Content uploaded by Vezio Cottarelli
Author content
All content in this area was uploaded by Vezio Cottarelli on Sep 22, 2014
Content may be subject to copyright.
This article was downloaded by: [109.73.70.165]
On: 19 March 2014, At: 18:54
Publisher: Taylor & Francis
Informa Ltd Registered in England and Wales Registered Number: 1072954 Registered office: Mortimer
House, 37-41 Mortimer Street, London W1T 3JH, UK
Bolletino di zoologia
Publication details, including instructions for authors and subscription information:
http://www.tandfonline.com/loi/tizo19
Taxonomical and ecological observations on
Pycnogonida from Apulian coasts (southern Italy)
Carla Chimenz a , Marco Tosti a & Vezio Cottarelli b
a Dipartimento di Biologia Animale e dell'Uomo , Università di Roma ≪La Sapienza≫
, viale dell'Università 32, I‐00185, Roma, Italy
b Dipartimento di Scienze Ambientali , Università della Tuscia , via S. Camillo De
Lellis, I‐01100, Viterbo, Italy
Published online: 28 Jan 2009.
To cite this article: Carla Chimenz , Marco Tosti & Vezio Cottarelli (1993) Taxonomical and ecological
observations on Pycnogonida from Apulian coasts (southern Italy), Bolletino di zoologia, 60:3, 339-347, DOI:
10.1080/11250009309355834
To link to this article: http://dx.doi.org/10.1080/11250009309355834
PLEASE SCROLL DOWN FOR ARTICLE
Taylor & Francis makes every effort to ensure the accuracy of all the information (the “Content”)
contained in the publications on our platform. However, Taylor & Francis, our agents, and our licensors
make no representations or warranties whatsoever as to the accuracy, completeness, or suitability
for any purpose of the Content. Any opinions and views expressed in this publication are the opinions
and views of the authors, and are not the views of or endorsed by Taylor & Francis. The accuracy of
the Content should not be relied upon and should be independently verified with primary sources of
information. Taylor and Francis shall not be liable for any losses, actions, claims, proceedings, demands,
costs, expenses, damages, and other liabilities whatsoever or howsoever caused arising directly or
indirectly in connection with, in relation to or arising out of the use of the Content.
This article may be used for research, teaching, and private study purposes. Any substantial or
systematic reproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in
any form to anyone is expressly forbidden. Terms & Conditions of access and use can be found at http://
www.tandfonline.com/page/terms-and-conditions
Boll. Zool. 60: 339-347 (1993)
Taxonomical and ecological
observations on Pycnogonida from
Apulian coasts (southern Italy)
CARLA CHIMENZ
MARCO TOSTI
Dipartimento di Biologia Animale e dell'Uomo,
Università di Roma «La Sapienza»,
viale dell'Università 32, I-00185, Roma (Italy)
VEZIO COTTARELLI
Dipartimento di Scienze Ambientali, Università della Tuscia,
via S. Camillo De Lellis, I-01100, Viterbo (Italy)
INTRODUCTION
The Pycnogonida described in this paper were isolated
from samples of benthos collected during ecological
researches carried out along an area of the Apulian coasts.
Pycnogonida of the southern Adriatic coast had never
previously been studied. The material described here
may provide useful taxonomic and biogeographic infor-
mation, as well as represent a basis for future biological
studies. In particular, the presence in our samples of both
males and females of Rhynchothorax alcicornis enables
us to describe the female, hitherto unknown, and to
point out the morphological differences between male
and female, thus completing the description of this
species.
MATERIALS AND METHODS
ABSTRACT
One-hundred-sixty specimens of Pycnogonida were collected
from the Apulian coast near Brindisi. Some taxonomically interesting
species are described in detail and the composition of the
populations of Apulian Pycnogonida discussed in relation to the
ecological characteristics of the sampled biocoenoses. None of the
adventitious species of Pycnogonida recorded from other
Mediterranean areas was included in these biocoenoses.
KEY WORDS: Pycnogonida - Taxonomy - Animal ecology -
Biogeography - Mediterranean sea.
