XLIV]
THE FISHES OF THE BRITISH OUIANA CONTINENTAL SHELF
66I)
THE FISHES OF THE BRITISH GUIANA CONTINENTAL SHELF,
ATLANTIC COAST OF SOUTH AMERICA, WITH NOTES ON THEIR
NATURAL HISTORY
ROSEMARY
H. LOWE(MCCONNELL)
Fisheries Division, Georgetown, British Guianal
(Communicated by the Zoological rSecret.er\-)
(Accepted for ptib1icat)iondune 1962)
CONTENTS
I. INTRODUCTION
: THE ARXAFISHED
Page
( a ) Collection of d a t a . . . . . . . . . . . . . . . . . . . . . . . . .
( b ) Physical characteristics of the area. . . . . . . .
(c) Invertebrates characteristic of the area.. . .
( d ) Fish fauna zones. . . . . . . . . . . . . . . . . . . . . . . .
11. FISH SPECIESCAUGHT
..........
( a ) Species and abundance of trawl-caught fish
( b ) The commoner fish species prcsent in the different zones. .......................
677
( c ) Comparison of numbers of species and fish in the different zones. . . . . . . . . . . . . . . . 680
111. SEASONALBIOLOGY
(a) Seasonal changeain zone I1 . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
681
682
( b ) Seasonal changes in gonad states of the fish.. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
(c) The occurrence of young in viviparous and ovoviviparous species. . . .
TV. FOOD
RELATIONSHIPS
................................................
v. HELATIONSHIPS O F T H I BRITISH GUTANA3fARINE FISH FAUNAWITH T
NEIQHROI~ILING
TERRITORIES
........................................
VI. SUMMARY
OF CONCLIJSIONS..
..........................................
VII. ACKNOWLEDGMENTS
.......
.....
VIII. REFERENCES. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
889
APPENDIXI. List of fish species caught off British Guiana, mainly in the trawl, showing
zone in which each species was moat frequently caught, and distribution
in neighhouring territories. .........................................
692
AITF:NI)TX11. Additional notes on cxert'ainlittle-known fish species found off British Guiana 697
T. INTRODUCTION.
THEAREAFISHED
(a) Collection of Data
THIS study of the fishes of the British Guiana coastal shelf, on the Atlantic coast of
South America between latitudes 6"-9" N. and longitudes 57"-60' W., was made
primarily during the trawl survey carried out by the R/V Cape St Mary in these
waters from April 1957-March 1959. It concerns thcrefore mainly the demersal
species of fish. A reference collection of all the fish species caught has been deposited
in the British Museum (Natural History), London, and the period March-October
1960 was spent working on these collections in London.
Off British Guiana the continental shelf averages about 80 miles wide out to the
100 fathom line. and during this two year survey the R/V Cape St Mary explored
this whole area, from about the eight fathoms line to the edge of the shelf, along the
whole 230 mile long coastline. though most of the fishing was done in water less than
40 fathoms decp. A summary dealing with the practical aspects of the results of
this survey has already been published (Mitchell & McConnell, 1959), but this contained very little about the species of fish caught and their biology. This is the first
comprehensive survey of this area ; previously the only trawling in the area comprised
some experimental hauls, mostly in water shallower than that fished by the R/V
Cape St Mary, made in July-August 1944 by Whiteleather and Brown (1945),
and from the ketch Arthur Rogers in ,July 1948-March 1949 (Hepworth, 1949 ;
Graham, 1949).
There is as yet no checklist of the marine fishes of British Guiana, and this present,
paper attempts to help remedy this lack of knowledge of the fish of this large area,
which links up the faunas of the West Indies and Brazil (Figure 1). For adjacent
1 Present nddren\ : Dept. Zoolag:, . British Muserim (Natui nl History). Cromwell Road.
I,ondon. S.W.7.
670
I~OSEMARY 11. LOW&; (MCCONNELL)
[J.L.s.z.
areas the literature that is available is mainly in the form of check lists of species
(Boeseman, 1948 a and b, 1952, 1963, 1956 ; Fowler, 1941 ; Meek & Hildebrand,
1923-28 ; Schultz, 1944, 1949), without any notes on the natural history of the fishes.
Brief field notes are given by Puyo (1949) for species found off French Guiana, and
since this present paper was prepared, IL paper by Durand (1961) has been published
Fro. 1.- Map of are&fiohpd by Cape St Mary trctwl, 1957- 1959.
listing the invertebrates and fishes off French Guiana and their relation with depth
and type of bottom.
The fishes and the corresponding data on which this paper is based were collected :
(1) on five cruises made with R/V Cape St Mar?/ in March, April and September
1958 and in February and March 1959.
(2) from routine examinations of samples of fish from all trips made by the ship
during the second year of survey (March 1958-March 1959).
(3) by analyses of the records of catches kept by the Skipper, W. G. Mitchell,
over the two year period April 1967-March 1959. It is a summary of the results of
these records which has already been published.
XLIV]
THE FISHES O F THE BRITISH GUIAh’A CONTINENTAL SHELF
67 1
During the two-year survey the R/V Cape St Mary made 35 cruises of 4-11 days
duration, on which 1070 stations were fished, and the trawl was towed for a total of
2246 fishing hours. Each station comprised one drag of the trawl, and generally
five two-hour hauls were made each day. Nearly all the fishing was during daylight
hours, as night fishing yielded little. The trawl used was a standard Peter Carey
trawl, 72 f t on the headline, 92 f t on the groundropes, with cod ends of 24 in. mesh
(Mitchell & McConnell, 1959). To ensure regular sampling in the different areas, the
chart of the 230 mile long coastline of British Guiana was divided into seven equal
and parallel-sided boxes, running out from the shore to the edge of the shelf (Figure
1). During the two year period 4 separate surveys were made, each covering the
whole area.
Some of the species caught in the trawl were also caught by local fishing methods ;
in the 2000 yard long pin seines set in an arc on the tidal mudflats ; in the baglike
Chinese seines staked in, and off, the river mouths ; on the hooked longlines (‘ cadells ’)
set in shallow water and in the river estuaries ; or on the hooks used for snapper over
hard ground in the deep water near the edge of the shelf (Allsopp, 1959). The fishes
caught by these local gear have as yet only been considered briefly elsewhere (Lowe
(McConnell)in press), but some catches were examined to see how many of the trawlcaught species were concerned, and to collect information about other stages in the
life-histories of these species. Young stages of some of the trawl-caught species were
also caught entering brackishwater swamps along the open coast a t Onverwagt,
where these species were then grown in brackishwater ponds (Allsopp, 1960).
(b) Physical Characteristics of the Area
The continental shelf off British Guiana slopes very gently to about 50 fathoms on
average about 75 miles from the coast, then there are abrupt drops to the 100 fathom
line on the edge of the shelf, from where the shelf falls away steeply to water over 1000
fathoms deep. The bottom is, on the whole, of soft mud inshore, then progressively
harder mud, sandy-mud, sand, and then becoming increasingly hard and calcareous
and rough towards the edge of the shelf, where there are many old coral blocks which
impede trawling (Nota, 1958 and data from R/V Cape St Mary echo soundings).
These coral blocks evidently represent an old, ice age, shoreline reef (Nota, 1958).
Shells are mixed in the sand and mud in certain areas. The areas of very soft mud
inshore are by no means stable, being moved over the harder mud by the complicated
interaction of tides and currents and water flowing from the many large rivers,
particularly after the water has been well stirred by the strong northeast trade
winds which prevail for much of the year (October-March) in this area. Offshore
the Guiana Current flows in a northwesterly direction a t from 1 to 4 knots ; the
effect of this ocean current is felt outside the 25 fathom line (about 30 miles offshore)
t o the edge of the shelf (Mitchell & McConnell, 1959). Inshore of this the effect of
the tides, profoundly modified by the strength and duration of the wind, may be felt
out to about 30 miles offshore (Whiteleather & Brown, 1945). The tidal rise and
fall is 3 metres a t springs, 1.3 metres a t neaps ; there are tidal mud flats almost all
along the coast, in some places half a mile wide.
Water movements along the coast are further complicated by the seasonal variation in flow from the many large rivers entering the sea along the coast, of which the
largest are the Courantyne River in the southeast, the Berbice, the Demerara (on
which Georgetown is situated), the Essequibo, the Ponieroon, and the Waini River
in the northwest, which lies just southeast of the many-mouthed Orinoco delta. These
rivers also cause great variations in the salinity of the sea along the roast. The
main rains are in May-August and the volume of freshwater entering the sea is
probably greatest from about June-September ; these are also the calmest months
during which the northeast trade wind is not blowing. These trade winds blow for
much of the rest of t h e gear, and are particularly strong in January-March, causing
heavy swells in from t h e Atlantic a t this time which stir up thri mnd along the coast,
078
ROSEMARY H. LOWE (MGCONNELL)
I J.L.S.Z.
From the air, zones of different coloured water, brown inshore, then green, then
blue ocean water, are clearly seen running approximately parallel with the coast. In
the wet seasons the brown water, clear brown from the rivers, and muddy brown where
the mud had been disturbed by wind and water movements, extends further out to
sea, and flotsum from the rivers often collects along the contacts between the zones.
There are remarkably sharp, but shifting, interfaces between these zones of water,
interfaces which fluctuate with the water movements, but on the whole the brown
water tends to lie over the mud and sandy-mud bottoms. The green water (presumably enriched by salts brought down the rivers) may be over mud stretching out over
sand, depending on the season, the amount of freshwater, the amount of mud being
stirred up, tidal movements, and whether wind and currents are reinforcing one
another's effect to strip off and stir up the soft mud. The clear blue ocean water lies
over the hard sand and coral bottoms.
Whiteleather & Brown (1945) further divided the water types off British Guiana
into : (1) clear brown water off the mouths of the larger rivers, estuarine water not
yet mixed with coastal mud ; (2) muddy brown water up to 10 miles offshore, this
fine mud kept constantly on the move by tides and wave action ; (3) muddy green
water ; (4)clear green water, sometimes turbid ; (5) continental blue or blueish
green water ; ( 6 )oceanic brilliant blue seawater.
