Helgolander Meeresunters. 47, 125-143 (1993)
Deep-water macroalgae from the Canary Islands: new
records and biogeographical relationships
R.J. H a r o u n 1, W. F. P r u d ' h o m m e v a n R e i n e 2, D. G. M f i l l e r 3,
E. S e r r a o 4, R. H e r r e r a 1
1Dpto. de Biologfa, Universidad de Las Palmas de Gran Canaria; 35017 Las Palmas,
Spain
2 Pujksherbarium/Hortus Botanicus, University of Leiden; P.O. Box 9514, 2300 RA Leiden, The Netherlands
3Fakultft ffir Biologie der Universit5t Konstanz; Postfach 5560, D-78434 Konstanz, Federal Republic of Germany
4 U.C.T.R.A., Universidade de Algarve; Campus de Gambelas, 8000 Faro, Portugal
ABSTRACT: Due to the geographical location and paleobiogeography of the Canary Islands, the seaweed flora contains macroalgae with different distributional patterns. In this contribution, the biogeographical relations of several new records of deep-water macroalgae recently collected around the Canarian archipelago are discussed. These are Bryopsidella neglecta (Berthotd) Rietema, Discosporangium mesarthrocarpum (Meneghini) Hauck, Hincksia onslowensis (Amsler et Kapraun) P.C. Silva, Syringoderma floridana Henry, Peyssonnelia harveyana J. Agardh, Cryptonemia semi- nervis (C. Agardh) J. Agardh, Botryodadia wynnei Ballantine, Gloiocladia blomquistii (Searles) R. E. Norris, PIahchrysis peltata (W. R. Taylor) P. Huv4 et H. Huv4, Leptofauchea brasiliensis Joly, and Sarcodiotheca divaricata W. R. Taylor. These new records, especially those in the Florideophyceae, support the strong affinity of the Canary Islands seaweed flora with the warm-temperate Mediterra- nean-Atlantic region. Some species are recorded for the first time from the east coast of the Atlantic Ocean, enhancing the biogeographic relations of the Canarian marine flora with that of the western Atlantic regions.
I N T R O D U C T I O N
As a result of the g e o g r a p h i c a l location of the C a n a r y Islands, their s e a w e e d flora comprises species with different b i o g e o g r a p h i c a l distribution types in the Atlantic O c e a n : c o l d - t e m p e r a t e , w a r m - t e m p e r a t e M e d i t e r r a n e a n - A t l a n t i c a n d tropical (Feld- mann, 1946; Afonso-Carrillo & Gil Rodriguez, 1982; Lfining, 1990; P r u d ' h o m m e v a n Reine & v a n d e n H o e k , 1990). T h e s e b i o g e o g r a p h i c a l analyses are b a s e d on the floristic i n v e n t o r i e s of the b e n t h i c m a r i n e m a c r o a l g a e . T h e s e inventories, h o w e v e r , contain mainly supralittoral a n d eulittoral species, b e c a u s e of m o r e i n t e n s i v e studies of i n t e r t i d a l platforms. Thus, only f e w sublittoral m a c r o a l g a e h a v e b e e n r e p o r t e d for t h e C a n a r y Islands (Borgesen 1925-30; G i l - R o d r / g u e z & Afonso-Carrillo, 1980; Sans6n et al., 1991). Only sporadic e f f o r t s - h a v e b e e n m a d e to i n v e s t i g a t e the sublittoral flora, the first contributions b e i n g by Vickers (1897) a n d B o r g e s e n (1938). In r e c e n t years, t h e e x t e n s i v e collections arising from the C A N C A P e x p e d i t i o n s to the M a c a r o n e s i a n Islands h a v e
126 R . J . Haroun, W. F. P r u d ' h o m m e v a n Reine, D. G. M~iller, E. S e r r a o & R. H e r r e r a a d d e d significant n e w d a t a for the a r e a ( P r u d ' h o m m e v a n Reine, 1988; P r u d ' h o m m e v a n Reine & v a n d e n H o e k , 1988, 1990).
In g e n e r a l , d e e p - w a t e r b e n t h i c a l g a l c o m m u n i t i e s atl o v e r the w o r l d h a v e only o c c a s i o n a l l y b e e n s t u d i e d in d e t a i l b e c a u s e of m e t h o d o l o g i c a l difficulties e n c o u n t e r e d in t h e s a m p l i n g of such r e m o t e e n v i r o n m e n t s . Until now, collections for t h e s e s t u d i e s h a v e b e e n b a s e d on d r e d g i n g m e t h o d s or SCUBA d i v i n g (Hiscock & M a g g s , 1984; Kajimura, 1987; S e a r l e s & Schneider, 1987). Recently, R e m o t e O p e r a t e d Vehicles (ROV) p r o v i d e d with a v i d e o c a m e r a a n d / o r collecting e q u i p m e n t as w e l l as the a v a i l i a b i l i t y of r e s e a r c h s u b m e r s i b l e s h a v e e n h a n c e d studies on d e e p - w a t e r c o m m u n i t i e s (Littler et al., 1986; H a n i s a k & Blair, 1988). N e v e r t h e l e s s , d e e p - w a t e r m a c r o a l g a e a r e still n o t w e l l known.
T h e p u r p o s e of the p r e s e n t contribution is to r e p o r t on s e v e r a l n e w l y r e c o r d e d d e e p - w a t e r m a c r o a l g a e r e c e n t l y collected in t h e C a n a r i a n a r c h i p e l a g o a n d to discuss their b i o g e o g r a p h i c a l relations, w h i c h m a y f u r t h e r c o n t r i b u t e to a n u n d e r s t a n d i n g of its s e a w e e d flora.