ACKNOWLEDGEMENTS
We are very grateful to Drs. J. H. Stock, F. Krapp and T. Munilla,
and to Mrs. F. Arnaud for their helpful information and comments on
the two species of Anoplodactylus. We also thank Dr. A. Belluscio
who entrusted the study of this material to us. Some «camera lucida»
drawings were made by Dr. N. Falchi. The SEM micrographs were
made by Dr. L. Del Caldo (Dipartimento di Biología Végétale, Univer-
sità di Roma «La Sapienza») and Dr. M. Carcupino (Dipartimento di
Scienze Ambientali, Università della Tuscia, Viterbo). The research
was partly supported by the Italian Ministero della Università e della
Ricerca Scientifica e Tecnológica (funds 60).
Sampling was carried out in July 1987 along the coast between
Punta Penne and Casalabate, approximately 10 km North and 30 km
South of Brindisi, respectively. The stations were set along thirteen
transects at depths ranging from 0.5 to 28 m (Fig. 1).
The bottoms monitored showed the typical biocoenotic features
of the Adriatic coast of the Salento region (Bedulli et al., 1986). The
various biocoenoses did not follow one another regularly according
to the depth and actually displayed a mosaic structure. Therefore,
the places in which Pycnogonida were collected will be referred to
using the following five biocoenotic classes identified according to
Pérès & Picard (1964) (and not to the single transects): (1)
photophilic algae (AP); (2) fine well-sorted sands (SFBC); (3) coarse
sands under bottom currents (SGCF); (4) Posidonia oceánica (HP);
(5) coralligenous (Cor). These biocoenoses are abbreviated in the
text for the sake of simplicity.
Sampling on hard bottoms was carried out by SCUBA diving, by
total «scraping» of a surface of 20x20 cm. A modified Charcot dredge
was used on soft substrates. The benthos samples were fixed in 10
neutralized formalin. After sorting, the specimens of Pycnogonida
40° 30' N
(Received
24 February 1992 - Accepted 14 May 1993) Fig. 1 - Location of the sampling stations along the Apulian coast,
southern Italy.
Downloaded by [109.73.70.165] at 18:54 19 March 2014
340
C.
CHIMENZ, M. TOSTI, V. COTTARELLI
were
transferred in 70 ethyl alcohol. Measurements were taken
using
a micrometric slide, according to the method proposed by
Munilla
(1986), and are given in mm.
For
observations under the Scanning Electron Microscope (SEM),
the
specimens were dehydrated through an ethanol series, dried
with
a critical point dryer (BALZERS CPD 020) and observed with a
Cambridge
Stereoscan 200 microscope. . •
The
specimens are preserved in one of the authors' (C. C.) collec-
tions
at the «Dipartimento di Biología Animale e dell'Uomo» of «La
Sapienza»
University of Rome.
RESULTS
AND DISCUSSION
Taxonomic account
Pycnogonida were found in 11 out of 13 transects.
From a total of 51 samples examined l60 specimens were
collected and identified (see Table I). Only the most in-
teresting taxa are here described.
Anoplodactylus cf. petiolatus (Kroyer, 1844)
Fig.
2
AP,
2 m: 1 ovigerous female.
Dimensions. Total length (from the tip of the proboscis
to the, tip of the abdomen, in dorsal view): 1:34.
Proboscis (ventral view): 0.42. Abdomen (dorsal view):
0.25.
Cheliphore (dorsal view): 0.40. First leg (dorsal
view, claw included): 1.90. Main claw: 0.19- Auxiliary
claw: 0.03. Cutting lamina: 0.11.
Remarks - A comparison with the typical form of A.
petiolatus (of which we also isolated some specimens
from our samples) reveals some evident differences:
smaller size, more compact trunk, shorter neck, absence
of tubercles on lateral processes, shorter appendages,
longer cutting lamina preceded by 2-3 spinules instead of
4-6.
On the other hand, similarities exist with A. petio-
latus as described by Marcus (1940) as well as with the
type of A. petiolatus as illustrated by Stock (1975) with
regard to lamina length, propodus spinulation, and ab-
sence of tubercles on the lateral processes. Among the
other species of Anoplodactylus known, our specimen
resembles only A. trispinosus Stock, 1951 as regards the
propodus spinulation and the cutting lamina length.
However, it differs for the absence of tubercles on the
lateral processes, in the shape of the proboscis (as the
subterminal constriction is missing), and in the chelae
wich bear two denticulations on the inner side of both
fingers, the immovable finger being slightly curved, and
the movable markedly curved.