Thus the ever-shifting water zones do not correspond exactly with bottom type
zones, but there is a general agreement. The way in which the water colour changes
rapidly in onc fixed spot was demonstrated dramatically while fishing in the northwest (Box VI) on 26 February 1959. The dhan buoy was anchored in 22 fathoms a t
8" 28' N. 89" 01' W. and was in light green water until 1300 hours, when a mass of
brown water, separated from tho green by a sharply defined interface, swcpt up on
the tide and travelled on northeast past the dhan ; the water around the dhan remained brown for a day and a half, then became green again during the night.
Since the R/V Cape St Mary survey, shrimp trawlers have started t o operate
from British Guiana, but these arc bringing in fish (as well as shrimps) mainly from
waters off Surinam, immediately Routheast of British Guiana, where, according to
Highman (1959), the bottom is of a somewhat different nature. Although it is mud
out to about 19 fathoms, beyond this is a shell ridge zone from 19-23 fathoms, then
a hard trawlable bottom, of predominantly grey mud and fine shell, extending out
to 40 fathoms, with fine ' moss-like Gorgonids, and deep blue water'.
Thus the factors which may affect fish distribution on the continental shelf off
British Guiana include (1) the type of bottom, (2) the distance from the coast, (3)
the depth, (4) the clarity of the water- and the physical, chemical and planktonic
conditions which affect this, and (5) the salinity of the water.
Some of these environmental characteristics are summarised in Figure 2, in which
the main fish fauna zones are also indicated. The Peter Carey trawl being a bottom
fishing device is, of course, selective of demersal fish, and the distribution of these
fish was found to depend on the type of bottom, rather than on the depth or position
along the coast. However, a number of more pelagic species were also caught,
which tcnded to keep to the clearer water and came and went with the type of water
rather than with the type of bottom, large numbers being caught together with the
demersal fish a t certain times.
Off British Guiana the distance from the coast is linked with the type of bottom,
soft mud inshore becoming progressively hard and sandy offshore and increasingly
calcareous approaching the old coral blocks a t the edge of thc shelf. So species
found close inshore round coral island8 in the West Indies are here only found forty
miles or so out from the coast. On this gradually sloping shelf depth is also linked
with distance from the coast and hence with the type of bottom, the shallow water8
having Soft bottom deposits, and the deep waters being over hard calcareous bottoms.
There is, however, some variation in the depth/bottom type relationship, for the slope
of the shelf i R Rteeper in t h e northwest and here mud is found out to 50 fathoms.
\
674
I. J . L . s . z .
JWYEMARY ir. LOWE (MCCONNELL)
whereas in the southeast the hard coral bottom is encountered in 20 fathoms, thougli
at about the same distance (about 45 miles) from the coast. Clarity and colour
of water is also somewhat linked with the distance from the coast, type of bottom and
depth. The salinity must also increase with distance from the coast, and this factor
evidently shows much seasonal variation, the clear brown water from the rivers
pushing much further out to sea, carrying with it much tell-tale debris, during and
after the heavy rains in June-September, whereas in very dry seasons brackish
water may cxtcnd fifty milcs or so u p tlic rivers.
BOX
m
PI
Y
Li
ri
iij
It
+
61
-in
51 - G O
41- 5 0
31 - 4 0 :
j
N W
KEY
TO
Mud
FIG.
g
!a
v
m
POSITION
PRINCIPAL BOTTOM
Mud R S a n d
ALONG
ii
Iij
COAST
I
S F
DEPOSITS
i7:1
Snnd
Coral
38.--Uiagrarn to show the principal bott.oin deposits at different depths along thc coast from
Box VII (northwest) to Box 1 (sciutheast) (see map, Pig. 1).
Figure in corner of each 5 or 10 fat,liom unit for each Box deriotes moan catch of odiblo
tidl (in kilo per fishing hour) in this unit over tho two-year trawl survcy.
(Note : distances riot to scale as shclf slopes more steeply in IiorthwesL ; to indicate
this the line RB represents a lino parallcl t o the coast about 45 miles offshore.)
A combination of' these many linked factors must control the distribution of the
diff'erent species of fish, and each environmental zone must be considered as a wholcb.
Figure 3 indicates the types of bottom found at various depths in tho different boxes
from northwest to southeast.
Invertebrates Characteristic oj the Area
On the very soft mud, particularly in the northwest, grey five-armed starfish
Astropecten marginatus (var.) were often brought up in thousands if the trawl dug
into the soft miid. Polychaett worm tubes were also brought up from the mud, and
(cb)
XLIV]
THE FISHES OP THE BRITISH GUIANA CONTINEKTAL SHELF
675
the polychaetes Eunice antennata (Savigny) 1818, and Onuphis magna (Andrews),
1891 were collected. Odd Sipunculids (Sipunculus multisulcatus Fischer) also came
up from the mud, and sea pansies Renilla reniforwk (Pallas). On the harder sandy
mud, the best trawling ground, basket stars Astrophyton muricatum (Link) were
abundant, also five-armed and nine-armed starfish Luidia clathrata (Say) and L .
alternata Say, and featherstars TropiowLetra carinata (Lamarck). Large areas of
multicoloured stringlike hydroids were found; mainly a t the interface of mud arid
sand. Various penaeid shrimps were fairly common ; species in the area included
Penaew aztecus Ives, P . schmitti Burkenroad, P. brasiliensis Latreille, P. setiferus
(Linn.) and Xiphopenaeus kroyeri (Heller). Squilla empusa (Say) was also very common in muddy and sandymud areas. Very few live molluscs were taken, but the
squid Boryteuthis brasiliensis (Blainville) was sometimes caught from clearer waters.
Few invertebrates were brought up from the pure sand, but in the deeper water
the harder sand and coral bottoms yielded numerous large sponges, dead corals,
seafans (gorgonids), seaperis (pennatulids), and an associated fauna of Crustacea,
including occasional squat lobsters Scyllarides aequinoctialis (Lund), spiny lobsters,
and various species of highly coloured swimming and other crabs (including Portunus
spinicarpus (Stimpson), P. spinimanus Latreille, Callinectes ornatus Ordway, Lupelln
forceps (Fabricius), Hepatus sp., Dromia erythropus ( G . Edwards), and others). The
boxcrabs Calappa flammea (Herbst)and C. sulcata (Rathbun) were also taken ; also
various spider crabs. Various echirioids and ophiuroids were also caught in this zone
(Mitchell & McConnell, 1959, appendix 3).
The brown rhizostome jelly fish Stomolophus meleagris (L. Agassiz) was a common
element in the trash in water less than 40 metres deep, in the brown water, and was
caught a t all times of year, though it varied in abundance ; these could be seen a t
the surface and a t times appeared to be moving to the southeast against the currents.
The invertebrates collected during the survey were very kindly identified a t the
British Museum (Natural History) and remain in their collections. Most of the invertebrates figured by Durand (1961) from French Guiana were occasionally seen off’
British Guiana.
(d) Fish Fauna zones
The fish faunas can best be considered separately from the four zones indicated
in Figure 2, namely :
Zone I , the ‘ brown fish ’ zone, inshore of the ten fathom line over very soft mud ;
this was inshore of the R/V Cape St Mary’s fishing area and has as yet becn little
studied. Here live species which rarely move out into the area fished by the Cape
St Mary, together with the young and adults of some of the species caught in the
trawl. The zone I fishes are known from local gear, pinseines, Chinese seines and
longlines, and from the Ketch Arthur Rogers trawl which was a t times fished in water
5-10 fathoms deep. The sea is here much diluted with rivcr water and subject to
great changes in salinity, and tides and winds keep the mud in constant suspension
except just off the river mouths where the water may be clear if much freshwater is
coming down to sea. Numerous species of catfish (nematognaths),rays, sciaenids and
fish such as toadfishes (Batrachoididae) abound. Many of the fish are brown in
colour and very soft-bodied.
Zone I I comprised the ‘golden fish ’ community of demersal fish found over mud
and sandy-mud bottoms, mainly in the brown water. Owing to the nature of the
bottom this zone extended out to about 20 fathoms in the southeast but to about 30
fathoms in the northwest. This was the most inshore zone fished by the R/V Cape
St Mary, and yielded most fish, so it was studied more comprehensively than the
other zones. Sciaenids predominated in this zone making up over 75 per cent of the
catch (by weight), and various species of catfish, carangids and grunts were abundant.
Many of the specics in this zone had a golden colour, and, probably in association
676
ROSEMARY ii. LOWE (MCCONNELL)
[J.L.s.z.
with the muddy opaque water, many of the species produced loud noises, croaking and
grunting loudly as the catch was landed on deck. Many of these species are also
found in the estuaries of the rivers, particularly in the rainy seasons.
Zone III,the ' silver fish ' community of the greener water, over sandy-mud and
sand bottoms contained more pelagic fish, many probably caught in the trawl as it
was shot or hauled. Carangids predominated, and Spanish mackerel, barracuda.
pomfret and herrings were abundant a t times. The presence of these fish appeared
to depend more on the water type than the type of bottom, and they tended to be
found closer inshore when the clearer greener water came further inshore, and were
then caught with the sciaenids (in zone TI) ; but they were also particulurly abundant
in 2 0 4 0 fathoms, for examplc in Boxes I V and V (see Figure 1). The pure sand
bottoms were almost barren of bottom-dwelling fish, though sandy-mud in these
clearer waters carried some lizard fish, grunts and flatfish.
ZoiLe I V comprised the ' red fish ' community ovrr hard coral arid sand bottoms
in deeper water (over 30-40 fathoms in Box 111, and over 20 fathoms in Box I in the
southeast.) Various species of pink and red snapper were caught here, mainly small
species, and the highly coloured fish characteristic of coral areas. The coral was.
however, dead a t this depth. The majority of the species caught in this zone were
bright red, many with spiny excrescenccs.
The species present in each zone are considered in a later section.
11. FISHSPECIES
CAUGHT
(u)Species and Abundance of Trawl-caught Fish
The species of fish found in these four main zones are listed in Appendix I. This
Appendix shows that some 72 families were represented in the area (64 in the trawl
catches), by about 213 species of fish (about 190 in the trawl). The families with the
most numerous species were the Sciaenidae (about 21 species), Carangidae (about 14
species), the nematognaths (about 12 species2), Yomadasyidae (about 13 species)
and Lutjanidae (about 7 species). The sciaenids were predominant, both in numbers
of species and in abundance of individuals, in zone I1 ; the carangids and pomadasyids
in zone 111, where the former lived a pelagic life and the latter on the bottom ; the
lutjanids predominated offshore in zone 1V over the hard calcareous bottoms.
