o u t 9 m d e p t h at Porto da Cruz, north coast of Madeira, e n d of March a n d e n d of April 1996; m i d d l e of May, no more a n i m a l s w e r e s e e n in the s a m e area.
Caloria elegans is c o n s i d e r e d a b o r e a l species (Cattaneo-Vietti et ai., 1990) a n d has previously b e e n recorded from G r e a t Britain into the M e d i t e r r a n e a n (Thompson, 1988). T h e colour photo in Wirtz (1995a, p a g e 183) shows a n i n d i v i d u a l from Faial Island, Azores.
Aeolidiella sanguinea N o r m a n , 1877
A single i n d i v i d u a l of this species was f o u n d w h e n t u r n i n g over stones at a d e p t h of 9 m at Canigal, s o u t h e a s t e r n coast of Madeira. This is the a n i m a l figured in a colour photo on p a g e 183 in Wirtz (1995a).
Aeolidiefla sanguinea is a b o r e a l species, c o m m o n a r o u n d the British Isles a n d pre- viously r e c o r d e d as far south as the Strait of G i b r a l t a r ( G a r c i a - G o m e z et al., 1989). Fin- d i n g the species at M a d e i r a r e p r e s e n t s a c o n s i d e r a b l e r a n g e extension.
Janolus cristatus (Delle Chiaje, 1841)
Two adult a n i m a l s of this species w e r e s e e n close together on a rock i n a b o u t 9 m d e p t h at Porto da Cruz, n o r t h coast of Madeira, e n d of April 1996.
T h e species h a s p r e v i o u s l y b e e n r e c o r d e d from the M e d i t e r r a n e a n S e a a n d from south N o r w a y to Morocco i n the e a s t e r n Atlantic (Thompson, 1988; C a t t a n e o - V i e t t i et al., 1990).
Aplysia dactylomela Rang, 1828
200 P. Wirtz
Since then, the species has n e v e r a g a i n b e e n m e n t i o n e d from Madeira. D u r i n g several h u n d r e d dives a r o u n d the i s l a n d from 1983 until 1993, the a u t h o r n e v e r e n c o u n t e r e d this large sea-hare.
Aplysia dactylomela
was definitely a b s e n t from the area n e a r Ponta Oli- veira, south coast of Madeira, d u r i n g the years 1989 to 1993. T h e species w a s first recor- d e d in shallow w a t e r (5-10 m depth) n e a r Ponta Oliveira in 1994 a n d is n o w c o m m o n there. A n a n i m a l of a b o u t 20 cm l e n g t h was o b s e r v e d s p a w n i n g in a b o u t 4 m w a t e r d e p t h n e a r Reis Magos, south coast of M a d e i r a i s l a n d on 30 M a y 1997.C r u s t a c e a D e c a p o d a
Batssia gasti
(Balss, I921)Balssia gasti
is c o m m o n on the z o a n t h a r i a nCerardia savaglia,
that f r e q u e n t l y grows onAntipathes
b u s h e s a n d on old pieces ofAntipathes
skeletons on the south coast of Madeira. W h e n w i p i n g b u s h e s ofGerardia
with a small h a n d n e t at d e p t h s g r e a t e r t h a n 25 m (but n o t w h e n s e a r c h i n g shallower), this d e c a p o d w a s f r e q u e n t l y e n c o u n t e r e d . Noel (1992) r e c o g n i z e s two different E u r o p e a n species in the g e n u sBalssia;
a c c o r d i n g to his key, M a d e i r a n s p e c i m e n s b e l o n g to the speciesB. gasti
as they h a v e t h r e e l a r g e spines o n the cephalothorax. Voucher s p e c i m e n s h a v e b e e n d e p o s i t e d not only in the M u s e u M u n i c i p a l (Hist6ria Natural) at F u n c h a t b u t also in the N a t i o n a a l N a t u r r h i s t o r i s c h Mu- s e u m at Leiden.Balssia
spp. live associated with various a n t h o z o a n s a n d s p o n g e s in t h e M e d i t e r r a - n e a n sea a n d in the E a s t e r n Atlantic from n o r t h - e a s t e r n Morocco a n d theCanary
Islands to G u i n e a (Zibrowius, 1985; Noel, 1992; Mori et al., 1994; Gonz~ilez P~rez, 1995; O c a n a et al., 1995).E c h i n o d e r m a t a E c h i n o i d e a
Schizaster canaliferus
(Lamarck, 1816)A n e m p t y test of
Scl'n'zaster canaliferus
w a s f o u n d in a b o u t 12 m d e p t h on s a n d y bottom in front of Ponta S a n t a Caterina, s o u t h e a s t e r n coast of M a d e i r a in M a y 1996. T h e species has up to n o w b e e n c o n s i d e r e d a M e d i t e r r a n e a n e n d e m i c (Schinner, 1993).T u n i c a t a A s c i d i a c e a
Clavelina lepadiformis
(M(iller, 1776)Clavelina lepadiformis
(in its Wpical w h i t e - r i m m e d form) c a n o c c a s i o n a l l y be f o u n d in the a r e a a r o u n d Funchal. T h e species is n o t at all c o m m o n .C. lepadiformis
has pro- b a b l y b e e n t r a n s p o r t e d to the Azores by boats (Monniot, 1974; M o n n i o t & M o n n i o t , 1983) w h e r e it has s u b s e q u e n t l y s p r e a d (Wirtz & Martins, 1993).C. lepadiformis
m a y also h a v e b e e n t r a n s p o r t e d to M a d e i r a b y boats. T h e photos ofC. lepadiformis in
Wirtz (1995a, p a g e 221) a n d Wirtz (1996a, p a g e 162/163) are from A z o r e a n animals.Twelve i n v e r t e b r a t e a n d eight fish species n e w to the m a r i n e f a u n a of M a d e i r a 201 mals are restricted to d a r k places such as cave e n t r a n c e s a n d overhangs. It is, of course, also possible, that the yellow form is the i n d i g e n o u s one, while the white form has b e e n i n t r o d u c e d fairly recently.