As no male specimens were available, account could
not be taken of the structure of the oviger and the shape
and position of the cement gland duct, which are
necessary for reliable taxonomic assignment in this
genus.
TABLE
I - Distribution of Pycnogonida at different depths (m) in the five biocoenoses studied along the Apulian coast.
Species
0.5-2
AP
Biocenosis
SFBC
SGCF
6-8
11-13
6-136-28
HPCor
7-11 15-21
21-28
Achelia
echinata Hodge, 1864 9
Achelia
langi (Dohrn, 1881) 4
Ammothella
biunguiculata (Dohrn, 1881)
Ammothella
longipes (Dohrn, 1881) 6 6
Ammothella
uniunguiculata (Dohrn, 1881) 7
Ammothella
longioculata (Faraggiana, 1940)
Ascorhynchus
simile Fage, 1942
Tanystylum
conirostre (Dohrn, 1881) 6 1
Anoplodactylus
angulatus (Dohrn, 1881) 3 2
Anoplodactylus
petiolatus (Krayer, 1844)
Anoplodactylus
ci petiolatus 1
Anoplodactylus
pygmaeus (Hodge, 1864) 2
Anoplodactylus
virescens (Hodge, 1864) 32
Anoplodactylus
compositus (Chimenz, Cot., To., 1991)
Anoplodactylus
sp. 4
Rhynchothorax
alcicornis Krapp, 1973
Rhynchothorax
mediterraneus Costa, 1861
Callipallene
cf pbantoma (Dohrn, 1881) 1
Callipallene
ind. juv.
Pycnogonum
plumipes Stock, I960 2
Pycnogonum
pusillum Dohrn, 1881 2
13
18
AP,
photophilic algae; SFBC, fine well sorted sands; SGCF, coarse sands under bottom currents; HP, Posidonia «matte»; Cor, coralligenous.
Downloaded by [109.73.70.165] at 18:54 19 March 2014
PYCNOGONIDA FROM APULIAN COASTS341
Fig. 2 - Anoplodactylus cf. petiolatus (female): a) habitus; b) chela;
c) propodus; d) first leg (right).
Anoplodactylus sp.
Fig. 3
AP,
0.5 m: 2 males (one ovigerous), 2 ovigerous females; HP, 7 m: 1
male.
Dimensions of the male. Total length: 1.70; Proboscis:
0.50; Abdomen: 0.33; Cheliphore: 0.48; Main claw: 0.21;
Auxiliary claw: 0.03; Cutting lamina: 0.15. Dimensions of
the female. Total length: 1.37; Proboscis: 0.46; Ab-
domen: 0.34; Cheliphore: 0.45; Main claw: 0.21;
Auxiliary claw: 0.02; Cutting lamina: 0.16.
Remarks - The specimens found can be assigned to
the pygmaeus «complex» {sensu Stock, 1975); they show
similarities to A. pygmaeus (Hodge, 1864), the only
species belonging to this complex so far collected from
the Mediterranean. However, the following charac-
teristics distinguish our specimens from this species: a
larger size; the presence of auxiliary claws (Fig. 3f); the
feebly or not segmented trunk (Fig. 3a, d); the shape of
the ocular tubercle; the oviger with 7 segments (instead
of 6) (Fig. 3h); the more prominent genital spurs on the
second coxa of the male (Fig. 3e); the less curved
propodus (Fig. 3f). On the other hand, some features of
our specimens recall A. turbidus Stock, 1975: the body
shape; the presence of tubercles on the lateral processes;
curved and dentate cheliphore fingers (Fig. 3g); the long
cutting lamina preceded by two spines; the presence of
auxiliary claws; the shape and position of the cement
gland duct (Fig. 3e). However, our specimens have a
longer abdomen, reaching the distal part of the first coxa;
proboscis lacking ventral spinules; oviger with 7 seg-
ments instead of 6; much more prominent genital spurs
on the seconda coxa of the male. Two other species of
the pygmaeus «complex», A. micros Bourdillon, 1955,
and A. trispinosus Stock, 1951, share with our specimens
the presence of auxiliary claws, cutting lamina, and
tubercles on the lateral processes; but the other features
are different. In conclusion, this Anoplodactylus cannot
be ascribed to any known species of the pygmaeus «com-
plex». It might be a new taxon, but we prefer to deter-
mine its systematic position through further comparisons
and surveys.