The families next most rich in species were the requiem sharks Carcharhinidac
(about 10 species) which tended to haunt the tidal contours between the brown and
green water. Serranids were represented by about eight species, mostly in the offshore
zones. Of the families represented by five or six species in the trawl, the Ariidae
catfish were characteristically inshore and estuarine fish, and numerous other species
of catfish were caught further inshore and in the estuaries by the local fishing methods
(Lowe (McConnell),in press). The chaetodontids (five species) and monacanthids
(five species) were only present in the most offshore zone I V ; the tetraodontids
(about four species) and ogcocephalids (three or four species) were represented by
different species in the different zones.
Aoundance.-'l'hc trawl catch was far and away greatest in zone II where thc
sciaenids predominated (Table I). Over the whole two-year period the sciaenids
made up over 75 per cent of the weight of fish landed in the trawl (Mitchell & McConnell, 1959). I n zone I1 the average catch over the whole survey was over 150
kilo per fishing hour of commercial species.g I n zone I11 the catches fcll off rapidly
to less than 70 kilo per fishing hour ; catches were very poor over pure sand. The
catches in the offshore zone I V were poorest, and negligible from a commercial point
Another 8 species from coastal waters have since been recorded-see Appendix I, Addendum.
These figures exclude trash dumped overboard ; this was also more plentiful in zones I and
11. An allowance was made for small fish (many sciaenids) in this trash when comparing trawl
ratehes off British Guiana and West Africa by Lowe t Longhurst (1961. Nature, 192 :620-622).
For details of Cape St Mary catches off' West Africa see references in Longhurst (1961, Nature,
192 : 622) and 8alzen, E.A. (1957, J . Cons. Ziat. Explor. Mar., 23 ( 1 ) : 72-83).
I.-i% comparison of trawl catches at different depths and in different boxes
along the coast, from eoutheast (Box I) to northwest (Box VII) (SW Figure 1).
TABLE
Figure8 givon u.ro the iiioan oatdl in kilos 1)er fitlhirig hour of all odihlt! q x c i i ~
tnkull iir
the R/V ('u.pe 8 t M a q ' h trawl over the two-year SIII'VAY, A p d I967 --Mnrrh lW!l
7
Depth
(fathoms)
NM.
VTI
Approximate
tish tauiin
ZOIitl
t
8-10
11-15
L 6-20
224
21-25
I42
2630
I65
111
3 1-40
41-5U
1
0
...
...
...
!)
...
...
...
...
\I
...
0
1)
...
...
51 -60
b l il)
7 1 -I
...
1V
...
...
of' view ; many of the species \\ere very sinall hpiny Mi a i d large snappcr were v e r ~ .
rarely caught in the traml.
The abundance of fish over different types of bottom ih shown in Figtir~3 : triiitl
and sandy-mud yielded far more fish than sand or coral bottoms.
(b) The CornmorLer k'ish Apwies PresetLt in the BijjererLt Zor~es
The zone in which each species is most commonly caught IS sliou n
in
hppentli \ I .
Zone J
The species caught by the R/V Cape St X U ~ ~ trawl
J ' A in k i ~ htliiLii ttw f'atlioriis.
or only in numbers when strong onshore ninds \\ere stirring u p tht. rnrrd and tht, fish
had moved offshore. included Chinese butterfish Lonchurus lutreolaks, I)ashav
Cynoscion acoupa, snook CeTLtroyomus eil~iferusand C. citzdrc.irturli.~.cat fish A r w s
(In these same catchw
,Fssus and A. parkeri, and toadfisli Batrachoidw .s%~ritia?r~rti.~~s.
the absencc, o f cararigids, niupterus rhombeus, C'onodon ,tohilis. and harracwln ant1
Scomberomorus was most marked). Stingrays llasyatis yuttutus and lirotryyott
rnicrophthalmi~~t~,
the butterfly ray Ggnmura tnicrura. the electric ray &Vat1.irie brusiliensis, and the butterfish Nebris microps, were also comnioner ill catrhtks from thew
very inshore waters. The yoimg, premature. stages of many fish were also c*aught
here, for example a ' nursery ground ' struck on 6 September 195s at I 1 fathoms in
Box 111 yielded small Nicropoyon furnieri (l6/18 urn. long), Getr!jatreirms l r ~ t ~ and
us
Conodon nobilis of 12/14 cm., (7haetodiptrrus ,faber X cni.. and small ('yrtosciott
jnnzaicensis, B . bugre and i l r i u s hi).
The Ketch Arthur Rogers, fishing these inaliorc* \z Ltters (five-tvii fathoms) in I ! M f i .
took many small catfish, including kukwarri Hezanernatichthys rugispinis, mud cuirass
Selenaspis herzbergii, and banjoman Aspredo uspredo, also various clupeids and engraulids, not recorded from the Cape S t Mary trawl (Graham, 1949). The catfish spines
got entangled in the trawl, delaying operations, so they \sere not a welcome catch.
JOURN. LT". ROC.--ZOOLOQ\ , w r , . SI,TV.
10
67 8
ROSEMARY H. LOWE (MCCONNELL)
[J.L.s.z.
The hcrrings taken included a Pellonn sp. (' shad '), and sciaenids included Cynoscion
moupa and Plagioscion sp. ; a lizard fish Synodus sp. was said to be abundant, and
the spotted sole Apionichthys durnerili fairly abundant ; a copper-coloured Oligoplites sp. was also taken from these more inshore waters. A reference collection of
species caught during this 1948 survey was sent t o the British Museum (Natural
History) by Miss V. E. Graham (Graham, lY49), and the species list was published as
Appendix 2 in Mitchell & McConnell, 1959, but subsequent work on fishes from this
area has thrown some doubt onto some of these identifications.
Other species not caught in the trawl but common in these inshore waters, as
shown by catches in local gear-Chinese and pin seines, longlines and a t Onverwagt
brackishwater fish culture station-include Tarpon atlanticus, Elop9 saurus, the
mullets Mugil liza (brasiliewis of authors, known locally as ' Querinian '), Mugil
curenaa and M . incilis, various catfishes such as kakinette Pseudauchenipterus nodosus
and the less common highwaterman HypophthalwLvn eden,tatus and many others.
pargcc Lobotes surinamensis, the large cownose ray Iihinoptera sp., and the foureyes
Anableps microlepis and A . anableps. Other catfishes and sciaenids from these gear
await further study. A round-the-year study of the species caught in these gear
has yet to be made, hut a preliminary account is given by Lowe (McConnell) (in
press).
Catches in all these inshore gear appeared to be highest during the rains, June to
September, when many of the sciaenids and other species from zone I1 moved inshore to zone I.
Among the sciaenids it was noticoabla that certain species werc taken in the
cstuarics (in Chinese seines) but were never caught entering brackish waters along the
open coast a t Onverwagt ; these included young Macrodon ancylodon, Nebris micropy,
Lonchurus lanceolatus, Isopisthus sp. Whereas other species, including Micropogon furnieri, Cynoscion acoupa and Oynoscion steindachneri, were taken entering
brackishwater swamps at Onverwagt but wcre not caught in the estuaries.
Zone I f
In zone I1 the most abundant species wrc? thc three sciaenids, the croaker Micropogon furnieri, which made up over 40 per cent of the weight of fish landed in the
trawl during the survey, bangamary Macrodon ancylodon (19 per cent of the weight,
of the total catch), and the seatrout Cynoscion virescens (11per cent of the weight of
the total catch) (Mitchell &, McConnell, 1959). Oynoscion jamaicensis (table bashaw)
and Larimus breviceps (rockhead bashaw) and two or three StelZifer species (also
called rockhead bashaw), and fsopisthus sp. (also called table bashaw) were very
common in the trash. Of the other sciaenids, numbers of butterfish Nebris microps
were occasionally caught, and among those taken one or two a t a time, mixed in
with the commoner species, Corvuln sanctae-luciae and Paralonchurus sp. came from
rather inshore waters, and Umbrina gracilicirrhus and Menticirrhus "1). from rather
deeper water over harder sandy-mud.
The fishes most commonly caught together with the sciaenids in this zone, between
8-20 fathoms were : requiem sharks, the dogfish Mustelus nov. sp., (Springer &
Lowe (in press)) and bonnet shark LSphyrna tiburo ; catfish including Bagre bagre
and B. marina, the former generally more inshore than the latter, and Arius species
of the strictir~ssis-parmocassis-grandicnssi~~
group (for which evidence was collected
suggesting that thcsc arc really one polymorphic species4); the cutlass fish Trichiurus lepturzcs, which was very abundant and also common i l l the estuaries where
the young are caught in Chinese seines ; thc carangids Vomer setapimis (moonfish)
and the bumper C h ~ r o s c o m h rcht.yswus
~
; the pomadasyids annafolk ' A ' Genyntretnus lrdeus mtl the less frequently caught annufolk ' B ' Conodon nobilis. The
following species were also frequently caught : two spccies of herrings, I f areiquln
' See Appendix I1
ursvj
T H E I'ISf-IKh VJ!' T H E HKITINH (:VfAiV i ('OXTlh {;\'I' 11, SHB3.4'
679
pemacolae and Ilisha argentata (and less often in this zonc thv thrcad herring O p k
thonema oglirivm), the Spanish mackerel Scomberomorus maculutus, the small barracuda
,Sphgraena gwdmncho, carangids such as the silver Oliyoplites snliens (locally called
mackerel), and (7aranx spp. and Hemicarctnx sp. When the water was rathclr clear
these silver fish mere iiumeroiis in thc catches, particularly C'omer, the barracuda,
and also the pomfret Peprikcs paru and seapatwa Diapterus rhombeus. The jackass
Ghuetodipterusfaber was common in the hydroid growths a t the interface of mud and
sand (many small ones carrying ectoparmitic isopods, Nerocib armuta Dana) ; one
haul in November 1957 took 2640 111s. of Chadodipterw, from 12 fathoms over mild
in Box IV.