C. lepadiIormis
has b e e n recorded from the M e d i t e r r a n e a n Sea a n d in the e a s t e r n Atlantic from w e s t e r n N o r w a y as far south as the C a n a r y Islands (Millar 1966; P~rez S~inchez & M o r e n o Batet, 1991; Wirtz & Martins, 1993).Clavelina dellavallei
(Zirpolo, 1925)This species is c o m m o n from low w a t e r s p r i n g tide level d o w n to at least 6 m d e p t h all a r o u n d Madeira. Photos of M a d e i r a n a n i m a l s are g i v e n i n Wirtz (1995a, p a g e 223, photo by P. Gomes) a n d in Wirtz (1996a, p a g e 164).
T h e species has so far b e e n c o n s i d e r e d a M e d i t e r r a n e a n e n d e m i c (Brunetti, pets. comm.) b u t T e m p l a d o et al. (1993) h a v e a l r e a d y r e p o r t e d it from the e a s t e r n Atlantic just outside the Strait of Gibraltar.
Pycnodavella taureanensis
Brunetti, 1991This species is very c o m m o n at Madeira, d o w n to at least 45 m depth. Photos of Ma- d e i r a n a n i m a l s are g i v e n in Wirtz (1995a, p a g e 221) a n d in Wirtz (1996a, p a g e 164).
Recently described, the species has so far only b e e n reported from the w e s t e r n Me- d i t e r r a n e a n .
O s t e i c h t h y e s T e l e o s t e i
Pomatoschistus pictus
(Malm, 1865)T h e p a i n t e d goby is c o m m o n on the south coasts of M a d e i r a a n d of Porto Santo, w h e r e it occurs on s a n d y bottom, f r e q u e n t l y close to rock walls, at depths from 10 to at least 40 m. Photos of M a d e i r a n a n i m a l s are g i v e n in Wirtz (1996b, page 238).
T h e species has b e e n recorded from the M e d i t e r r a n e a n Sea a n d in the e a s t e r n At- lantic from N o r w a y to the C a n a r y Islands (Miller, 1990).
Vaneaugobius cana~ensis
V a n Tassel, Miller & Brito, 1987A p o p u l a t i o n of this species was e n c o u n t e r e d on s a n d y - m u d d y bottom in a b o u t 10-20 m d e p t h o n the e a s t e r n side of the b a y of Machico, south coast of Madeira. Photos of M a d e i r a n a n i m a l s are g i v e n in Wirtz (1996b, p a g e 239). S p e c i m e n s h a v e b e e n de- posited in the M u s e u M u n i c i p a l (Hist6ria Natural) F u n c h a l a n d in the N a t u r k u n d e - m u s e u m Stuttgart.
T h e species has previously b e e n r e c o r d e d from the C a n a r y Islands a n d from G u i n e a (Miner, 1990).
Chromogobius
sp.202 E Wirtz
species is the s a m e as the u n d e s c r i b e d C a n a r i a n species. Photos of M a d e i r a n a n i m a l s are g i v e n in Wirtz (1996b, p a g e 239).
T h e species has so far only b e e n recorded from the C a n a r y Islands (Miller, 1990, as
Chromogobius
sp.).Nerophis ophidion
(Linnaeus, 1758)A single i n d i v i d u a l of this species was c a u g h t in a
Cymodocea nodosa
m e a d o w in the w e s t e r n half of the b a y of Machico in a b o u t 12 m d e p t h in S e p t e m b e r 1995.T h e species has previously b e e n r e c o r d e d from the M e d i t e r r a n e a n Sea a n d in the e a s t e r n Atlantic from N o r w a y as far south as the C a n a r y Islands (Dawson, 1986; Brito, 1991).