Rhynchothorax alcicornis Krapp, 1973
Figs 4-8
SGCF,
28 m: 1 female; Cor, 21 - 28 m: 6 males, 5 females, 2 juv.
Dimensions of the male. Total length: 1.40; Proboscis:
0.45;
Abdomen: 0.26; Palp: 0.57; Oviger: 0.72; Fourth
leg:
1.23. Dimension of the female. Total length: 1.38;
Proboscis: 0.37; Abdomen: 0.25; Palp: 0.77; Oviger: 1;
Fourth leg: 1.25.
Description of the female. Tegument strewn with
numerous little tubercles; each one bears a pair of curved
setules (represented in only a few drawings). The body
tapers from the first to the fourth segment. Segmentation
lines are present between all trunk segments. The dorsal
ornamentation consists of six small tubercles on the
distal part of cephalon; one (sometimes two) mid-dorsal
tubercle(s) lie on the first segment, two tubercles on the
second and third segments; only one tubercle on the
fourth one (Figs 5, 6). The proboscis is barrel-shaped,
with well-developed dorsal antimere and dorsal and ven-
tral sides with wrinkles and wide flat bosses (Fig. 7a).
Eyes are absent; the ocular tubercle is reduced (Figs 5a,
7a).
As already described above, in the zone of the ocular
tubercle there are six small roundish protuberances
arranged as illustrated in Figures 5a and 7c. The lateral
processes are armed with distal multiple apophyses; the
first segment shows only one apophysis on the posterior
side;
the fourth segment bears one apophysis on the an-
terior side; the second and third segment bearing two
apophyses on the anterior and posterior sides (Figs 5a, 6).
The abdomen is cylindrical, long, almost reaching the
distal extremity of the second coxa of the fourth leg; its
extremity is slightly bifid (Fig. 5a). True cheliphores are
absent; they are replaced by two formations resembling
those of other species of the same genus, defined by
Child (1979) as «cephalic extensions»; they show a basal
constriction (Fig. 7d). Palps are five-segmented, longer
than the proboscis (Fig. 5b); they have a shape like that
described for the male by Arnaud & Krapp (1990). Oviger
ten-segmented; fourth and sixth articles the longest; the
seventh article bearing 2 compound and 2 simple spines,
the eighth 4 compound spine and 1 simple, the ninth and
tenth 2 compound spines and 1 simple one each; in ad-
Downloaded by [109.73.70.165] at 18:54 19 March 2014
342C. CHIMENZ, M. TOSTI, V. COTTARELLI
Fig. 3 - Anoplodactylus sp., female:
a) habitus, trunk, proboscis, abdomen;
b) fourth leg (right), c) chela. Male:
d) habitus; e) fourth leg (right);
f) propodus; g) chela; h) oviger.
dition to the usual claw, article ten is armed with a
serrated lamina (Fig. 5c). Almost all articles bear the
above described tubercles with paired setules. Legs
distally tapering, with subequal coxae; first coxa of all
legs showing two multiple apophyses (Fig. 5a); the
second coxa of the first, second and third leg with one
apophysis on the outer side and one long tubercle on the
inner side; lacking apophyses, but enlarged and with a
large swelling facing the inner side, bearing the opercule
of the genital opening in the fourth leg (Fig. 8a, b). Third
coxa without apophyses but with tubercles. Femur and
tibiae almost subequal in length; femur and tibia 2
bearing a long dorsal seta near distal extremity; tibia 1
with a similar seta at about half its length. Tarsus short,
with two setules on the ventral side; propodus long, slen-
der, with four spine-like setules on the sole and four
similar setules plus two distal paired setules on dorsal
side (Fig. 5d).
The main differences between male and female are
illustrated in Figures 4, 5, and 8. Summarizing: the fourth
article of the male oviger has four stout setae, lacking in
the female; the last four articles of the male oviger show
different spinulation from the female (see Figs 4c and 5c);
the second coxa of the fourth leg in the male, not
enlarged, bears one apophysis on the outer side and
only one long tubercle on the inner side; lastly, the
tegument of the male shows a larger number of little
tubercles.