Species caught t w o or three a t a time included the electric rays Nurcine brusilierL&
and Diplobatis pictus, thc: stingrays Dasyatis giittatus (D. geipkesi on one occasion
only) and Urotrygon microphthalmum, the butterfly ray Gy~nnuranzicrura, various
Haemulou species, the threadfin Polynernus virginicuu, the striped puffer Colomesus
psittacus (the young of which are seen in the estuaries at high tide), thr. porcupine fish
Ghilomycterus spinosux, and the batfish ' Ogcocephalus ti uncutu,v 7.s Large sawfish
of two species, Pristis pectinatus and perotteti, were occasionally caught, particii larly
near the estuariw : they w r v taken in every month of the year.
Zone I11
Zone 111 was much less w ell-defiiied and merged u ith zones I1 and 1V as the bottom
was relatively barren and the catch mainly of the more pelagic, silver, fish, which
moved with the water types into zones I1 and IV. Carangids predominated in zone
111, about nine species being found here (Appendix I),of which the commonest en^
Vomer setapinnis and Chloroseombrus chrysurus, Hemicurunz amblyrhpchus and
Oligoplites salieiw., with somt' Caraiix hippoe and C . crysos ; a few Selene uomer were
caught in clearer waters (this is the common species around Trinidad). Barracudii
all apparently Sphyraenu guachatcho, and the mackerels #comberomorus muculatu.\
and cavalla (mvalla in the clearer more offshore waters ; no S. regalis were secn)
herrings, particularly Opisthonenaa oglinuin but also Hareiiyula pensamlap and Ilisha
argentata, and pomfret Peprilus paru, were also very common here. (See ' MackerelBarracuda area ' shown in Figure 1). The slenderer seapatwa Eucinostomw gub
and argenteus, the cobia or ' cod ' Rachycpntrorb cfinadus and the ' Salmon ' Pom,utovn7~.\
saltatrix, were also caught, in small numbers a t times. Groutid fish were few, but
included the flatfish Cyclopwtta jMwiata and C. chittendeni, and, further out, thc
flying gurnard Dactylopteruv volitans, a cuskeel Lepophidium brevibarbe, and occasional
Porichthys porosissimus. Sciaenids were rare in this zone, but Mrnticirrhw spp.
(' deepwater butterfish ') werc caught futher offshow than any o t h r clpecitxs of
Sciaenid except &pees 1nrrc~eolnztw.sfound in zone I]'.
ZotLe
IV
In Zonr 1V, in the oBshore haters over hard sand and coral, uliere large spongrh
and seafans grew in many places, the fish in the trau 1 were mostly small. very spiny
and many were bright red in colour. Eques lanceolatus was the only sciilenid to bv
taken in this zone, a feu a t a time. This zone is fished by hooks on handlinrs from
the snapper schooners, which catch large red snapper, mainly Lutjanus ayu. also
large grouper, and occasional dolphin Coryphaeim hippurus, and other large fish ;
the species caught in this way off British Guiana have not yet been studied systematically or listed. Except for one haul in which over 300 kilo of large red snapper were
taken, the trawl rarely caught large snapper (and on this occasion a dhan buoy was
put down but further fishing in the area yielded no more snapper) ; i t seemed thc
snapper were too high off the bottom to be caught in the trawl, and the ground was
very rough, causing much damage to trawls in this zone. Small species of snapper.
See Appendix 11.
680
HOYEMARY H
LOWE (Mc(’OKh’EL1,)
[J.L.s.z.
a few a t a time, were however taken in the trawl, mainly Lutjanus synayris, Ocyurus
near chrysurus, Rhombopliteu aurorubcm, and several others not yet identified ; all
wcrc pink or red in colour.
On the sbhole, catches in zone 1V produced numerous species but relatively few
iridividuals of any one species. The other red fish caught included : three speaies
of holocentrid (Holocentrus ascensionis was commonest, H . bullisi taken a t several
stations, and Myripristis jacobus from one station) ; the bigeye Priacanthus arenatus ;
a t least three species of red mullet (A!Iullus sp. Upmeus rnucdatus and W . parvus,
the latter with its stripcd yellow and black caudal fin) ; four gurnards (Yrionotus
alipionis, two other species of Prionotus and Bellator wditaris) ; about three scorpaenids (including the huge-eyed Scorpaenn agassizii) ; and some brilliant red batbli
resembling Ogcocephalus vespertilio. Other fish found here included tlie blacknose
shark Carcharhinus acronotus, the guitarfish Rhinohatos percellens, five species of
lizard fish (Trachinocephalus myops, three Syuodus spp. and one Saurida s p ) , three
flatfish (Bothus sp., Syucium papillosum and a vividly striped Gymnachirus sp., a n
odd Ilippocumpus longirostris, Fistularia yetimha, two interesting stromateids (a
Cubiceps h p . and a Psenes-like fish). Carangidae in this zone were represented by the
goggle-eyed scad S‘elar crumenophthultnus, and the small round and rough scads
Decapterus punctatus and macarellus (less common) and Trachurus lathami. Three
species of Diplectrum and Prionodes phopbe, and a t least six species of grunt were taken
(including three Bathystoma specicLs, two Iiaemulon and Anistotremus virginicus).
Very few sparids, all one Calanzus sp., were taken on the whole survey. Caulolatilus
{pppyi, a species previously known only from Trinidad was taken on two occasions,
several of a similar size (20/23 cm. long) a t a time ; one Antigonia capros was caught
on one of these occasions. At least six chaetodontids were caught, including P o w canthus arcuatus and paru, taken together, and an odd Holacanthus tricolor. Catches
also included a few specimens of Acunthurus hepatus, Balistes carolinus and B. vetuh,
Sphoeroides spp. and one Antenwurius scaber. Labrids and scarids were each represented by a few individuals of two species. Five species of monacanthids were
caught ( M . ciliatus less commonly than M . hispidus). Cowfish Lactophrys tricornis
of varying sizes were fairly common, and the round batfish Iialieutichthys sp. was
taken a t two stations.
(c) Comparison of N u d e r s of Species arLd Pish in the Dijjerent Zones
As the zones overlapped so much it is difficult to equate the number of species
living in each zone. However, summarising from Appendix I, it seems that from
zone I about 62 species were recorded (and further work in this zone is sure to reveal
many more) ; from zone I1 came 80 species : from zone 111 43 species ; and from zone
I V about 80 species.
The fish were, however, much more ahundant in zone I1 than zone ZV, as has been
brought out in Table I and Figure 3, arid on the whole (with the exception of the larger
snapper and grouper) larger. Owing to the irregular bottom in zone I V this could
not be trawled as effectively as zone 11, where the hard mud made an ideal trawling
ground, but even taking this into account, it seems that zone I1 is intensely more
productive of bottom fish than zone TV. This is to be expected as zone I1 is constantly enriched with salts brought in by the rivers, and kept in constant circulation
by the complicated and continual water movements in this area. The green water
zone of plankton growth immediately offshore of this area, so clearly seen from the
air, shows the enriching effect of these nutrients. The bottom in zone IV, on the other
hand, quite probably lies below a thernloclinc ; no sea temperature data are available
for the west side of tlie Atlantic off British Guiana, but on the east sidc of the Atlantic
in these latitudcs a thermocline is known to extend between 10/15 and 15/% fathoms
(Watts, 1958).
It is interesting that so many of the zone I and I1 fish are softbodied species from
t l w early part of Appendix 1.i n \\hidl thc specks arc arranged in ‘ systematic order ’,
YLTV]
THE I”1SHES OF THE HItlTXSH GUIANA CONTINXNTAL SHEIAF
ti8 1
whereas the zone IV fish are predominantly spiny fish from the latter part of the
systematic table.
111. SEASONAL
BIOLOQY
Seasonal Changes in Zone I1
Sciaenids and other spccies characteristic of zone 11 were caught all the year
round and there were no regular seasonal fluctuations in numbers (Mitchell & McConnell, 1959). There mere, however, some local movements of the fish. The analyses
of catches of the main commercial species already described (Mitchell & McConnell,
1959) showed that there is a general tendency for fish of the sciaenid zone (zone 11)
to move inshore during June-August, the months of the main rains when most water
is discharging down the rivers to the sea ; these are also the calm months when the
northeast trade winds are not blowing. Many of the fish spawn a t this time, and
catches in the estuaries in Chinese seines, and on the tidal mudflats in pin seines, are
particularly heavy a t this time of year. The R / V Cape St Mary’s skipper commenting on the profound effect of heavy rains in August 1959 noted ‘ dark water and
flotsum being carried far out to sea (in an easterly direction disturbing the normal
tidal southeast/northwest currcnts), and catches near the shore after this were very
good ; it seems that the rains attract the fish ashore and the catches increase, then
the “ dogfish ” (Carchnrhinusspp.) move in after the fish and the good fish disappear’.
During much of the rest of the year the tradewinds blow onshore, stirring up the
mud, and the fish tend to move offshore. The northeast trades are particularly
strong in January-March, and the longer and stronger they blow, the more the fish
move offshore, and catches are then particularly good in the northwest, the area
where they can move into deeper water and still remain over mud.
Particularly good examples of the movements of fish offshore during heavy
tradewinds were witnessed in February 1959 in Box I, when the trawl took species
which a t other times live inshore of the Cape St Mary’s inner limit (eight fathoms),
such as Lonchurus lanceolatus, Arius jissus, and Batrachoides surinamensis, with
many Narcine brasiliensis and small stingrays such as Urotrygon microphthulmum
(see page 677). Also in January and February 1959 in the northwest district (Box
VI) the fish were found to have moved out between two visits to the area ; this
appeared to be related to a change in the bottom, as quantities of very soft ‘ sling ’
mud had movcd out over the harder mud which had produced good yields of fish on
the earlier visit, and the fish had then moved out further.These same good fishing
grounds in 20-25 fathoms in the northwest proved barren when visited in J u n e J u l y
1958, and there was evidence from catches in more inshore gear (pin seines and Chinese
seines) that the fish had moved inshore. However, the previous year, June 1957,
catches in slightly shallower water (17-19 fathoms) of these same northwest grounds
had been good, which suggests either that the fish may not move very far inshore,
or that, there is some variation in seasonal movements from year to year (probably
depending on the advent of the rains.)