MATERIAL A N D M E T H O D S
S a m p l i n g b y i n t e r t i d a l collecting, S C U B A d i v i n g a n d v i d e o - a i d e d d r e d g i n g w a s e x e c u t e d at 63 stations b e t w e e n S e p t e m b e r - O c t o b e r 1991 from t h e G e r m a n R.V. H e i n c k e (Fig. 1). All i s l a n d s of the a r c h i p e l a g o w e r e visited, b u t s a m p l i n g on n o r t h e r n coasts of s o m e i s l a n d s a n d on the coasts of G o m e r a I s l a n d w a s not p o s s i b l e b e c a u s e of a d v e r s e w e a t h e r conditions. A d d i t i o n a l s a m p l e s w e r e o b t a i n e d b y d r e d g i n g on a sea- m o u n t n e a r F u e r t e v e n t u r a I s l a n d (stations 53-55). T h e following a b b r e v i a t i o n s a r e u s e d from e a s t to w e s t for the C a n a r y Islands: L, Lanzarote; F, F u e r t e v e n t u r a ; C, G r a n Canaria; T, Tenerife; G, G o m e r a ; P, La P a l m a a n d H, Hierro.
T h e c o l l e c t e d m a t e r i a l w a s s o r t e d out on s h i p b o a r d , a n d i n t e r e s t i n g s p e c i m e n s , as w e l l as the crustose coraUine algae, w e r e p r e s e r v e d in 5 % F o r m a l i n i n s e a w a t e r a n d s t o r e d in the dark. M o s t s p e c i m e n s w e r e p r o v i s i o n a l l y i d e n t i f i e d on b o a r d u s i n g keys, d e s c r i p t i o n s a n d r e c o r d s from p u b l i s h e d p a p e r s on m a r i n e a l g a e for M a c a r o n e s i a a n d a d j a c e n t a r e a s (Borgesen, 1925-30; Taylor, 1960; G i l - R o d r i g u e z & A f o n s o - C a r r i l l o , 1980; L a w s o n & John, 1987; Afonso-Carrillo & Sans6n, 1989; S c h n e i d e r & S e a r l e s , 1991), a l t h o u g h m a n y a d d i t i o n a l r e f e r e n c e s w e r e u s e d as well. D a t a on d i s t r i b u t i o n of the c o l l e c t e d t a x a w e r e o b t a i n e d from the d a t a m a t r i x on M a c a r o n e s i a n m a r i n e a l g a e ( P r u d ' h o m m e v a n Reine, 1988; P r u d ' h o m m e v a n Reine & v a n d e n H o e k , 1988, 1990). S p e c i e s n a m e s are m a i n l y a r r a n g e d following the s y s t e m a t i c a p p r o a c h of W y n n e (1986). M o r e t h a n 1245 v o u c h e r s p e c i m e n s of 278 m a c r o a l g a l s p e c i e s w e r e p r e p a r e d a n d m o u n t e d ; v o u c h e r n u m b e r s c o r r e s p o n d to t h e R i j k s h e r b a r i u m L e i d e n c o l l e c t i o n n u m b e r s of the C A N C A P e x p e d i t i o n s . The m a i n v o u c h e r collection is k e p t in t h e Las P a l m a s d e G r a n C a n a r i a University H e r b a r i u m , D u p l i c a t e s of m o s t s p e c i e s a r e d e p o s i t e d in the R i j k s h e r b a r i u m of L e i d e n a n d s o m e in the A l g a r v e H e r b a r i u m . D e t a i l e d s t u d i e s on the c o l l e c t e d s p e c i m e n s w e r e u n d e r t a k e n in the l a b o r a t o r i e s in Las P a l m a s d e G r a n C a n a r i a a n d in Konstanz, G e r m a n y .
L a b o r a t o r y cultures w e r e u s e d to o b t a i n s m a l l e r t a x a not directly d e t e c t a b l e in field collections. T h e y w e r e i n i t i a t e d b y i s o l a t i n g m i l h m e t e r - s i z e d f r a g m e n t s of v a r i o u s s u b - strata, such as m a c r o a l g a e , shells, c o r a l l i n e algae, small p e b b l e s or s a n d g r a i n s into p o l y p r o p y l e n e t u b e s with 4-ml culture m e d i u m (sea w a t e r w i t h PES e n r i c h m e n t ) . U p o n arrival at the K o n s t a n z laboratory, the i n o c u l a n t s w e r e t r a n s f e r r e d to p o l y s t y r e n e Petri
~eee u2 N ,.-1 0 I.M 0 Z _..1 b-- 9 o o e~ o . ~ 6~
128 R . J . Haroun, W. F. P r u d ' h o m m e v a n Reine, D. G. Mfiller, E. Serrao & R. Herrera
dishes a n d i n c u b a t e d at 12 or 18 _ 1 ~ u n d e r white fluorescent light with p h o t o n fluence rates b e t w e e n 11 a n d 17 ~mol m -2 s -1. I n t e r e s t i n g taxa d e v e l o p i n g in s u c h r a w cultures w e r e m a d e u n i a l g a l b y isolation of c l e a n thallus fragments. For d o c u m e n t a t i o n , algal thalli w e r e fixed a n d s t a i n e d in acetocarmine, c o n t a i n i n g a trace a m o u n t of a n i l i n e blue, a n d m o u n t e d i n 50 % Karo | corn syrup.
RESULTS
C h l o r o p h y t a : B r y o p s i d a c e a e
Bryopsidella neglecta
(Berthold) RietemaC o 11 e c t i o n s : # 14282 a n d # 14288, Sept. 30, Stat. 14, 28o49'54" N, 13o51'32" W, Estrecho La Bocafna b e t w e e n L a n d F, d e p t h 65-70 m; # 14365, Sept. 30, Stat. 15, 28~ N, 13~ W, Estrecho La Bocaina b e t w e e n L a n d F, d e p t h 70-80 m; # 15119, Oct. I1, stat. 53, 28~ 14~ S e a m o u n t n e a r F, d e p t h 45-50 m; # 15250, Oct. 11, Stat. 56, 28~ 14~ Pta. Barlovento, F, d e p t h 64 m.