Remarks - Our material shows some differences com-
pared with the specimens described by Krapp (1973) and
Arnaud & Krapp (1990). The cephalic extensions seem to
be more developed than can be seen in the drawings of
Arnaud & Krapp (1990), as well as in the descriptions of
other species of the genus (see for example Child, 1979)
and show a basal constriction; the palps are longer than
the proboscis (in Arnaud & Krapp's drawings they seem
as long as the proboscis); the process on the palp fourth
Downloaded by [109.73.70.165] at 18:54 19 March 2014
PYCNOGONIDA FROM APULIAN COASTS343
Fig. 4 - Rhynchothorax alcicornis, male: a) habitus, trunk, abdomen;
b) palp; c) oviger; d) fourth leg (right).Fig. 5 - Rhynchothorax alcicornis, female: a) habitus; b) palp;
c) oviger; d) fourth leg (left).
segment is not articulated; the spinulation of the male
oviger is different.
ECOLOGICAL AND BIOGEOGRAPHICAL REMARKS
The differences in abundance and specific com-
position among the Pycnogonida assemblages of the
sampled biocoenoses need to be discussed in relation to
the ecological features of these same biocoenoses (Table
I).
The richest and most diversified assemblage (95
specimens, 13 species) was that of the photophilic algae
(AP).
Even though partly accounted for by the larger
number of samples collected, this richness seems to be a
typical feature of this biocoenosis, which is characterized
by numerous microenvironments entailing diversified
and plentiful trophic resources (Arnaud & Bamber, 1987).
The species found, among which A. virescens was the
most representative, have been usually reported as living
on hard substrates covered with algae, in particular at
shallow depths (Krapp, 1973). In order to reveal any
possible depth-linked difference, biocoenosis AP was
divided into three bathymétrie belts: 0-2 m, 6-8 m, and
11-13 m. No changes of species connected with depth
were noted; the differences were mainly due to the
greater abundance and specific richness at a shallower
level, probably linked to the richer algal assemblage.
The coralligenous assemblage, although poorly diver-
sified, was fairly abundant: 34 specimens, identified
mainly as R. alcicornis and R. mediterraneus, that com-
monly occur in concretionary environments (Arnaud,
1987);
P. plumipes was also typical of this biocoenosis,
where also the new species A.
compost
tus was collected
(Chimenz et al., 1991).
In the Posidonia «matte», 22 specimens were collected
and assigned to nine species. Some of these, A. echinata,
A.
longipes, T. conirostre, A. angulatus, typical of hard
algae covered substrates, also occur in AP biocoenosis;
others, such as A. biunguiculata, are psammophilous, or
occur on sandy or muddy substrates, like A. longioculata
and A petiolatus. In this case, the coexistence of species
Downloaded by [109.73.70.165] at 18:54 19 March 2014
344C. CHIMENZ, M. TOSTI, V. COTTARELLI
ob
Downloaded by [109.73.70.165] at 18:54 19 March 2014
Mm-wie^m
W w/wtät
•?>-:-
T3
n
z
o
o
o
5
•a
33
O
2
>
z
n
o
>
Fig. 7 - Rhynchothorax alcicornis: a) proboscis; b) palp; c) tubercles of the cephalic area; d) cephalic extensions
Downloaded by [109.73.70.165] at 18:54 19 March 2014
346C. CHIMENZ, M. TOSTI, V. COTTARELLI
s?'A$t®£?
Fig. 8 - Rhynchothorax alcicornis. Female: a) II coxa of fourth leg (arrow); b) the same enlarged, showing the genital opening. Male: c) II coxa
of fourth leg (arrow); d) the same, enlarged.
typical of diversified biocoenoses or substrates is found,
rather than a clearly characterized assemblage. This
aspect has been observed in the other groups living in
the Posidonia meadow and suggested to Bianchi et al.
(1989) the hypothesis that Posidonia should be con-
sidered a «complex climatic mesoecosystem», rather than
a biocoenosis.
The assemblages of the soft bottom biocoenoses were
very poor: in the coarse sands under bottom currents
biocoenosis five specimens assigned to five species were
collected, including the psammophilous A. biungui-
culata and the detritus-loving Ascorhynchus simile;
whilst the assemblage of the fine well sorted sands
biocoenosis included four specimens and three species,
two of which (A. petiolatus and R. mediterraneus) are
linked to the sandy or muddy component of the sub-
strate.