Other local movements of the fish appeared to be caused by the appearance of
numerous earcharinid sharks on the good fishing grounds. Although some were
present throughout the year, these appeared in great numbers during the rains when
the rivers were carrying much debris down t o sea, and were particularly abundant from
June-September when they haunted the tidal contours. Not only did they damage the
trawl, but they appeared to chase the other fish from the area. On many occasions
good hauls of the usual sciaenid species were made for several hauls, then packs of
sharks converged on the ship-perhaps attracted by the smell of fish in the trawland all the other fish abruptly disappeared, the following hauls being blank. It
proved necessary to move a mile or so for a further good haul to be made before the
sharks reappeared and the process was repeated, so good fishing became very patchy.
Up to 32 sharks were taken in one haul of the net, hut most of the damage to t’henet
((1)
6Wz'
I.J.L.s.~
LLOSEMANY 11. LOIVV; ( ~ l c ~ Y ) N X M ~ L )
was done from oittside. the sharks tearing the gilled fish out of the trawl as i t was hauled
inboard.
(b) H e r n o i d Cliarcge.r i n Gonad States of the Fish
The gonad states of the species of fish most commonly caught in the trawl were
examined throughout the year March 1958-March 1959, with the exception of
November and December when the ship was out of action. Larger numbers of fish
were examined in the months when tht. biologist accompanied thc ship, namely
February, March. April and September ; data for the other months were from much
smaller samples, of thr commonest specie8 only. examined after the ship returned
from each cruise.
TABLE
2.- 'I'tw sr-:a~oiial
occiirreiw of ripe fish in the trawl.
x denotes prewnce of ripe fish, 2 most of fist1 caught ripe, ( x ) ripening fish, 0 no
ripn fish in Rctmples examined. - species not prrsmt' in Ramplw kept for gonad
examination.)
(
Slw
h s
-\.lirro~ogonPcplLieri . . . .
ilfacrodonancylodon . . . .
Cpwscion wireirescen.9 . . . .
('ynoacion jumnicennis . .
Larinkus breair
!hlaroscombma c h r y a m t ~
(:haetodiptenur faher. . . .
Sph,yrae)ia guachamho . .
~~comberomor11.8
maculntm
'rrichiuffb8 kpturua . . . .
Peprilun paru .
Ragre marina. . . . . .
Hnrengulapenaacolae. . .
Opisthonema ogtininlrm.. . . .
Y
i
:
":
,
'
Y
X
.
-.
Y
(Y)
x
( ;.:)
(x)
x
-
~'
v
.
.
,.'
Y
--
-
(Y ) (X)
X
--
(
...
~~_-2
~.
-. -
-
-_
.,
i
i
.. . . . .
~
---i
.Ian. Fob. $far. Apr. May .Tune July Aug. Sep. Oat. Nov. Dec.
Y
:i
x
x
x
x
x
v
x
x No fishing
0
x
(i
0
0
0
0
x
0
x .,
,
'./
Y
x
0
0
- -x
Y
(:onoilon nobilis
L'07ner nstapitbnis
Month
. -.
,~
' -
..
x
x
(X)
-
x
-
(Y
- -
>.'
(X)
x
-_
--
.
r:
~
.
-.
->:
x
Y
'X
,..'
(X)
_
-
-
x
-
-
-.
.
x
X
' -
.~
__
-
Y
--
~.
(X)
(X)
..
-
-
--0
(X)
j ,
A
x
>c
- x
x
x
X
-
~x
~- x
(x)
.
x
. ~.
~.
(X)
.,
x
,,
,,
x
x
x
x
x
~-
x
~-
(X)
. -
,.
,>
..
., ..
,.
,.
,,
,,
1,
-
....
9.
,.
,,
..
..
*,
.,
7,
1.
.,
..
,,
9.
..
..
.I
..
The months in which ripe fish of various species were recorded are summarized
in Table 2 . This shows that ripe fish were present in samples examined at all times
of year in the croaker Aficropogon .furnit+, the table bashaw Cynoscion jamaicensis,
the rockhead bashaw Larimu.9 breviceps, the sea patwn Diapterus rhombeus, annafolk
Genyatrernus luteus, and the moonfish Vorner setapinnis. I n some other species, such
its L'onodon nobilin, Para1onchuru.q sp., -Venticirrhw sp., and Bagre m r i n a , ripe fish
were fonnd whenever these species appeared in the catches, and in Chaetodipterun
,[tiher whenever the fish caught were large enough to be mature.
In the other common sciaenida, on the other hand, bangarnary Macrodon ancylodo?i w n d ' seatrout ' Cymscion virescens, ripe fish, or any fish with active gonads, were
rarcly caught in the trawl, and were only taken when the trawl was fished very close
inshore. as it was in the southeast in August-October 1958 and February 1959.
&\rnongfish of these species caught in t h e same months in the northwest, in deeper
water, no active gonads could be found. It seems therefore that a ' place effect *
liere obwcures any seasonal changes in breeding, that the fish when ripe must congregate in certain places, evidently moving inshore into shallower water, and unless these
particular places are fished by the trawl ripe fish are not caught. Until samples
round the year arr available from such places it is not possible to determine whether
there are marked seasonal variations in gonad ar.tlivity. though t he l a r p niimherx
YLIV j
THE FISHES OF THE BRITISH GUIANA CONTINENTAL SHELF
083
caught offshore with inactive gonads suggests that spawning is by no means continuous
or prolonged in the mature fish of these species,
Extreme examples of this ‘ place effect ’ were seen in the butterfish Nebris microp8
and the table bashaw Isopisthus sp., as no fish of these species with active gonads
were ever found in the trawl ; the young of both these species are caught in Chinese
seines, and i t seems that these may move into estuaries t o spawn.
Among the less abundant species, which were only examined when the biologist
was aboard (February, March, April and September), ripe and ripening fish seemed to be
more abundant in September than in March ; this included the bumper Chloroscombrus chrysurus, barracuda Sphyraena guachancho, the mackerel Scomberomorus
mculatus, and the herrings Harenguk pensacolae and Opisthonema oglinum. September comes after a period of good feeding during the rains for most fish ; it also
corresponds with the end of the summer for the more northern species. The Skipper’s
records state that ‘ sardines ’ and large mackerel were particularly abundant in the
southeast in very shallow water in August, and that the large mackerel (up t o about
three kilo weight) then contained eggs ; further north off the U.S.A. Scomberomorun
spawn ‘ as late as September ’ (Breder, 1948).
Small fish of many species were, however, taken both in February and September
in the trawl (and during the rains June-September in the estuaries) ; 5 cm. long
Chloroscombrus chrysurus and Yeprilus paru were caught in February, together
with small Macrodon, B. bagre, and Achirus (in 12 fathoms in Box 11),and, as mentioned above (page 667) a ‘ nursery ground ’ fished in February contained small fish
of many species.
Off French Guiana Puyo (1949) reports that many species, (including Macrodon)
spawn in the main rains June-August, and in British Guiana many young fish are
caught in Chinese seines in the estuaries at this time. This may, however, be partly
due to the greater attraction to the estuaries when more freshwater is coming down
the rivers, rather than a seasonal fluctuation in numbers of fish spawning. Young
croaker, Micropogon furnieri, were taken all the year round a t Onverwagt, entering
brackish water from the sea; this species was never taken from the estuaries.
Possibly the estuarine fish have better defined seasonal breeding than fish of the open
coast such as Micropogon.
Large ova were found in Arius fissus in February, and in Bagre marina (16 mm.
diameter ova) in both March and September. Males of these species are said
to practice oral incubation of the eggs (Puyo 1949). Counts of ripe ova in three
B. marina showed that only about 20 eggs are laid a t a time.
Thus in this area there seems to be great diversity in spawning rhythms, some
common species, (belonging t o a t least four families, Sciaenidae, Gerridae, Pomadasyidae and Carangidae), e.g. the croaker, seem to be capable of spawning at any time of
year. Others, including mcmbers of the same families, such as the common sciaenids
Macrodon ancylodon and Clynoscion virescens, appear to have much more restricted
breeding seasons.
(c) The Occurrence of Young in Viviparous and Ovoviviparous Species
Most of the Elasmobranchs caught by the R/V Cape St Mary are viviparous
or ovoviviparous. These were only examined in February, March, April and September, when the biologist was a t sea. I n September the only fish seen carrying young
were a Carcharhinus obscurus, containing three young 22 cm. long, and a Rhinobatos
percellens female (48 cm. long) carrying two young still with yolk sacs (another R.
percellens caught in April had three embryos 65 mm. long). I n February-April
many species were found carrying young ; these included many Mustelus nov. sp.,
Carcharhinus maculipinnis, other requiem sharks, Sphryna tiburo and S. tudes
(diplanu), Pristis pectinatus, Rhinobatos percellens, and Urotrygon microphthalmum ;
Narcine brasiliensis were found containing long yolk strings, and a Qmnura micrum
(iH4
ItOSERlhKY 11. J,C)\VE
( MaCOSNF,LIJ)
I J.L.S.7..
had long fi1ament.s (trophonemata) in the ovidnct, in a condition suggesting that yonng
had just been released.
stingrays (previously
Of particular interests were. thc (J.rotrygo)L ,)ri;i~rophthal~?,?~~~.
lamvn only from the t,ype collected off the Amazon), as in February nearly all the
females caught contained young, three at; a time, about ready to be born, suggesting
that there is a definite seasonal breeding in this species, (see Appendix 11, page 698).
The Mustelus nov. sp. embryos from different females (generally four in each) were
also all of a size. ready tjo Ire horn. tnilfirst. in March, suggesting scrasonal breeding
in this species too.
Thc young S. tudes, four all about 28 cm. long w,nd nearly ready for birth, were from
ti 107 cni. female. Both thcse, and the embryos of Carcharhinu,P maculipinnis (four
of 19 em. still with extrrnal gills, from a 116 cm. female) appear to be the first records
of embryos for these species ; thc mat,crial is in the British Museum for further study.