Small plants of a coenocytic g r e e n alga were collected as e p i p h y t e s on b l a d e s of various
Halymenia
species. T h e y were originally identified b y us asBryopsis halymeniae
Berthold. Rietema (1972) d e m o n s t r a t e d that this is a stage i n the life cycle ofDerbesia
neglecta
Berthold. T h e correct n a m e was later c h a n g e d toBryopsidella neglecta
(Rietema, 1975). All p l a n t s w e collected were sterile (Fig. 2). Later,B. neglecta
p l a n t swere also o b t a i n e d in culture from spores of field-collected
D. neglecta
s p e c i m e n s from the same d e p t h range.World distribution: M e d i t e r r a n e a n Sea a n d Atlantic coast of Morocco.
Notes on biogeography: These records r e p r e s e n t a n e x t e n s i o n of the k n o w n distribu- tion area of this species (Fig. 12).
B. neglecta
a p p e a r e d i n our samples from b e t w e e n 4 5 - 8 0 m d e p t h onHalymenia
blades; this is the d e e p e s t k n o w n record for this species in its distributional range. AsD. neglecta
it w a s reported for T (Borgesen, 1925), b u t as a tide pool plant.P h a e o p h y t a : E c t o c a r p a c e a e
Hincksia onslowensis
(Amsler et K a p r a u n ) P. C. SilvaC o 11 e c t i o n s : # 15248, Sept. 30, Star. 14, 28049'54 " N, 13~ W, Estrecho La Bocaina b e t w e e n L a n d F, d e p t h 65-70 m; # 15249, Oct. 8, Stat. 46, 28~ " N, 15o42'72 " W, Sardina del Norte, C, d e p t h 30-40 m.
Filamentous, uniseriate ectocarpoid thalli up to 5 cm long, were f o u n d e n t a n g l e d with other species. Basal filaments are sparsely b r a n c h e d , w h e r e a s apical filaments are profusely b r a n c h e d . Plurilocular sporangia, sessile or o n one- to several-celled pedicels, are laterally attached. Additional features such as cell d i m e n s i o n s a n d chloroplast m o r p h o l o g y allowed us to identify these s p e c i m e n s as
Hincksia onslowensis,
a species recently d e s c r i b e d from North Carolina d e e p - w a t e r flora (Amsler & K a p r a u n , 1985, asGiffordia onslowensis).
Plants were isolated from i n o c u l a n t s ofSyringoderma floridana
H e n r y a n d
Sporochnus bolleanus
M o n t a g n e fragments.In older cultures, plants formed a b u n d a n t lateral outgrowths of 2 - 5 cells in length, which e x t e n d at right angles a n d have a rhizoidal character. This feature a n d the p r e s e n c e of u n i l o c u l a r sporangia are s h a r e d with
Hincksia sordida
(Harvey) P.C. Silva, which is a related taxon reported from T a s m a n i a a n d s o u t h e r n Australia (Womersley, 1987, asGiffordia sordida).
A n isolate, collected in 1991 from the coast of Chile, shows i n t e r m e d i a t e characteristics, w e l l - d e v e l o p e d p s e u d o h a i r s a n d u n i l o c u l a r s p o r a n g i a (Miil- ler, unpubl.). A comparative culture study with isolates from the type locality of each taxa, as well as the C a n a r i a n a n d C h i l e a n populations, is n e e d e d to clarify the relationship b e t w e e n these geographically s e p a r a t e d entities.World distribution: w e s t e r n Atlantic O c e a n : North Carolina a n d Georgia. As
Hinck-
sia sordida
i n the Pacific Ocean: T a s m a n i a , s o u t h e r n Australia a n d n o r t h e r n Chile.Notes on b i o g e o g r a p h y :
Hincksia onslowensis
is a b r o w n alga first d e s c r i b e d from the North Carolina c o n t i n e n t a l shelf a n d later reported from n e a r b y areas. O u r collec- tions, also as d e e p - w a t e r plants at depths of 3 0 - 7 0 m, r e p r e s e n t n e w records a n d greatly e x t e n d its distributional r a n g e to the E a s t e r n Atlantic Ocean. Its area of distribution is rather similar to that figured forSyringoderma floridana
H e n r y a n dGloioctadia blom-
quistii
(Searles) R.E. Norris (Fig. 12).C h o r i s t o c a r p a c e a e
Discosporangium mesarthrocarpum
(Meneghini) H a u c kC o 11 e c t i o n s : # 14964, Sept. 30, Stat. 14, 28o49'54" N, 13051'32 " W, Estrecho La Bocafna b e t w e e n L a n d F, d e p t h 65-70 m; # 14450, Sept. 30, Stat. 17, 28o49'70" N,
130 R. J. Haroun, W. F. P r u d ' h o m m e v a n Reine, D. G. Mfiller, E. Serrao & R. H e r r e r a
don,
b
:,_..
Fig. 3.
Discosporangium rnesarthrocarpum
(Meneghini) Hauck. a: General morphology; b: plurilocuIar sporangia. Scale: 2 mm (a) and 50 ~rn (b}13o40'20 " W, Estrecho La Bocaina b e t w e e n L a n d F. depth 30-40 m; # 14971, Oct. 8, Star. 46, 28~ N, 15~ W, S a r d i n a del Norte, C. depth 30-40 rn.