From the biogeographical point of view, in the area un-
der consideration 17 out of the 50 species listed by
Arnaud (1988) for the whole Mediterranean sea have
been found. Some of these species, including R. alcicor-
nis,
where previously considered rare. The Brindisi
assemblage showed a lower affinity with those of the
northern and central Adriatic than with those of the
eastern and eestern Mediterranean Basins, confirming
also for the Pycnogonida what had been observed for
other taxonomical groups (Pérès, 1967). It is worth poin-
ting out that no adventitious species previously recorded
in other Mediterranean areas, such as Ammotbea hilgen-
dorfi (Böhm, 1879), found by Krapp & Sconfietti (1983),
and Anoplodactylus portus Caiman, 1927, recorded by
Chimenz et al. (1979), was collected.
REFERENCES
Arnaud F., 1987 - Les Pycnogonides (Chelicerata) de Méditerranée:
distribution écologique, bathymétrique et biogéographie.
Mésogée, 47: 37-58.
Arnaud F., Bamber R. N., 1987 - The biology of Pycnogonida. Adv.
mar. Biol., 24: 1-96.
Downloaded by [109.73.70.165] at 18:54 19 March 2014
PYCNOGONIDA FROM APULIAN COASTS347
Arnaud F., Krapp F., 1990 - The genus Rhynchothorax (Pycnogonida)
in the Mediterranean Sea. Beaufortia, 41: 1-6.
Bedulli B., Bianchi C. N., Morri C, Zurlini G., 1986 - Carat-
terizzazione biocenotica e strutturale del macrobenthos delle
coste pugliesi. In: M. Viel & G. Zurlini (eds), Indagine ambientale
del sistema marino costiero della regione Puglia. ENEL Publ.,
Roma, pp. 227-255.
Bianchi C. N., Bedulli D., Morri C, Occhipinti Ambrogi A., 1989 -
L'herbier de Posidonies: écosystème ou carrefour éco-
éthologique? In: C. F. Boudouresque, A. Meinesz, E. Fresi, V.
Gravez (eds), International Workshop on Posidonia Beds. GIS
Posidonie publ., pp. 257-272.
Bourdillon A., 1955 - Les Pycnogonides de la croisière 1951 du
«President Theodore Tissier». Rev. Trav. Inst. Pèches marit., 19:
581-609.
Child C. A., 1979 - Shallow-water Pycnogonida of the Isthmus of
Panama and the coasts of Middle America. Smithson. Contr.
Zool., 293- 1-86.
Chimenz C, Brignoli P. M., Basciano G., 1979 - Pantopodi del porto
di Civitavecchia e dintorni (Italia Centrale). Cah. Biol. mar., 20:
471-497.
Chimenz C, Cottarelli V., Tosti M., 1991 - Researches on the
Mediterranean Pycnogonida. I. Anoplodactylus compositus
n. sp. (Arthropoda, Chelicerata). Fragm. entomol., 23: 7-13.
Krapp F., 1973 - A fourth Mediterranean Rhynchothorax and remarks
on the genus (Pycnogonida). Bull. zool. Mus. Univ. Amst., 3:
119-124.
Krapp F., Sconfietti R., 1983 - Ammothea hilghendorji (Böhm, 1879),
an adventitious pycnogonid new for the Mediterranean Sea. Mar.
Ecol., 4: 123-132.
Marcus E., 1940 - Os Pantopoda brasileiros e os demais sul-
americanos. Bol. Fac. Filos. Cienc. Let. Univ. Sao Paulo, 19, Ser.
Zool. 4: 3-144, 17 pls.
Munilla T., 1986 - Biometria de una poblaciòn de Ammothella
longipes (Hodge, 1864) (Pycnogonida). Orsis, 2: 103-114.
Pérès J. M., Picard J., 1964 - Nouveau manuel de bionomie benthique
de la mer Méditerranée. Rec. Trav. St. mar. Endoume, 47:
1-137.
Stock J. H., 1951 - Pantopoda. Résultats scientifiques des croisières
du Navire-école belge «Mercator». Mém. Inst. Sci. nat. Belg., 43:
1-23.
Stock J. H., 1975 - Biological results of the University of Miami Deep-
Sea Expeditions: 108. Pycnogonida from continental
shelf,
slope
and deep sea of the tropical Atlantic and east Pacific. Bull. mar.
Sci.,
24: 957-1092.
Downloaded by [109.73.70.165] at 18:54 19 March 2014