A female sawfish Pristis pactin.ntus, 3.1 metres long, caught in 11 fathoms in Box
IT1 in March 1958, contained eleven young 42 cm. total length (30 cm. body and
12 rm. saw) still with about R pint of yolk in the 8wr attached to each embryo.
I V. I~ooouRELATIONSHIPS
From the general picturc which ('merges from the analyses of stomach contentb
of the different species, the principal sources of fish food in the main trawling areas off
British Gniana appear to kw :
( I ) inverteln-atw in the bottom mud, ptrticularly polychaetes, hiit also odd
siprtnculids, small crabs, hermit crabs and molluscs ;
( 2 ) cnistacen over the bottom mud, particularly various species of penaeid shrimp
( s w page 675), Squilla empusa, and crabs ;
(3) the more pelagic small white shrimps (family Sergestidae) of the clearer waters :
(4)fish.
(1) Invertebrates from the bottom mud were found in the following species :
the croakers Micropogon furniwi and Paralonchurus sp. (polychaete worms), the
grnnts Gengatremun Zuteus (mixed invcxrtebrates) and Conodon nobilis (molluscs),
batfish Ogcocrphalus ' truncatus ' (tiny molluscs and crabs), the catfishes Bagre
marina and bagre, Arius parkmi and '4. jissus (all rather general feeders). The
Reapatwa Diapterus rhombeus from the shape of its extensible mouth would appear to
probe in the bottom and intestines contained sand or mud and worm remains.
*Varcinr brasiliensis contained polyrhactes, Sipwmclus and small brown eels from
trhebottom mud.
( 2 ) Crnstacea abovt. t h e bothom, mainly penacicl shrimps and Squilla, were
found in the Ktomachs o f : the bonnet shark Sphyrna tiburo, stringrays Dasyatis
{liittntus and Urotnygon microphthnhnum, butterfly ray Q m n u r a rnicrura, catfishes
Bagre m a r i m and B. hagre, Arius of the stricticas,cis-purmocassis-grandicassis
group
(which also contained small round leucosid crabs), the threadfin Polynemus virgiriicus
(which also contained small fish), small Trichiurus Zepturzts (larger ones ate fish).
.-lchirus sp. (contained ~S'quilla),Serranids such as Diplectrum spp., Lutjanus synaqris
(some of \t hich also had fish). Among the sciaenids, penseid shrimps were found in
.Mrntirirrhits (togcther with crab remains), limbrina gracilicirrhus, Nebris m i c r o p
arid Lonch urus lanceolalus. Penacids were also f o l d in young Macrodon m q l o d o n .
. i d togrthc~with fish in larger ones ; in Cynoscion jarrinicensis, and in C . uirescms
they wcre found together with Squilla and fish.
Crab remains were found in MusteluR nov. q).(together with squid), Muraenesoi
.savanna, A'yacim papi1lotsu9ti and Menticirrhus spp. Other bottom invertebrates
t2att.n included hydroids, sponges and feather stars (Tropiometra carinata) crammed
into Chmetodipterus faber on diflerent occasions. Molluscs were found in h'phoeroiden
tmtudinew. Ogcorephalus truncntwx ttnd Conodon nobilis.
(3) ' I ' h c b mow pelagic. white shrimps (Srrgestidar) were found in . the Manta raj
XLIVI
THE 1~’lSIJES O W THE BRITISH GtrIAYA C’ONTINISNTAL SHELF
wi
Xobulu sp., sonie herrings. carangids such as Oligoplites mliens, Curunx hippos,
C. crysos and Heniirarans ambl~yrhynchun,the moonfish Vomer setapinnis (together
with small herrings and anchovies) and Selme vomer, the bumper Chloroscombrus
chrysurus (together with anchovy larvae). Among sciaenids small C. jumaicensis.
and Larimus breviceps with its upturned mouth, seemed t o feed on sergestids and
’
when very plentiful sergestids were sometimes found in Macrodon and C. 2wescens.
The herrings Harengula pensacolae and Opisthonema oglinum had been feeding on
zooplankton, and the greenish stomach contents of Peprilus paru suggested this had
lwen feeding on plankton too.
(4) Fish were found in the stomachs o f : sharks, which seem to eat whatever
species they can catch and have a varied fish diet ; guitarfish Rhinobatos percellens
(small fish eaten) ; the two species of Bugre, which both included some of the commonest small fish in their diet ; barracuda 8phyraena guachuncho which were taking
mainly pelagic fish (clupeids, anchovies and Ablennes) but, also eat Macrodon and
Trichiurus ; the threadfin Polyn ~ m u svirginicus, which contained mainly anchovies
and clupeids (one was found with a small flounder) ; the mackerel Swmberomorus
spp. which take pelagic species, clupeids and anchovies, (and other smaller Scomberomorus) ; the lizard fishes which all take small fish ; some of the carangids, such as
Vomer setupinnis which included small pelagic fish, clnpeids, anchovies, and odd
Trichiurus, in their diet. Some snapper Lutjunus spp. were found with small fish
among the invertebrates in their stomachs-all much chewed, particularly about the
head, and hard to identify (one L. qnoxjris contained a hatfish Ogcocephalus sp.
together with Squilla).
The common sciaenids Macrodon ancylodon and C. virescens are primarily shrimp
and fish eaters, the bigger Macrodon taking Stellifer spp., Trichiurus and small
Polynemus as well a8 shrimps. The larger predator C. virescellnp, one of the main
predators of the inshore waters, feeds mainly on other sciaenids, and stomachs
contained Macrodon ancylodon, Micropogon .furnieri, C. jamicensis, numerous
Stellifer spp., Umhrina sp., and non-sciaenids such as Trichiurus lepturus, clupeids.
odd catfish (B. bagre and Arius sp.), and odd lizard fish (Sauridu sp.) and brown
sandeels. C. vircscens of all sizes seem t o take shrimps, Squillu or fish, whichever
are most easily available, the largest virescens eating shrimp a t times ; Squilla were
particularly abundant in their stomachs at one time and place (e.g. in September
1958 in Box I11 in 14 fathoms), and Trichiurus lepturus a t another time and place
(e.g. in February 1959 in Box VI in 22 fathoms) ; these Trichiurus were surprisingly
long to be swallowed whole, for example a 49 cm. G’. virescens contained a 45 cm.
Trichiurus folded in its stomach, headfirst and one virescens contained two cutlass
fish 45 and 50 em. long. C. jumuicensis also eats small fish frequently, and Lonchurus
lanceolatus includes some small fish in its diet. (The only fish ever found in a croaker
Micropogon was a brown sandecl of a kind taken along with bottom invertebrates
by Narcine.)
I n Figure 4 an attempt in made to illustrate the main food relationships in the
main trawling zones (I1 and 111).
Further offshore over hard sand and coral bottoms : the snappers contained
invertebrates and a few small fish ; the only sciaenid in this area, Eques lanceolatus,
contained tiny, Y’riton-like univalve molluscs. Sponges were found in both black and
French angel fishes (Porncanthus arcwctus and paru). Chuetodon spp. contained
tiny shrimps, Scorpaena spp. had Squilla and small crabs. The few scarids taken
contained gritty brown mush-the coral in this area, all in water deeper than 20
fathoms, appears to be dead. The various monacanthids contained hydroids,
feather stars and pieces of coral. Few predatory species were caught in the trawl
in this zone (hook line catches show they are here, but above the bottom). Perhaps
the spininess of these bottom fish deters predators ; one gurnard had a small flatfish
in its stomach. The only shark caught over the hard bottom. Carcharhinus acronotus.
contained octopus legs and fish.
IJ .L.B.z.
tt O H EM AK Y H . I ,O\V K ( M N W K N ELI ,)
liHti
It is interesting that sharks, barracuda, Scorr~brromorus and Yomatomus, all fish
of relatively clear water, all chop their prcy before swallowing it, so stomachs sometimes only contain part of a fish (different parts of one garfish Ablennes hians were
found in two Sphyraena guachancho caught a t the samc time). The snapper, which
idso live in clear water, chew and fold tlieir prey. The sciaenids, on the other hand,
whose main habitat is very muddy watcr, where locating pieces of chopped prey
might not be so easy, all swallow their prey whole, generally headfirst. The Burprisingly large Trichiurus swallowed whole by G. virescena have already been mentioned.
T H E M A I N FOOD R E L A T I O N S H-~
I P S O F T H....E COMMONER T R A W L - C A U G H T
FISHES
~
~~
~~.~~
. ..~
~
__
~~~~~~
....
~
FIQ.4.-The main food relationships of the commonest species caught in the trawl.
V. RELATIONSHLW
OF
THE
BRITISH GUIANA MARINE FISHFAUNA
WITH
THOSE OF
NEIQHBOURINQ !hRRITORIES
The South American continental shelf is continuous from the Gulf of Mexico, off
Panama, south to Brazil. Large rivers enter the sea dong this whole coastline from
the Orinoco to the Amazon. These rivers bring down mud, and conditions would
appear to be rather similar throughout this whole area. There are, of course, some
local differences ; but there appear to be no major barriers to the dispersal of these
coastal shelf fish along the whole coast from the Gulf to the Amazon. The distributions of the British Guiana fishes found in neighbouring tcrritories are indicated in
Appendix I , from data taken from t h e following sources: Panama from Meek &
Hildebrand (1923-28), Venezuela from Schultz (1944, 104Y), Surinam from Boeseman
(1948 a and b, 1952,1953, 1956), French Guiana from Puyo (1949) and Durand (196l),
and Brazil from Fowlcr (1941), Tortonese (1948).
The West Indian island arc, with its clear blue water over hard coral bottoms,
offcrs very different conditions to the fish. Many of the fish species here are also
caught off Florida (Breder, 1948, Joseph & Yerger, 1966 ; Robins. 1958) and are also
recorded from Brazil (see Appendix T).
YLIV 1
T H E PISIIIW OF T H E BlWI’ISH UtJJANA (’ONTISJCNTAI~YHELI:
(is7
In British Gniana the . coastal shelf’ fauna is found in zones 1-11 (111),and t h e
island arc ’ fauna in zone IV (111); the latter approaching nearest to the coast in
the southeast of British Guiana where it comes into 20 fathoms, about 40 miles from
the coast, in Box I.