Hippocampus hippocampus
(Linnaeus, 1758)Both E u r o p e a n species of seahorses,
Hippocampus hippocampus
(Linnaeus, 1758) a n dHippocampus ramulosus
Leach, 1814 are c o m m o n at Madeira, b u t o n l yH. ramulo-
sus has so far b e e n n o t e d in the literature. T h e photo of a M a d e i r a n a n i m a l i n Wirtz (1994, p a g e 49) does in fact showH. hippocampus
a n d notH. ramulosus,
as e r r o n e o u s l y stated.Hippocampus hippocampus
has b e e n recorded from the M e d i t e r r a n e a n Sea a n d in the e a s t e r n Atlantic from the W a d d e n Sea south to the Gulf of G u i n e a (Dawson, 1986).A canthocybium solandn
(Cuvier, 1831)T h e Wahoo is occasionally sold at F u n c h a l fish m a r k e t ("Mercado dos Lavradores"). E n q u i r i n g a b o u t the origin of these fish, I w a s told that they came from M a d e i r a n waters. To check on this, I i n t e r v i e w e d the s p o r t s f i s h e r m a n c o m m u n i t y a n d w a s told the fol- lowing: Wahoo are not u n c o m m o n in late s u m m e r ; it seems that these fish m i g r a t e from the south with w a r m e r water, as in some years they h a r d l y a p p e a r at all, p a r t i c u l a r l y w h e n the water is u n s e a s o n a b l y cold (R. H a y e s a n d M. H e n r i q u e z , pets. comm.). N u m e r o u s photos d o c u m e n t the c a p t u r e of this species. Nevertheless,
Acanthocybium solandri
has not yet b e e n recorded from M a d e i r a i n the scientific literature a n d the FAO species c a t a l o g u e s u m m a r i z i n g our k n o w l e d g e on the scombrids of the world (Collette & N a u e n , 1983) notes the p r e s e n c e of this species at the C a p e Verde Islands, b u t n o t at the C a n a r y Islands, w h e r e it is of c o n s i d e r a b l e e c o n o m i c i m p o r t a n c e . T h e species h a s r e c e n t l y also b e e n recorded from the Azores (Azevedo & Heemstra, 1995).Sphyraena viridensis
(Cuvier, 1829)T h e only b a r r a c u d a reported from M a d e i r a n waters until n o w is t h e E u r o p e a n Bar- r a c u d a
Sphyraena sphyraena
(Linnaeus, 1758). As noted by Ben-Tuvia (1986) and de Silva (1990) the Striped BarracudaSphyraena vindensis
has in the past frequently been confused withSphyraena sphyraena.
Brito (1991) already noted thatS. sphyraena
pro- bably does not occur at the Canary Islands and that all previous records of this species at the Canaries are probably misidentifications of the very c o m m o nS. viridensis.
This is apparently also true for the Azores (Wirtz, 1990: unpublished report to the Departamento de Oceanografia e Pescas of the University of the Azores) and for Madeira: During several hundred dives at both these archipelagos I have seen and photographed S. dr/-densis
quite frequently, while never encounteringS. sphyraena. The
specimen reportedas
Sphyraena sphyraena
in Edwards (1993) was checked and confirmed to beS. viriden-
T w e l v e i n v e r t e b r a t e a n d e i g h t fish s p e c i e s n e w to the m a r i n e f a u n a of M a d e i r a 203
Sphyraena viridensis
is also k n o w n from the M e d i t e r r a n e a n Sea, b u t b e c a u s e of p a s t confusion withS. sphyraena
the d i s t r i b u t i o n of this s p e c i e s is insufficiently d o c u m e n t e d (de Silva, 1990; Bizsel & Cihangir, 1996).Sphyraena barracuda
( W a l b a u m , 1792)T h e G r e a t e r B a r r a c u d a
Sphyraena barracuda
h a s a l a r g e d i s t r i b u t i o n in t h e Atlantic, t h e I n d i a n O c e a n , a n d t h e Pacific (de Silva, 1990). In the e a s t e r n Atlantic it is c o n s i d e r e d rare a n d k n o w n w i t h c e r t a i n t y only from S e n e g a l to N i g e r i a (de Silva, 1990). W h e n i n t e r v i e w i n g the s p o r t s f l s h e r m a n c o m m u n i t y in F u n c h a l h a r b o u r (see s e c t i o n onAcan-
thocybium solandri,
above), I w a s told that, w h i l e the G r e a t e r B a r r a c u d a is e x t r e m e l y rare, l a r g e i n d i v i d u a l s of this s p e c i e s h a v e b e e n c a u g h t n e a r M a d e i r a (R. H a y e s a n d M. H e n r i q u e z , pers. comm.). Unfortunately, t h e r e s e e m s to b e no p h o t o to d o c u m e n t this.Grammoplites gruveli
(Pellegrin, 1905)W h e n b u y i n g cuttlefish
(Sepia o[ficinalis)
at F u n c h a l fish m a r k e t ( " M e r c a d o dos L a v r a d o r e s " ) the a u t h o r e n c o u n t e r e d twice, in 1994 a n d in 1996, an i n d i v i d u a l of the crocodile fishGrammoplites gruveli
in the m a n t l e c a v i t y of the cuttlefish. T h e f i s h e r m e n selling t h e cuttlefish h a d , however, not c a u g h t it t h e m s e l v e s . T h e y h a d b o u g h t t h e c u t t l e - fish from a firm t h a t h a d b o a t s c a t c h i n g t h e m in the a r e a of G u i n e a - B i s s a o . Thus, G.gruveli
c a n n o t b e c o u n t e d as a M a d e i r a n fish.T h e s p e c i e s h a s so far b e e n r e c o r d e d from M a u r i t a n i a to A n g o l a (Knapp, 1990).