Filamentous, b r o w n i s h thalli, 2 - 7 m m in length, were collected e n t a n g l e d with other species. Several characters, as, for example, the p r e s e n c e of discoid l a t e r a l plurilocular sporangia, b r a n c h i n g p a t t e r n a n d cell morphology, were u s e d to identify t h e s e s p e c i m e n s as D.
mesarthrocarpum,
a rare b r o w n alga with u n c e r t a i n p h y l o g e n e t i c affinities (Fig. 3a, b).World distribution: M e d i t e r r a n e a n Sea Adriatic Sea, e a s t e r n A t l a n t i c O c e a n : M a d e i r a Archipelago, a n d s o u t h e r n Australia
Notes on b i o g e o g r a p h y :
D. mesarthrocarpum
was d e s c r i b e d from the A d r i a t i c i n the last century, a n d has since b e e n collected in other M e d i t e r r a n e a n localities, as well as i n S o u t h e r n Australia, w h e r e it m a y h a v e b e e n i n t r o d u c e d (Womersley, 1987). In addition, Audiffred & P r u d ' h o m m e v a n R e i n e (1985), collected it at Porto Santo, M a d e i r a Archipelago (north of the C a n a r y Islands). O u r collections, b e t w e e n L a n z a r o t e a n d F u e r t e v e n t u r a Islands at 3 0 - 7 0 m depth, e x t e n d its s o u t h e r n limit to t h e C a n a r i a n Archipelago a n d confirm its p r e s e n c e in the M a c a r o n e s l a n Islands (Fig. 12).S y r i n g o d e r m a t a c e a e Syringoderma floridana H e n r y
C o 11 e c t i o n s : # 14271, Sept. 30, Stat. 14, 28~ N, 13051'32" W, Estrecho La Boca~na b e t w e e n L a n d F, d e p t h 65-70 m; # 14350, Sept. 30, Stat. 15, 28~ N, 13051'32" W, Estrecho La Bocafna b e t w e e n L a n d F, d e p t h 70-80 m; # 1 4 3 9 6 , Sept. 30, Stat. 16, 28~ N, 13~ W, Estrecho La Bocafna b e t w e e n L a n d F, d e p t h 90-96 m; #14783, Oct. 7, Stat. 38, 28~ N, 16~ W, Pto. de Alcal~ T, d e p t h 80-90 m; #15093, Oct. 11, Stat. 53, 28013'78" N, 14044'77" W, S e a m o u n t n e a r F, d e p t h 45-50 m; #15144, Oct. 11, Stat. 56, 28~ N, 14~ W, Pta. Barlovento, F, depth 64 m.
In several d r e d g e hauls, small b r o w n algae identified as S. floridana w e r e collected. The thallus is flabellate, up to 1.5 cm long, monostromatic, with distinct m a r g i n a l growth, a t t a c h e d to the rocky substrate b y rhizoidal filaments a n d covered b y sessile u n i l o c u l a r s p o r a n g i a (Fig. 4a, b, c).
World distribution: w e s t e r n Atlantic Ocean: e a s t e r n Florida.
b
Fig. 4. Syringoderma floridana Henry. a: General morphology; b: T. S. near the margin; c: plurilocu- lar sessile sporangia. Scale: 3 mm (a); 100 ~rn (b) and 50 ~rn (c)
132 R . J . H a r o u n , W. F. P r u d ' h o m m e v a n Reine, D. G. Miiller, E. Serrao & R. H e r r e r a
N o t e s on b i o g e o g r a p h y : This s p e c i e s is a typical m e m b e r of the d e e p - w a t e r flora of the e a s t e r n Florida c o n t i n e n t a l shelf. Until now, collections of s p e c i m e n s t e n t a t i v e l y a s s i g n e d to this g e n u s w e r e r e c o r d e d from C a n a r i a n a n d o t h e r M a c a r o n e s i a n islands (Oosterbaan, Univ. of Leiden, u n p u b l i s h e d report), but an a b s e n c e of fertile structures did not allow e x a c t identification. D u r i n g the p r e s e n t expedition, s e v e r a l s a m p l e s c o n t a i n e d s o m e fertile material. H e n c e its p r e s e n c e in the e a s t e r n Atlantic O c e a n is confirmed, g r e a t l y e x t e n d i n g its k n o w n distributional r a n g e (Fig. 12).
R h o d o p h y t a : P e y s s o n n e l i a c e a e
Peyssonneh'a harveyana
J. A g a r d hC o 11 e c t i o n s : # 15107, Oct. 11, Stat. 15107, 28~ '' N, 14~ " W, S e a m o u n t n e a r F, d e p t h 4 5 - 5 0 m ; #15191, Oct. 11, Stat. 56, 28~ " N, 14~ P t a . Barlovento, F, d e p t h 64 m.
S p e c i m e n s of a clark-red crust i d e n t i f i e d as
P. harveyana
w e r e c o l l e c t e d from the s e a m o u n t n e a r F. T h e thallus w a s strongly a d h e r e n t to the s u b s t r a t u m e x c e p t at its margins, entire or lobed, with a v e r r u c o s e a n d radially striated surface (Fig. 5a, b). In s o m e s p e c i m e n s , i r r e g u l a r sori with cruciate t e t r a s p o r a n g i a w e r e o b s e r v e d .b
a
Fig. 5.
Peyssonnelia harveyana
J. Agardh. a: General morphology; b: T. S. of the thallus. Scale: 10 mm la) and 300 ~rn (b)World distribution: M e d i t e r r a n e a n Sea, e a s t e r n Atlantic O c e a n : British Isles to Portugal.
N o t e s on biogeography~ Previous records of epilithic or e p h i p h y t i c
Peyssonnelia
harveyana
are all from the u p p e r sublittoral to 12 m, not from such d e p t h s as found d u r i n g the p r e s e n t expedition. T h e distribution a r e a of this species i n c l u d e s the M e d i t e r - r a n e a n Sea, a n d the coasts of Portugal a n d the British Isles. O u r records e x t e n d the distributional r a n g e of the species further to the south (Fig. 12).H a l y m e n i a c e a e
Cryptonemia seminervis
(C. Agardh) J. A g a r d hC o 11 e c t i o n s : # 14375, Sept. 30, Stat. 15, 28~ N, 13o51'32 '' W, E s t r e c h o La Bocaina b e t w e e n L a n d F, d e p t h 70-80 m; # 15193, Oct. 11, Star. 56, 28~ '' N, 14~ W, Pta. Barlovento, F, d e p t h 64 m.