-4s so much exploration remains to be done in these areus it is perhaps premature
to compare species lists. However, summarising the data on the distribution of
bpecies caught in British Guiana in neighbouring territories, given in Appendix I.
it seems that :
(1) of British Guiana’s approximately 213 coastal and marine species considered
here, about 103 (48 per cent) are species known from Panama to Brazil.
(2) Of these 103 species recorded from Panama to Brazil, about 69 (67 per cent)
are also recorded from the FloridaIWest Indies region.
(3) Brazil has 22 species in common with British Guiana not found in Panama,
1 J I l t only 10 of these are not recorded north of British Gniana, (the other 12 being
recorded from the island arc or Venezuela), i.e. the ‘ southern element ’ of British
Giiiana’s fauna seems to be only about 5 per cent.
(4) Twentythrec (ten per cent) of British Guiana’s species recorded from the
West Indies and Florida are not recorded from Brazil, i.e. seem to represent a northern
and island arc element of the fauna. Of these 23 species 13 are only recorded from
Florida and the West Indies (and not from Panama), (three of these from the West
rndies only). This suggests that the ‘ island arc ’ element comprises between 5-10
per cent of British Guiana’s marine fish fauna.
(5) Of the 213 odd British Guiana species, about 85 have so far been recorded
from Venezuela to the immediate northwest (Schulz, 1944, 1949), about 32 from
Surinam to the immediate southeast (Boeseman, 1948a and b, 1952, 1953, 1956).
and about 82 from French Guiana (Puyo, 1949 ; Durand, 1961). It is understood
that further studie8 are in progress in these territories, so these numbers are likely to
be augmented in the near future.
(6) About eleven species which appear t o be undescribed or new, and on which
further systematic work is required, were collected during the Cape St Mary survey.
The more pelagic fish, thc carangids and Scomberomorus, appear to move about
a great deal, but little evidence was obtained from the trawl catches of regular
seasonal movements into and out of the area. Further north, off Florida and Maine,
the literature suggests that there is a southward movement of some species in the
winter months. Hookline catches off British Guiana from the snapper schooners
do suggest that species such as the dolphin Coryphaena hippurus visit the area a t
certain times of year (for example a run appeared in August 1958). Among the
inshore and bottom dwelling fish a number of British Guiana species show local variations, suggesting the popihAons do not range far, and future work may show that
some should be considered subspecies, or representatives of a cline along the coast.
The latitudinal distribution from the eauator northward of some of the srJecies
occurring off British Guiana is shown by ioseph and Yerger (1956) in their platcfi
I11 and IV.
‘
VI. SUMMARY
OF CONCLUSIOKS
This survey, mainly by trawl, of the coastal and marine fishes of the continental
shelf off British Guiana, an area hitherto unexplored, has shown that a t least 213
species, representing 72 families of fish, occur here. As systematic studies have yet
to be made of the offshore handline fishery and of the roastal pin seine, Chinese seine
and longline fisheries, the number of species is likely to be much higher. No check list
exists of the marine and coastal fishes off British Guiana and this present paper
attempts to help remedy this. Rather incomplete checklists are available for some
of the neighbouring territories. but very little is on record about the natural history
of any of the species.
AHH
R O S E M A I ~ Y 1 1 , i , o w ~(MVCV)NNWI,I,)
1J.L.LI.B
2. Thc fish faunas of the area considered off British Guiana were found to inhabit
four rather overlapping zones :
Zone 1, the vcry muddy coastal watcrs out to ten fathoms over very soft, shifting
rnud (rarely visited l ~ the
y trawler and as yet little studied) ; zone I I , the harder mud
itnd sandy-mud bottomed areas from 10-20 or 30 fathoms (20 in thc southeast,
90 in the northwest), which proved the best areas for trawling, producing an average
of over 150 kilo of fish per fishing hour ; ; m e 111,the clearer, greener, waters, generally
over rather barren sandy bottoms ; zone 18,the clear blue water over hard coral
and sand bottoms, (out from about the 20 fathom line in the extreme southeast and
from 3 0 4 0 fathoms further north). There were some seasonal changes in water type
and corresponding scasonal movements of fish, particularly between zones I and 11,
and zones I1 and 111 ; in zone 111 niorc pelagic fishes were caught in the trawl.
3. The species of fish caught off British Guiana are listed in Appendix I, which
indieat.esthe zone in which each specics was most commonly found, and its distribution in neiglibouring territories. Zone I, the ' brown fish ' zone yielded some 02
species, mainly soft-bodied nematognat,hs, sciaenids and rays, zone I1 the ' goldcn
fish ' zone, from which about 80 species came, was predominantly a sciaenid zone.
these making up over 75 percent of the wcight of the trawl catch. Tn zone I11 the ' silver
fish ' zone only about 43 species were found, the cstch was largely of pelagic fish,
carangids, Spanish mackerel, barracuda, hcrring and pomfret, which came and went
with the water types. In zone IV, t'he red fish zone, catches, though rich in
numbers of species (at least 80) were poor in numbers of individuals and weight of
fiRh caught (see table I) ; large red snapper were rarely caught in numbers in the
trawl and most of the catches were of small spiny red species.
4. Most of the species were present throixghout the year. There was however, a
gcneral inshore movement,during the rains, J u n e to September, the calm months when
much fresh water is pouring down thc rivers into the sea, and an offshore movement
in thc vrry windy months January-March. This offshorc movement brought some
of the zone I fish into zone 11, and catches were particularly good from the more
offshorc grounds in the northwest after prolonged windy periods. Whenever packs
of carcharhinid sharks appeared in the neighbourhood, the fish evidently scattered
;Lnd catches became very patchy ; the. profound influence of the presence or absence
of sharks on the distribution of the ot'lier fish demonstrates well the enhanced importance of biotic pressures in the tropics where seasonal changes are so slight.
(5) Gonad stat#esshowed that in some species ripe fish ca'n be caught a t any time
of year ; these included the croaker Micropogon furnieri, the bashaws Cynoscion
jamuicensis and Larirnus Dreviceps, t,he seapatwa Diapterus rhombeus, the grunt
'anafolk Cen,yatremu.s luteus and the nioonfish Vomer setapinnis. Some ripe fish were
also recorded from all tlic months that the following species were caught, Conodon
nobilis, Paralonchurus sp., Menticirrhws sp. and Bagre m.arina. Thorc was, however,
ovidencc of more restricted breeding seasons in the vcry common sciaenids Macrodon.
nncylodon and Cynoscion virescens, but ' place effects ', the fish with active gonads
moving away from the trawled zone, probably inshore, obscured scasonal changes in
the numbers breeding. Among Nebris mn,icrops and Isopisthus sp. caught in the trawl
none were ever found wit,h a'ctive gonads. I n French Guiana, Puyo (1949) maintained that many specics spawn during the rains, wlion they are caught abundantly
in Chinese seines in the estuaries, but the British Gniana data suggested that spawning
periods may be much more protracted. Young Micropogon furnieri were caught
throughout the year entering brackish swamps a t Onvcrwagt.
(6) Many species of viviparous and ovoviviparous sharks and rays were found
carrying young, particularly in February-April, when the young of several species
were about ready to be born.
(7) Stomach content analyses showed that the most abundant fish, the croaker
Micropogon furnieri were feeding mainly on polychaete worms from the bottom mud.
The next. most. abuutlant: specieR. Macrotlon nncylodon. and C'~~n.osc?:orr.
.tiirescem ate
T H E FISHES (JY THE I3KITISH ( 4 U t A K A ( N N T I S KKTAL HHELV
XLIV
W!)
penaeid shrimps and fish, Macrodon taking shrimps when small and including fish
in their diet when larger. C. vire.scens was one of the main predators of the sciaenid
zone but took penaeids and Squillu at intervals throughout life ; the fish eaten were
mainly other sciaenids and included Macrodon and Micropogon . Penaeid shrimps
were found in a t least 18 species. Other predators Mere the sharks which had a very
varied fish diet, and barracuda and mackerel, which ate more pelagic fish such as
clupeids and anchovies. Like Pomatomus these chopped up their prey before eating
them, in contrast with the predatory sciaenids living in muddy waters which swallowed
their prey whole.
(8) Comparison of the species recorded from British Guiana with tliose from
neighbouring territories shows tliat 48 per cent of British Guiana’s marine fishes considered here are also found from Panama to Brazil ; and of these 67 per cent are also
recorded from Florida and the West Indies. Only 10 species in common with Brazil
are not recorded north of British Cuiana (i.e. about five per cent of British Guiana’s
fauna). Of 23 (ten per cent) species found to the north of British Guiana but not
recorded from Brazil, 13 were not in Panama and appeared to be purely ‘ island arc ‘
species. Thus British Guiana’s marine fauna appears to be made up of about 50
per cent widely distributed coastal shelf species, and an island arc component (between
five-ten per cent of the fauna) of clearer-water loving species, coming inshore to
within about 40 miles of the coast in Box I in the southeast where there is a hard
coral and sand bottom in 20 fathoms, and about five per cent appears to be a ‘ southern
element ’ in common with Brazil. About 11 possibly new species on which further
study is required, were collected during the survey.
In the complexity of the fauna, the large numbers of species present, many
closely related and often appearing to eat much the same food, and the presence in
several species of ripe fish throughout thc year, British Guiana’s marine fish fauna
demonstrates characteristics of tropical as opposed to temperate communities. The
large numbers of fishes from these very turbid waters which produce loud noises,
particularly among the sciaenids, grunts and catfishes, is very striking and provides
an exciting subject for further study.
VII. ACKNOWLEDGMENTS
I am very grateful to the past and present Directors of Agriculture, British Guiana.
M i A. F. Mackenzie and Mr G. B. Kennard for the opportunity to make this study.
I would like to thank the Master of the Cape Bt Mary, Mr W. 13. Mitchell, and his
officers and crew, and the Fishery Officers Messrs W. H. L. Allsopp and E. Shepherd
and their staffs for help in many ways, particularly my assistant Mr E. Wong, who
made collections when I was not able to accompany the ship and carried out much of
the tedious work in the analyses of the catch records. Miss V. E. Graham very
kindly helped to pack the invertehrates for dispatch to the British Museum (Natural
History) for identification.