D I S C U S S I O N
E a c h n e w l y r e c o r d e d s p e c i e s can b e c o n s i d e r e d a test c a s e for the theories on the zoo- g e o g r a p h y of the a r e a in question. In r e c e n t p u b l i c a t i o n s (Wirtz et al., 1988; A b r e u , 1991; A b r e u , 1994; A b r e u & Biscoito, 1994; Biscoito & Wirtz, 1994; Guerreiro, 1994; H o u a r t & Abreu, 1994; Stock, 1994; Wirtz, 1994, 1995 a, 1995 b; K n i g h t - J o n e s & K n i g h t - J o n e s , 1995; M a l a q u i a s , 1996; Nfinez et al., 1995; N d n e z & Talavera, 1996; O r t e a et al., 1996; R o g e r s & Nash, 1996; d ' U d e k e m d ' A c o z , 1996; Vald6s et al., 1996; W i t t m a n n & Wirtz in press) a n d in this one, a total of 134 s p e c i e s has b e e n n e w l y r e c o r d e d for the s h a l l o w w a t e r m a - rine f a u n a of M a d e i r a a n d the n e i g h b o u r i n g i s l a n d of Porto Santo (several more, a p p a - r e n t l y u n d e s c r i b e d s p e c i e s h a v e b e e n f o u n d r e c e n t l y a n d w e r e sent to s p e c i a l i s t s for description). For 30 of t h e s e , I w a s u n a b l e to find p r e c i s e d a t a on their d i s t r i b u t i o n . F o u r s p e c i e s f o u n d e x c l u s i v e l y in or n e a r h a r b o u r s a n d t h u s p r o b a b l y t r a n s p o r t e d b y m a n are e x c l u d e d from t h e f o l l o w i n g analysis. Table 1 s u m m a r i z e s t h e k n o w n d i s t r i b u t i o n of the r e m a i n i n g 100 s p e c i e s , u s i n g t h e t e r m i n o l o g y of Briggs (1974).
204 P. W i r t z
Table 1. Z o o g e o g r a p h i c affinity of 100 a n i m a l species recently recorded from the shallow waters of Madeira
Currently only k n o w n from M a d e i r a 7
Currently only k n o w n from M a d e i r a a n d the C a n a r y Islands 9 Currently only k n o w n from Madeira, the C a n a r y Islands a n d the Azores 2
Cosmopolitan 11
Eastern Atlantic boreal 19
Lusitanian, M e d i t e r r a n e a n a n d M a u r i t a n i a n 24
Lusitanian, M e d i t e r r a n e a n , M a u r i t a n i a n a n d West African 9
Tropical a) e a s t e r n Atlantic 2
b) amphi-Atlantic 9
c) circumtropical 7
Tropical amphi-Atlantic a n d e a s t e r n Atlantic Lusitanian a n d e a s t e r n Atlantic Boreal 1
t h o s e e x t e n d i n g i n t o t h e w e s t e r n M e d i t e r r a n e a n ) is a l s o q u i t e s t r o n g . A d d i n g to t h i s a s u b s t a n t i a l c o m p o n e n t of c o s m o p o l i t a n s p e c i e s ( 1 1 % ; b u t s e e K n o w l t o n , 1 9 9 3 a n d M i y a & N i s h i d a , 1997 o n t h e p r o b a b i l i t y t h a t m a n y s o c a l l e d c o s m o p o l i t a n s p e c i e s a r e u n r e c - o g n i z e d c o m p l e x e s of s i b l i n g s p e c i e s ) , t h e p i c t u r e t h a t e m e r g e s is t h a t of a m u l t i p l e o r i g i n of t h e M a d e i r a n s h a l l o w w a t e r b i o t a .