In two d r e d g e s a m p l e s from depths b e t w e e n 60-80 m, s e v e r a l fleshy, r e d a l g a e w e r e collected. Thalli w e r e rose-red, up to 5 cm in length, arising from a s m a l l discoidal holdfast, with a short t e r e t e stipe of 0.8 m m d i a m e t e r in b a s a l parts, e x p a n d i n g into fan- s h a p e d b l a d e s w i t h c u n e a t e bases, in w h i c h the stipes e x t e n d as e v a n e s c e n t midribs. M a r g i n s w e r e irregular, with s o m e proliferations (Fig. 6a, b).
b
Fig. 6.
Cryptonemia seminervis
(C. Agardh) J. Agardh. a: General morphology; b: T. S. of the thallus. Scale: 5 mm (a) and 30 ~m (b)134 R . J . H a r o u n , W. F. P r u d ' h o m m e v a n R e i n e , D. G. Mfiller, E. S e r r a o & R. H e r r e r a
F o l l o w i n g P r i c e et al. (1986),
Cryptonemia seminervis
a n dC. luxurians
(C. A g a r d h ) J. A g a r d h a r e t r e a t e d h e r e as a s i n g l e s p e c i e s , w i t h C.seminervis
as t h e e a r l i e s t a v a i l a b l e name.Cryptonemia lomation Berthold, reported by Borgesen (1938) from a deep-water
collection near C, w a s also encountered during this expedition, as well as C. crenulata (J. Agardh) J. Agardh, w h i c h w a s recently reported for T (Gil-Rodriguez et al., 1985) as an intertidal species, Apart from other morphological differences, both species lack the conspicuous midrib observable in C. seminervis.
W o r l d distribution: A s C. seminervis, this species is reported in the East Atlantic ocean, from the Scilly Isles in south-western Britain ( M a g g s & Guiry, 1987), d o w n to southern Spain a n d the Atlantic coast of Morocco; it is also k n o w n from W e s t Africa. It has also b e e n reported as C. luxunans from several localities of the tropical a n d w a r m - temperate regions of the western Atlantic Ocean.
Notes on biogeography: This n e w record fills the gap in the distributional range of this species along the eastern Atlantic coast, resulting in a n e w distributional range which includes the tropical a n d warm-temperate regions of both sides of the Atlantic O c e a n (Fig. 12).
R h o d y m e n i a c e a e
Botryocladia wynnei
B a l l a n t i n eC o 11 e c t i o n s : # 14816, Oct. 7, Stat. 38, 28007'66" N, 16~ W, Pto. Alcal~, T. d e p t h 8 0 - 9 0 m; # 15179, Oct. 11, Stat. 56, 28009'35" N, 14~ W, Pta. B a r l o v e n t o , F, d e p t h 64 m.
T h i s small, r e d a l g a w a s i d e n t i f i e d as
B. wynnei
d u e to its e x t e r n a l m o r p h o l o g y w i t h a s h o r t s t i p e a n d t h e p r e s e n c e of 1 - 7 g l a n d ceils p r o t r u d i n g i n t o t h e i n n e r c a v i t y . T h e c o r t e x is i n c o m p l e t e (Fig. 7a, b, c).a
Fig. 7.
Botryocladia wynnei
Ballantine. a: General morphology; b: surface view of outer wall of a vesicle; c: inner vesicle wall showing a gland-supporting cell with 6 gland cells. Scale: 5 mm (a);O t h e r
Botryodadia
species, n a m e l yB. boergesenii
F e l d m a n n ,B. chia]eana
(Meneghini) Kylin a n dBotryocladia occidentafis
(B~rgesen) Kylin w e r e also c o l l e c t e d d u r i n g this cruise, b u t their g e n e r a l m o r p h o l o g y a n d i n t e r n a l structures differed from B.wynnei
(Ballantine, 1985). A n o t h e r r e l a t e d s p e c i e s isB. lawsonfi
John, w h i c h often has a b r a n c h e d stipe a n d vesicles w h i c h are m u c h l o n g e r (Lawson & John, 1987).W o r l d distribution: w e s t e r n Atlantic O c e a n : Puerto Rico a n d North Carolina. Notes on b i o g e o g r a p h y : B. w y n n e i is a d e e p - w a t e r s p e c i e s first d e s c r i b e d from Puerto Rico a n d l a t e r r e p o r t e d from the c o n t i n e n t a l shelf of N o r t h Carolina. O u r r e c o r d s r e p r e s e n t a n e x t e n s i o n of its distributional r a n g e to the e a s t e r n Atlantic O c e a n (Fig. 12).
Gloiocladia blomquisti
(Searles) R.E. NorrisC o l l e c t i o n s : # 15160, Oct. 11, Stat. 54, 28~ 14~ S e a m o u n t n e a r F, d e p t h 7 0 - 8 0 m ; # 15186, Oct. 11, Stat. 56, 28~ 14~ Pta. Barlovento, F, d e p t h 64 m.
Two s a m p l e s of G.
blomquistii
w e r e c o l l e c t e d n e a r F. The thallus w a s c r e e p i n g , d i c h o t o m o u s l y a n d i r r e g u l a r l y b r a n c h e d , with i r r e g u l a r a n d proliferous m a r g i n s . T h e i n t e r n a l structure is c o m p o s e d of one l a y e r of l a r g e m e d u l l a r y cells s u r r o u n d e d b y a cortex of s e v e r a l l a y e r s of small p i g m e n t e d cells (Fig. 8a, b).!
Fig. 8.