B i a n c h i e t al. (in p r e s s ) h a v e p o i n t e d o u t t h a t t r o p i c a l s p e c i e s a r e t h e d o m i n a n t s p e c i e s i n t w o s u b l i t t o r a l c o m m u n i t i e s . T h e y a l s o d i s c u s s e d t h e p o s s i b i l i t y of a r e c e n t w a r m i n g t r e n d i n f l u e n c i n g a n d c h a n g i n g t h e s h a l l o w w a t e r f a u n a l a n d f l o r a l c o m - p o s i t i o n of M a d e i r a . T h e i n c r e a s e of t h e a b u n d a n c e of t h e c i r c u m t r o p i c a l s e a - h a r e Aplysia dactylomela o n t h e c o a s t of M a d e i r a , w h i c h u n d o u b t e d l y is a r e c e n t p h e n o m e - n o n , is a n o t h e r i n d i c a t o r t h a t t h e s h a l l o w w a t e r b i o t a of M a d e i r a m i g h t b e i n a p h a s e of c h a n g e .
T w e l v e i n v e r t e b r a t e a n d e i g h t f i s h s p e c i e s n e w to t h e m a r i n e f a u n a of M a d e i r a 205
L I T E R A T U R E C I T E D
Abreu, A. D., 1991. First record of Mitra z o n a t a Mayrrat, 1818 a n d Cancilla turtoni (Smith, 1890) (Gastropoda: Mitridae) from the archipelago of Madeira. - Bocagiana 142, 1-2.
Abreu, A. D., 1994. O n the p r e s e n c e of Mitrella p a l l a r y i (Dautzenberg, 1927) (Mollusca: Gastropoda) in the archipelago of Madeira. - Bocagiana 172, 1-4.
Abreu, A. D. & Biscoito, M. J., 1994. The family Cassidae (Moilusca: Gastropoda) in the archipelago of Madeira. - Boeagiana I64, 1-3.
Azevedo, J. M. & Heemstra, P. C., 1995. N e w records of marine fishes from the Azores. - Arquip6lago. Life Mar. Sci. 13 A, 1-10.
Ben-Tuvia, A., 1986. Sphyraenidae. In: Fishes of the North-eastern Atlantic and the Mediterranean. Ed. by P. J. P. Whitehead, M. L. Bauchot, J. C. Hureau, J. Nielsen & E. Tortonese. UNESCO, Paris, 1473 pp.
Biscoito, M. & Wirtz, P., 1994. Two n e w records of stingrays (Pisces Dasyatidae) from the archipelago of Madeira (NE Atlantic). - Bocagiana 169, 1-4.
Bizsel, K. & Cihangir, B., 1996. A n e w fish record from the Turkish seas; yellow-mouth barracuda (Sphyraenidae; S p h y r a e n a v i r i d e n s i s Cuvier, 1 8 2 9 . - Turkish J. Zool. 20, 357-359.
Briggs, J. C., 1974. Marine zoogeography. McGraw-Hill, N e w York, 475 pp.
Brito, A., 1991. C a t , l o g o de los peces de las islas Canarias. F. Lemus, La Laguna, 230 pp.
Brunetti, R., 1987. Species of C l a v e f i n a in the M e d i t e r r a n e a n Sea. - Annls Inst. oc~anogr., Paris 63, 101-118.
Brunetti, R., 1991. P y c n o c l a v e l l a t a u r e a n i n a n.sp. (Ascidiacea) from the M e d i t e r r a n e a n Sea. - Vie Milieu 4 1 , 2 4 5 - 2 4 8 .
Cattaneo-Vietti, R., Chemello, R. & Gianuzzi-Savelli, R., 1990. Atlas of M e d i t e r r a n e a n nudibranchs. Editrice La Conchiglia, Roma, 264 pp.
Collete, B. & Nauen, C. E., 1983. FAO species catalogue. Vol. 2. Scombrids of the world. - F A O Fish. Synops. 125(2), 1-137.
Dawson, C. E., 1986. Syngnatidae. In: Fishes of the North-eastern Atlantic and the Mediterranean. Ed. by P. J. P. Whitehead, M. L. Bauchot, J. C. Hureau, J. Nielsen & E. Tortonese. UNESCO, Pa- ris, 1473 pp.
Edwards, A. J., 1993. A small collection of fishes from Madeira Island deposited in the Hancock Mu- seum. - T r a n s . nat. Hist. Soc. Northumbria 56, 35-40.