Gloiodadia blomquistii
(Searles) R.E. Norris. a: General morphology~ b: T.S. of the thallus. Scale: 5 mm (a) and 100 ~rn (b)136 R.J. Haroun, W. F. P r u d ' h o m m e v a n Reine, D. G. Miiller, E. Serrao & R. Herrera
World distribution: w e s t e r n Atlantic O c e a n : e a s t e r n Florida a n d North Carolina. Notes on b i o g e o g r a p h y : T h e g e n u s
Gloiocladia
s e n s u Norris (1991) is a d e e p - w a t e r g e n u s with a disjunct distribution a r o u n d the Indo-Pacific a n d Atlantic Oceans. O n the c o n t i n e n t a l shelves of e a s t e r n Florida a n d North Carolina, threeGloiocladia
species are k n o w n ; two of these have b e e n collected for the first time from d e e p - w a t e r habitats in the C a n a r y Islands. In this contribution, only the material a s s i g n e d to G.blomquistii
is reported (Fig. 12). A similar species is t e n t a t i v e l y identified asCloiocladia atlantica
(Searles) R.E. Norris b e c a u s e of its p i n n a t e habit; b u t that s a m p l e n e e d s further study.
Hafichrysis peltata
(W. R. Taylor) P. Huv~ et H. Huv~C o l l e c t i o n s : # 1 5 1 5 3 , Oct. 11, Stat. 54, 28~ 14~ S e a m o u n t n e a r F, d e p t h 70-80 m; # 15189, Oct. 11, Stat. 56, 28~ " N, 14~ " W, Pta. Barlovento, F, d e p t h 64 m.
Small plants a s s i g n e d to
H. peltata
w e r e collected g r o w i n g a t t a c h e d to the sub- stratum b y small discoidal holdfasts. The thallus is shortly stipitate with i r r e g u l a r or l o b e d blades, fleshy, u p to 2.5 cm tall a n d with a n i n t e r n a l structure of v e r y large i n n e r m e d u l l a r y cells a n d a cortex of 2 - 4 layers of small p i g m e n t e d cells (Fig. 9a, b).a m
" ~ ~ : ~ , ~ ~ ~ ~ b
Fig. 9.
FIalichrysis peltata
(W. R. Taylor). P. Huve et H. Huv6. a: General morphology; b: T. S. of the thallus partly through a cystocarp. Scale: 5 mm (a) and 50 ~m (b)A n o t h e r
Halichrysis
species,H. depressa
(J. Agardh) Bornet is reported for the West African coasts as a n intertidal a n d rarely sublittoral species occurring d o w n to d e p t h s of 8-16 m (Lawson & John, 1987). It is also recorded at similar depths i n the M e d i t e r r a n e a n Sea. This latter species differs from our s p e c i m e n s in g e n e r a l m o r p h o l o g y as well as i n its i n t e r n a l structure, which gives rise to some doubts a b o u t the taxonomic status of the g e n u s . See also P. Huv4 & H. Huv4 (1977) a n d R. E. Norris (1991).World distribution: Tropical a n d subtropical coasts of the w e s t e r n Atlantic Ocean. Notes on b i o g e o g r a p h y : The p r e s e n t species w a s first described as
Fauchea peltata
b y Taylor (1942) from Tortuga Island (Venezuela), as a d e e p - w a t e r m a c r o a l g a collected at 40 m. It is n o w k n o w n i n other parts of the tropical a n d w a r m - t e m p e r a t e r e g i o n s of the w e s t e r n Atlantic Ocean. Our s p e c i m e n s r e p r e s e n t a n e w record for the e a s t e r n Atlantic O c e a n (Fig. 12).
Leptofauchea brasiliensis
JolyC o 11 e c t i o n s : # 14845, Oct. 7, Stat. 39, 28~ " N, 16046'62 '' W, Los Cristianos, T, d e p t h 90-110 m.
In a dredge s a m p l e from d e p t h s b e t w e e n 90-110 m, s p e c i m e n s of a small r e d alga with fleshy lanceolate blades w e r e collected. Its i n t e r n a l structure agrees with that of
L. brasiliensis,
with a m e d u l l a r y layer of large h y a l i n e cells s u r r o u n d e d b y 1-2 i n c o m p l e t e layers of m e d i u m - s i z e d cells a n d a cortex with only one layer of small p i g m e n t e d cells (Fig. 10a, b).b - -
Fig. 10.
Leptofauchoa brasfliensisJoly,
a: General morphology; b: T. S. of the thallus. Scale 5 mm (a) and 100 gm (b)138 R . J . Haroun, W. F. P r u d ' h o m m e v a n Reine, D. G. Mfiller, E. S e r r a o & R. H e r r e r a
World distribution: w e s t e r n A t l a n t i c O c e a n : Brazil, Curagao, e a s t e r n Florida a n d N o r t h Carolina.
N o t e s on b i o g e o g r a p h y : This species, d e s c r i b e d as a d e e p - w a t e r p l a n t from t h e Brazilian coast by J o l y (1957), has b e e n r e p o r t e d as a m e m b e r of the d e e p - w a t e r flora from the c o n t i n e n t a l s h e l v e s of e a s t e r n Florida a n d North Carolina. Its o c c u r r e n c e n e a r T r e p r e s e n t s a n e w r e c o r d for the e a s t e r n Atlantic O c e a n (Fig. 12).
Solieriaceae
S a r c o d i o t h e c a d i v a r i c a t a W. R. Taylor
C o l l e c t i o n s : # 1 5 1 2 5 , Oct. 11, Stat. 53, 28~ 14~ S e a m o u n t n e a r F, d e p t h 4 5 - 5 0 m.
T h e thallus of this a l g a is erect, up to 2 cm in length, d i c h o t o m o u s l y d i v i d e d a n d p i n k - r e d in colour. Its i n t e r n a l structure consists of a m e d u l l a of h y a l i n e filaments e m b e d d e d in a g e l a t i n o u s matrix, b o u n d e d b y two layers of large, r o u n d e d , colourless cells a n d a cortex of two layers of small p i g m e n t e d cells (Fig. 11a, b).