Emson, R. J. & Whitfield, P. J., 1991. Behavioural a n d ultrastructural studies on the s e d e n t a r y platy- c t e n e a n c t e n o p h o r e Vallicula m u l t i f o r m i s . - Hydrobiologia 216/217, 27-33.
Freeman, G., 1967. Studies on r e g e n e r a t i o n in the creeping ctenophore, Vallicula m u l t i f o r m i s . - J. Morphol. 123, 71-83.
Garcia-Gomez, J. C., Cervera, J. L., Garcia, I=. J. & Cuadra, C. L. M., 1989. Resultados de la Cam- pafia Internacional de Biologia Marina "Ceuta 86": Moluscos Opistobranquios. - Boll. malac. 25, 125-138.
Gonz~ilez P~rez, J. A., 1995. Crust&ceos dec&podos d e las Islas Canarias. Publicaciones Turquesa, Santa Cruz d e Tenerife, 282 pp.
Gostiner, T., 1987. N u d i b r a n c h s of Southern Africa. Sea Challengers, Monterey, 136 pp.
Guerreiro, J., 1994. On bivalve molluscs of the island of Porto Santo (Madeira, Portugal). - Bocagi- ana 173, 1-4.
Gronow, N. & Hayward, P. J., 1991. O p i s t h o b r a n c h e s d e l'he Maurice, a v e c l a description de d e u x esp~ces nouvelles. - Revue fr. Aquariol. I8, 1-30.
Houart, R. & Abreu, A. D., 1994. The Muricidae (Gastropoda) from Madeira with the description of a n e w species of O c e n e b r a ( O c i n e b n n a J (Muricidae: Ocenebrinae). - Apex 9, 119-130.
Knapp, L. W., 1990. Platycephalidae. In: Check-list of the fishes of the eastern tropical Atlantic. Ed. by J. C. Qu~ro, J. C. Hureau, C. Karrer, A. Post & L. Saldanha. UNESCO, Lisbon, 1492 pp. Knight-Jones, P. & Knight-Jones, E. W., 1995. Spirorbidae (Polychaeta) from Madeira including a
n e w species a n d s u b g e n u s of Spirorbis. - Mitt. hamb. zool. Mus. Inst. 92, 89-101. Knowlton, N., 1993. Sibhng species in the sea. - A. Rev. EcoL Syst. 24, 189-216.
Malaquias, M. A. E., 1996. Opisthobranquios (MoUusca Gastropoda) do arquip~lago d a Madeira e ilhas Selvagens. Relat6rio de est~gio, Universidade da Algarve, Faro, 72 pp.
2 0 6 R W i r t z
Millar, R. H., 1966. T u n i c a t a - A s c i d i a c e a . M a r i n e i n v e r t e b r a t e s of S c a n d i n a v i a 1, 1-123.
Millen, S. V., 1980. R a n g e e x t e n s i o n s , n e w d i s t r i b u t i o n sites, a n d n o t e s o n t h e b i o l o g y of s a c o - g l o s s a n o p i s t h o b r a n c h s (Mollusca: G a s t r o p o d a ) in British C o l u m b i a . - C a n . J. Zool. 58, 1 2 0 7 - 1 2 0 8 .
Miller, P. J., 1990. G o b i i d a e . In: C h e c k - l i s t of t h e f i s h e s of t h e e a s t e r n tropical A t l a n t i c . Ed. by J. C. Q u ~ r o , J. C. H u r e a u , C. Karrer, A. P o s t & L. S a l d a n h a . U N E S C O , Lisbon, 1492 p p .
M o n n i o t , C. & M o n n i o t , F., 1983. N a v i g a t i o n o u c o u r a n t s ? La c o l o n i s a t i o n d e s A q o r e s et d e s Ber- m u d e s p a r les a s c i d i e s (Tuniciers b e n t h i q u e s ) . - C. r. Soc. Biogeogr. 59, 5 3 - 5 8 .
M o n n i o t , F., 1974. A s c i d i e s littorales e b a t h y a l e s r 4 c o l t 4 e s a u c o u r s d e la c a m p a g n e B i a c o r e s : A p l o u - s o b r a n c h e s . - Bull. M u s . n a t n . Hist. nat., P a r i s (3 e s&r. n ~ 251, Zoologic) 173, 1 2 8 7 - 1 3 2 6 . Mori, M., Morri, C. & Bianchi, C. N., 1994. N o t e s o n t h e life h i s t o r y of t h e p o n t o n i n e s h r i m p Balssia
gasti (Balss, 1921). - O e b a l i a 20, 1 2 9 - 1 3 7 .
N o b r e , A., 1937. M o l u s c o s T e s t ~ c e o s M a r i n h o s d o A r q u i p 4 1 a g o d a M a d e i r a . M e r e s E s t u d . M u s . zool. Univ. C o i m b r a (Ser. 1) 98, 1-101.
Noel, P., 1992. C14 p r 4 l i m i n a i r e d ' i d e n t i f i c a t i o n d e s C r u s t a c e a D e c a p o d a d e F r a n c e et d e s p r i n c i p a - les a u t r e s e s p S c e s d ' E u r o p e . - Coll. p a t r i m o i n e s nat. S e c r e t a r i a t d e la F a u n e et d e la Flore. M u s . Nat. d ' H i s t . N a t u r e l l e , Paris, 9, 1-145.
Ntinez, J., P a s c u a l , M., D e l g a d o , J. D. & S a n M a r t i n , G., 1995. Interstitial p o l y c h a e t e s f r o m M a d e i r a , w i t h a d e s c r i p t i o n of Syllides bansei P e r k i n s , 1981. - B o c a g i a n a 179, 1-7.
Nfinez, J. 8, T a l a v e r a , J. A., 1996. F a u n a of t h e p o l y c h a e t o u s A n n e l l d a f r o m M a d e i r a . - B a l m M u s . M u n . F u n c h a l (Supl.) 4, 5 1 1 - 5 3 0 .
O c a n a , O., Brito, A., N f i n e z , J. & B a c a l l a d o , J., 1995. R e d e s c r i p c i 6 n d e Gerardia savaglia (Bertolini, 1891) ( A n t h o z o a : Z o a n t h a r i a : G e r a r d i i d a e ) . - V i e r a e a 24, 153-164.
O r t e a , J. & M a r t i n e z , E., 1991. M o l l u s c o s O p i s t h o b r a n c h i o s d e C a b o Verde: A n a s p i d e a (Aplysio- m o r p h a ) . - Publ. O c a s . Soc. Port. M a l a c . 15, 17-42.
O r t e a , J., Vald~s, A. & G a r c i a - G d m e z , J. C., 1996. R e v i s i d n d e las e s p e c i e s a t l f i n t i c a s d e la f a m i l i a C h r o m o d o r i d i d a e (Mollusca: N u d i b r a n c h i a ) d e l g r u p o cromfitico azul. - A v i c e n n i a (supl.) I, 1-165.
P 6 r e z Sc~nchez, J. M. & M o r e n o Batet, E., 1991. I n v e r t e b r a d o s M a r i n o s d e C a n a r i a s . E d i c i o n e s d e l C a b i l d o I n s u l a r d e G r a n C a n a r i a , L a s P a l m a s , 335 pp.
P o p p e , G. T. & Goto, Y., 1991. E u r o p e a n s e a s h e l l s . H e m m e n , W i e s b a d e n 1, 1-352.
Q u i g n a r d , J. R & Pras, A., 1986. L a b r i d a e . In: F i s h e s of t h e N o r t h - e a s t e r n A t l a n t i c a n d t h e M e d i - t e r r a n e a n . Ed. b y R J. R W h i t e h e a d , M. L. B a u c h o t , J. C. H u r e a u , J. N i e l s e n & E. T o r t o n e s e . U N E S C O , Paris, 1473 pp.
R a n k i n , J. J., 1956. T h e s t r u c t u r e a n d b i o l o g y of Vallicula multiformis, g e n . et sp. nov. a p l a t y c t e n i d c t e n o p h o r e . - J. L m n . Soc. (Zool.) 43, 5 5 - 7 1 .
R o g e r s , A. D. & N a s h , R. D. M., 1996. A n e w s p e c i e s of Ochetostoma (Echiura, E c h i u r i d a e ) f o u n d in t h e A z o r e s , w i t h n o t e s o n its ecology. - J. m a r . biol. A s s . U. K. 76, 4 6 7 - 4 7 8 .
S c h i n n e r , G. O., 1993. B u r r o w i n g b e h a v i o u r , s u b s t r a t e p r e f e r e n c e a n d d i s t r i b u t i o n of Schizaster canaliferus ( E c h i n o i d e a S p a t a n g o i d a ) in t h e n o r t h e r n A d r i a t i c sea. - Mar. Ecol. 14, 129--145. Silva, D. R de, 1990. S p h y r a e n i d a e . In: C h e c k - l i s t of t h e f i s h e s of t h e e a s t e r n t r o p i c a l Atlantic. Ed.
b y J. C. Q u 4 r o , J. C. H u r e a u , C. Karrer, A. P o s t & L. S a l d a n h a . U N E S C O , L i s b o n , 1492 pp. Sterrer, W., 1986. M a r i n e f a u n a a n d flora of B e r m u d a . Wiley, N e w York, 742 p p .
Stock, J. H., 1994. A n e w m e m b e r of t h e f a m i l y B o g i d i e l l i d a e ( C r u s t a c e a , A m p h i p o d a ) f r o m poikilo- h a l i n e w a t e r s of M a d e i r a . - B o c a g i a n a I71, 1-8.