World distribution: Pacific O c e a n : G a l a p a g o s Islands; w e s t e r n Atlantic O c e a n : e a s t e r n Florida a n d North Carolina.
~,
~
Fig. 11. Sarcodiotheca divaricata W. R. Taylor. a: General morphology; b: T. S. of the thallus. Scale: 5 mm (a) and 100 ~m (b)
N o t e s on b i o g e o g r a p h y : This species w a s first d e s c r i b e d from d e e p w a t e r off the G a l a p a g o s Islands in the Pacific O c e a n (W. R. Taylor, 1945). S u b s e q u e n t l y , it w a s f o u n d on the c o n t i n e n t a l shelves of e a s t e r n Florida a n d N o r t h Carolina. In the p r e s e n t p a p e r it is r e c o r d e d for the first time, as a d e e p - w a t e r plant, in the e a s t e r n Atlantic O c e a n (Fig. 12).
8rJ~opsis halj~rneniae Cr?ptonemia sernineruls Halichr~sis peltata Dt$cosporangiurn mesarthrocarpum G[oloc/adia b/ornquistii 9 Leptofauchea brasiliensis ,. Pepssonne/ia horuejJana Botryocladia w?nnei .. Sorcodiotheca d/varir
Fig. 12. Distributional maps of newly recorded species in the Canary Islands9 Hatched area: previously known area; thick arrow: Canary Islands; thin arrow: Madeira Archipelago
140 R . d . Haroun, W. F. P r u d ' h o m m e v a n Reine, D. G. Mfiller, E. S e r r a o & R. H e r r e r a
D I S C U S S I O N
T h e C a n a r y I s l a n d s are y o u n g volcanic i s l a n d s f o r m e d less t h a n 20 m i l l i o n y e a r s a g o (20 Ma), a n d a r e s e p a r a t e d b y d e e p trenches. H o w e v e r , the e a s t e r n m o s t i s l a n d s , L a n z a - rote a n d F u e r t e v e n t u r a , m a y h a v e o r i g i n a t e d as f r a g m e n t s of t h e A f r i c a n p l a t e , w h i c h drifted to d e e p e r w a t e r s . T h e r i c h n e s s of the C a n a r y m a r i n e a l g a l flora is t h o u g h t to b e the result of p a l e o c l i m a t i c events, its specific g e o g r a p h i c a l position, a n d the l a s t i n g a v a i l a b i l i t y of m a n y different m i c r o c l i m a t e s a l o n g its coasts (Afonso-Carrillo & Gil- Rodrlguez, 1982; P r u d ' h o m m e v a n Reine & v a n d e n Hoek, 1990).
Most of t h e s t u d i e s on s e a w e e d b i o g e o g r a p h y a r e d e v o t e d to t h e a n a l y s i s of p a l e o - or p r e s e n t distribution a r e a s of i n t e r t i d a l a n d s h a l l o w - w a t e r m a c r o a l g a l s p e c i e s . D u r i n g the last 20 years, a rich d e e p - w a t e r s e a w e e d flora w a s d i s c o v e r e d on s c a t t e r e d offshore reefs in b e t w e e n N o r t h C a r o h n a a n d e a s t e r n Florida. M o r e s t a b l e e n v i r o n m e n t a l c o n d i t i o n s p r e v a i l in such d e e p habitats, m a k i n g t h e m s u i t a b l e for the survival of s e v e r a l tropical to w a r m - t e m p e r a t e s p e c i e s ( S c h n e i d e r & Searles, 1973; Searles, 1984, 1987; H a n i s a k & Blair, 1988). The m o r e c o n s t a n t a n d m o d e r a t e t e m p e r a t u r e conditions i n d e e p - w a t e r e n v i r o n m e n t s allow t h e survival of s p e c i e s on b o t h sides of the A t l a n t i c O c e a n . O u r findings also e x t e n d t h e d e p t h r a n g e s for s p e c i e s p r e v i o u s l y l i s t e d o n l y from s h a l l o w - w a t e r , such as
Peyssonnelia harveyana
a n dBryopsidella neglecta,
w h i c h s u g g e s t s t h a t t h e s e m a c r o a l g a e a r e only a b l e to survive in d e e p - w a t e r h a b i t a t s w h e n t h e y r e a c h their l a t i t u d i n a l limits. A similar p h e n o m e n o n was r e p o r t e d b y S c h n e i d e r & S e a r l e s (1973) for N o r t h C a r o l i n a offshore m a c r o a l g a e w i t h a C a r i b b e a n origin. T h e d e e p - w a t e r e n v i r o n - m e n t s e r v e s as a safe p l a c e a n d is also a r e f u g e for m a n y g r o u p s of a n i m a l s , such as m o n o p l a c o p h o r a n s a n d s t a l k e d crinoids, that in m o r e a n c i e n t times w e r e c o m m o n in s h a l l o w w a t e r (Vermeij, 1980). T h e p e r s i s t e n c e of t h e s e g r o u p s c a n b e a t t r i b u t e d to t h e fact that in the d e e p sea, no m a s s extinction e v e n t s o c c u r r e d as t h e y did in s h a l l o w w a t e r s d u r i n g c o l d e r p e r i o d s (Vermeij, 1987; Breeman, 1990). Besides, m a n y d e e p - w a t e r m a c - r o a l g a e s h o w p h y s i o I o g i c a l a n d m o r p h o l o g i c a l a d a p t a t i o n s for survival u n d e r t h e s p e c i a l conditions of d e e p - w a t e r h a b i t a t s (Littler et al., 1986; Lfining, I990).The n e w d i s t r i b u t i o n a l r e c o r d s r e p o r t e d h e r e s u p p o r t the h y p o t h e s i s t h a t t h e d e e p - w a t e r flora of the C a n a r y I s l a n d s is closely r e l a t e d to the w a r m - t e m p e r a t e w e s t e r n Atlantic flora (eSpecially
Hincksia onslowensis, Syringoderma floridana, Gloiodadia
blomquistii, Leptofauchea brasiliensis
a n dSarcodiotheca divaricata)
as w e l l as to the W. M e d i t e r r a n e a n flora (especiallyB. neglecta, Discosporangium mesarthrocarpum
a n d P.harveyana).