T e m p l a d o , J., G u e r r a , A., B e d o y a , J., M o r e n o , D., R e m 6 n , J. M. M a l d o n a d o , M. & R a m o s , M. A., 1993. F a u n a m a r i n a circalitoral d e l S u r d e la P e n i n s u l a Iberica. M u s e u N a c i o n a l d e C i e n c i a s N a t u r a l e s . M a d r i d , 181 p p .
T h o m p s o n , T. E., 1988. M o l l u s c s : B e n t h i c o p i s t h o b r a n c h s . Brill, L e i d e n , 356 p p . ( S y n o p s e s of t h e British F a u n a N.S. 8.).
T r o w b r i d g e , C. C., 1995. N e w Z e a l a n d o p i s t h o b r a n c h s a s s o c i a t e d w i t h t h e l o w i n t e r t i d a l , c r u s t o s e g r e e n a l g a Codium convolutum. - V e l l g e r 38, 1 1 6 - 1 2 5 .
T w e l v e i n v e r t e b r a t e a n d e i g h t f i s h s p e c i e s n e w to t h e m a r i n e f a u n a of M a d e i r a 2 0 7
Vald#s, A., O r t e a , J., Avila, C. & B a l l e s t e r o s , M., 1996. R e v i e w of t h e g e n u s Oendrodoris E h r e n b e r g , 1831 ( G a s t r o p o d a : N u d i b r a n c h i a ) in t h e A t l a n t i c o c e a n . - I . moll. Stud. 62, 1 - 3 1 .
W a t s o n , R. B., 1897. O n t h e m a r i n e m o U u s c a f r o m M a d e i r a ; w i t h d e s c r i p t i o n s of t h i r t y - f i v e n e w s p e - cies, a n d a n i n d e x - l i s t of all t h e k n o w n s e a - d w e l l i n g s p e c i e s of t h a t island. - I. Linn. Soc. (ZooL) 26, 18-320.
Wirtz, P., 1994. T h r e e s h r i m p s , five n u d i b r a n c h s , a n d two t u n i c a t e s n e w for t h e m a r i n e f a u n a of M a d e i r a . - B o l m M u s , M u n . F u n c h a I 46 (257), 167-172.
Wirtz, P,, 1995a. U n t e r w a s s e r f f t l i r e r M a d e i r a K a n a r e n A z o r e n W i r b e l l o s e - U n d e r w a t e r G u i d e M a d e i r a C a n a r y I s l a n d s A z o r e s I n v e r t e b r a t e s . N a g l s c h m i d , S t u t t g a r t , 248 pp.
Wirtz, P,, 1995b. O n e v a s c u l a r p l a n t a n d t e n i n v e r t e b r a t e s n e w for t h e m a n n e flora a n d f a u n a of M a d e i r a . - A r q u i p ~ I a g o . Life Mar. Sci. I3 A, 1 1 9 - I 2 4 .
Wirtz, P., 1996a. N e u e K e u l e n s e e s c h e i d e n vort M a d e i r a . - A q u a r . Terrar. Z. 1996, t 6 2 - t 6 4 . WLr~Z, P., 1996b. Die G r u n d e l n y o n M a d e i r a . - A q u a r . Terrar. Z. I996, 2 3 6 - 2 3 9 .
Wirtz, P. & M a r t i n s . H., 1993. N o t e s o n s o m e r a r e a n d h t t l e k n o w n m a r i n e i n v e r t e b r a t e s f r o m t h e A z o r e s - w i t h a d i s c u s s i o n of t h e z o o g e o g r a p h y of t h e a r e a . - A r q u i p e t a g o . Life Mar. Sei. 11 A, 5 5 - 6 3 .
Wirtz, P,, M~ller, B. & N a h k e , P., 1988. T h e C a r i b b e a n s h r i m p Tuleariocaris neglecta C h a c e 1969 f o u n d in a s s o c i a t i o n w i t h Diadema antillarum at M a d e i r a , a n d two n e w r e c o r d s of d e c a p o d c r u s t a c e a n s f r o m t h e C a p e V e r d e i s l a n d s . - Cour. Forsch. Inst. S e n c k e n b e r g I05, 169-171. W i t t m a n n , K. & Wirtz, P. A first i n v e n t o r y of t h e m y s i d f a u n a ( C r u s t a c e a : M y s i d a c e a ) in c o a s t a l
w a t e r s of t h e M a d e i r a a n d C a n a r y A r c h i p e l a g o s . - Bolm M u s . M u n i m p . F u n c h a l {in p r e s s ) . Z i b r o w i u s , H., 1985. Gerardia savaglia ( C n i d a n a : A n t h o z o a : Z o a n t h a r i a ) - n o u v e l h o t e d e Balssia