T h e d i s t r i b u t i o n a l r a n g e ofSarcodiotheca divaricata
c a n b e e x p l a i n e d b y the r a t h e r r e c e n t closure of the P a n a m a Isthmus (3.1-3.5 Ma) a n d illustrates a close affinity b e t w e e n the m a r i n e biota of b o t h coasts of C e n t r a l A m e r i c a (Vermeij, 1980; Liining,1990).
A long d i s p e r s a l r a n g e for d e e p - w a t e r s e a w e e d s is r a t h e r difficult to a c c e p t ; the l i k e l i h o o d of d e e p - w a t e r s p e c i e s b e c o m i n g p a r t of a drifting s e a w e e d raft is r a t h e r small (van d e n H o e k , 1987; S e a r l e s & S c h n e i d e r , 1987; Lfining, 1990). F u r t h e r m o r e , g l a c i a t i o n periods, with t e m p e r a t u r e s a few c e n t i g r a d e s l o w e r t h a n today, w e r e sufficient to shift the n o r t h e r n l e t h a l b o u n d a r i e s of s o m e strictly s u b t r o p i c a l or t r o p i c a l - t o - s u b t r o p i c a l s p e c i e s to the south of the M a c a r o n e s i a n I s l a n d s s e n s u stricto (Madeira, S a l v a g e s a n d Canaries). Therefore, the n e w d i s t r i b u t i o n a l p a t t e r n s of the d e e p - w a t e r s p e c i e s r e p o r t e d h e r e m a y s u p p o r t the v i c a r i a n c e h y p o t h e s i s a l r e a d y p r o p o s e d b y P r u d ' h o m m e v a n R e i n e & v a n d e n
}-Ioek (1990) for t h e s e a w e e d f l o r a of t h e M a c a r o n e s i a n I s l a n d s . T h e n e w r e c o r d s s u g g e s t a c l o s e r a f f i n i t y b e t w e e n d e e p - w a t e r f l o r a s p r e s e n t i n o f f s h o r e r e e f s o r c o n t i n e n t a l s h e l v e s of b o t h s i d e s of t h e A t l a n t i c ( S e a r l e s , 1984; H a n i s a k & Blair, 1988). F u r t h e r m o r e , F r e d e r i c q e t al. (1993) r e p o r t e d s o m e n e w r e c o r d s of s u b t i d a l m a c r o a l g a e f r o m t h e A z o r e s I s l a n d s , m o s t l y w i t h a t r o p i c a l e a s t e r n A t l a n t i c ( W e s t A f r i c a n ) o r M e d i t e r r a n e a n d i s t r i b u - t i o n r a n g e . T h e s e f i n d i n g s c a n a l s o b e e x p l a i n e d i n t h e f r a m e w o r k of a T e t h y a n r e m n a n t s h y p o t h e s i s , t h a t is, v i c a r i a n t f l o r a s o c c u p y i n g t h e w e s t e r n a n d e a s t e r n c o a s t s of t h e A t l a n t i c O c e a n ( P r u d ' h o m m e v a n R e i n e & v a n d e n H o e k , 1990). A d d i t i o n a l r e s e a r c h is n e e d e d , b o t h a t floristic a n d p h y s i o l o g i c a l l e v e l s , to i n c r e a s e o u r u n d e r s t a n d i n g of t h e b i o g e o g r a p h i c a l r e l a t i o n s h i p s of t h e d e e p - w a t e r f l o r a s p r e s e n t i n t h e A t l a n t i c O c e a n .
A c k n o w l e d g e m e n t s . This p a p e r is b a s e d on the collections m a d e during a n expedition of the R.V. Heincke of the Biologische Anstalt Helgoland, Hamburg. We are grateful to Dr. K. L6ning a n d Dr. G. G a s s m a n n (B.A.H., Hamburg), who organized this expedition, a n d to the smiled efforts of the captain a n d crew of the ship.
L I T E R A T U R E C I T E D
Afonso-Carrillo, J. & Gil-Rodriguez, M. C, 1982. Aspectos biogeogr~ficos de la flora ficol6gica m a r i n a de las Islas Canarias. - Actas 2. Simp. ib~r. Estud. Bentos mar., 3, 41-48.
Afonso-Carrillo, J. & Sans6n, M., 1989. Clave ilustrada p a r a la d e t e r m i n a c i 6 n de los macr6fitos marinos bentSnicos de las Islas Canarias. Dpt. Biol. Veg. Univ. La Laguna, La Laguna, 55 pp. Amsler, C. D. & Kapraun, D. F., 1985. Giffordia o n s l o w e n s i s sp. nov. (Phaeophyceae) from the
Northern Carolina continental shelf and the relationship b e t w e e n Giffordia a n d Acinetospora. -
J. Phycol., 2I, 94-99.
Audiffred, P. A. J. & P r u d ' h o m m e v a n Reine, W. F., 1985. Marine algae of tlha do Porto Santo a n d Deserta G r a n d e (Madeira Archipelago). - Bolm Mus. munic. Funchal 37, 20-51.
Ballantine, L., 1985. Botryocladia w y n n e i sp. nov. and B. spinuiifera (Rhodymeniales, Rhodophyta) Taylor & A b b o t t from Puerto Rico. - Phycologia, 24, 199-204.
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