At the British Museum 1 would like to tharik the folloning members of staff for
naming the invertebrates: Dr I. Gordon and Mr R. W. Ingle (Crustacea), Miss
A . M. Clark (Echinoderms), Miss A. C. Edwards (Annelids), Mr 8 . Prudhoe (parasitic
worms), Dr W. J. Rees (coelenterates). I would also like to thank all the members of
the Fish Section who helped while 1 was working there on these collections, particularly
Dr. E. Trewavas for her encouragement and advice, and Mr N. H . Marshall and Mr
G . Palmer for invaluable assistance with the literature.
VIII. REFERENCES
~ L S O P P ,W.
11.L.
I: 1-52.
~ L S O P P ,W. H. 1,.
Gitiuiin i-’i.sli.
195% H.eview of the fisheries of British G U I ~ I I M
Byit.
. ffzliana&%ah.Dzv. Bull.,
1960. Oiiverwagt brackishwater fish culture station British Guisna. Brit.
I&*. Bull., 3 : 1 4 3 .
ti!#
KOYEM.4K.Y I I
)
LOW6 (McC’OUNlCLL)
1J .L.S .Z.
13. A. 1894. i3rtentdir rasults of explorations 1)y the U.S. Fish Coniinimon steamer
Albatross. No. XXXIII. Dosrriptions of two new flonndors, @astropsefta frontolis and
Cyclopsettu chittendeni. I’roc. U.S. 1Vl;ot. Mtia., 17 (No.1030) : 633-636.
I(ICEBIS, W. & TEE-VAN,
J. 1928. Tho finhey of PorL-au-Prince Bay, Haiti, with a suinniary of
tho known species of inai in(^ fish of the islaid of Haiti and Santo Doiningo. %07ogirrr.
N . Y . , 10 (1) : 1-279.
-.
1933. Pield book of the ahorc jislics oJ’ Ber/rrutfn. 337 pp. l’utnain’s, N.Y.
- 1937. A now specieb of ( ’ ( z i ~ I o l d I i / 8
from TIiriidad, British Wost Indier. Zoolopcri,
I:PAN,
149 pp. Anglo-American Caribb. Coinni. Washington, D.C.
1948 & 1953. b’ishes of the Western North At’lantic. P t . 1: Sharks No. 1, Pt. 1,
194X : 5B~540 ; Yt. I1 : Sawlishes, Guitarfishes, Skahes, Rays. No. I, pt. 2, 1953 : 1--514.
M e m . Seurs P o u t d Marine Hesearch.
.
-~
1953. Fishes of tho Gulf of Maine. U S . B i a h . & Wildlife Fish. Bull., 74 : 1 -577.
HoEsamfAx. bf. 1945 a. On new, littk known, or othorwise intoresting species of surinatii
fishes. Proc. Kori. Ned. Akatl. Il‘et. Ainsterdain, 51 : 3--11.
. - - - 1948 b.
Home preliminary notes on Huririani stirigrays, inoluding a cIewrit&1i of u u t ~ w
xpecics. Zool. Medad. Mu.9. Leiden, 30 ( 2 ) : 31-47.
ii*s
of
1952. A preliminary list of Surinam fishes not included in ~ ~ i g ~ n n i a nonuiiieratioit
1012. 2001.Merlerl. Alm. Leiden, 31 (17) : 179-200.
.-1953. Scientific results of thc Surinain Expcdition 1048-1949, Part. TI. Zoology, No. 2. Tliv
Fishes (I). Zool. Meded. Mua. Leideri, 32 (1) : 1--24.
1 W i On recent accessions of Surinarii fishes. ZooZ. Meded. Mus. Leiden, 34 (12) : 183-199.
BOHLKE,
J. 1956. A srnall collection of new ocls from Western Ptiorto Rico. Not. Nut. Acorl.
Sci. Philad., 289 : 1-13.
HKEDER, C . M. 1927. Scientific results of tho first oceanographic expedit.ioii of the ‘ P a w n ~ eI ,
1925. Fishes. BUZZ.Binyharri. Oceutiog. CoZZ., 1 (1) : 1-90.
h!!.
13it7,yham
-~ . . 1938. A contributjon t,o the life histories o f Atlantic ocean flying fishes.
Oceamog. Coll., 6 (5) : 1-126.
- 1948. Field book of itwrina jislies oJ’ / h e A t h i t i c Coast j r o m Labrarlor to T e r m . 352 pp.
Putriain’s Sons, N.Y.
Hxowr;, H. H. 1942. The fisheries of British Guiana. Dew. & Welfare in West Iiidies (AnyloAmericwi Caribtsan Comm.) Bull, No. 4 : 1-69.
ODRAND,J. 1961. Notes siir le platcau Guyaimis. Las elements priiieipaaux de la fauiic et
leurs rdations avec Ie fond. Cahiers qf.Rech. Sci. Tech. Outpe-Mer, I’uris, Nu. 3 : 1-33.
E r c ~ I ~ t a i A ~C.
N , H.
1912. The freshwater fishes of British Guiana, iucluding a study of tho
ecological grouping of species and tho relation of the fauna, of the plateau to that of the
lowlands. Mem. Canzegic J!!w.,5 (No. 67) : 1L.564. (Includes brackishwater species.)
I<:KDnMN, D. s. 1956. Recent $sh records f r o m Yuerto Rican waters. Cyclostylod report., Carib hean Commission, Part-of-Spain. Trinidad.
Food of pelagic fishes frort, P~uarLoRico nrid the Baliunia Ish,nds. Cyclostyled report,
.~1958.
Caribbean Coniin., Port-of-Spain, Trinidad. (Reprinted from 3rd Intcriiational Cam: Fish
Conforcnoe, Miami Beach, Florida.)
FOWLER,H. W. 1939. Notes on fishes from Jamaica with descriptions of thrcc nuw speoioli.
,Not. Nut. Acad. Sci. Phila., 35 : 1-16,
.- 1941. A list of the fishes known from tho coast of Brazil. ArpfLiTO8. Zool. Sao l’cccclo, 3 (6) :
115-184.
- -1946. Notes on B collecfioit of fidies h m Trinidad. Not. Nut. h a d . Sci. Phila,, 166 : 1-11,
1947. Notes on Bahama fishes obtained by Mr Charles 0. Chaplin in 1947, with descriptions
of two new species. Not. Nat. Acad. Sci. PhiZa., 199 : 1-14.
.
1950. Rcsults of the Catherwood-C~iaplinWest Indies Expodition. 1948, Part 111, Tliv
fishes. Proc. Acad. Nat. Sci. Philo., Ill: 69-93.
1952. Fishes front deep water off southern Florida. Not. Nut. A i d . Sci. Philu.,
246 : 1-16.
FKASER-BRDNNER,
A. 1940. Notex OIL t h c Plectognath fishes. 11.1. On dfotaacanthus sst,$er
Bennett and related species, with a key to tho genus Stephanolepis and descriptions of font.
new species. AWU.Mag. tmt. Hist. ser. 11, 5 : 518-535.
1941. Notes on the plectognath lishcn. 1‘1. A synopsis of the gencra of the family Aluteridao, and descriptions o f 6even new specks. Ann. Mag. m t . Hist., ver. 11, 8 : 176-199.
Ursss, R. €1. 1959. 4 synopsis of t h o pstlarvae of Western Atlant,ic lizard-fidws (8ynodontidae). Copeia, 3 : 232-236.
)
on tho coasts of t.he h n w r i c e ~
O r ~ s s u mI.
, 1937. Itoviow of the Seahoruotx ( H i p p o c w m p ~found
continents and Europc. Proc. U.LS’.Nut. M m . Washirqton, 83 : 497-594.
- 1951. The eels of tho northern Gulf Coast of the IJnited States and soinu relat,ed q w c i w .
Texas Jour. Sci., 3 : 431-485.
- 1982. Fishes of the family Carangidae ol’tlin riorthern Gulf of Mexico and three related qm:ioH.
J’rtb. Inetit. Mariwe S c i . tTn&r. 7’exas. 2 (2) : 43-117.
~
~
-
~
XLIV]
TIIE FISHES O F THE BRITISH G U I A N A C0P;TTNENTAL SHELI'
(59 1
GINSBURG,
I. 1954. Four new fishrs and one little h o w u species from the east coast of the U.S.
including the Gulf of Mexico. J . Wash. Acad. Sci., 44 : 256-264. (Cubiceps spp.)
GRAHAM,
V. $1. 1949. lieport o n the marine biological specimens collected during the simiey of
British Guiana food jishea in 1948 and 1949. Unpublished report, Georgetown.
HErwORTR, T. A. C. 1949. h'xperirnental fishery survey of the coastal uaters of British Guiana.
Unpublished report, Georgetown.
HIGMAN,J. B. 1959. Snrinani fishery explorations May 1 1 July 31, 1937. P o m w . Fish. Rev.,
21 (9) : 8-15.
HILDEBRAND,
S. F. & CABLE,L. $2. 1934. Reproduction and developincnt of Whitings, Kingfishos,
Drurns, Spot, Croaker, and Weakfishes or Seatrouts, family Sciaenidae, of the Atlantic
roast of the United States. Bureau Fish. Washington Bull., 48 (16) : 41-117.
JORDAN,D, 8 . & EIGENMANY,
C. H. 1889. A review of the Nciaenidrte of Anlei-ice and Europe.
Ann. Rep. Comm. Pkh & Fisheries Washinqtoia for 1886 : 343-451.
.JORDAN, D. 8. & EvnanlA", €3. W. 1896, 1898, 1900. The fishes of North and Middle America
Parts I-IV. Brill. U.S. A a t . MILS.washing to?^. 47 : 1-3313.
.JORDAN, D. S., EVKRMASN,
R . W.&L CLARK, H. W. 1930. Checklist of the fishes and fidilike
vcrtebrates of North and Middle America north of the northern boundary of Venezuela
and Columbia. Rept. U.S. Comrn. Fzshcries Washingtonfor 1928, part I1 : 1 670.
Josiwa. E. B. & YERGER,
It. W. 1936. The fishes of Alligator Harbour, Florida, with notea
on their natural history. Florida State Uniu. Studies 22, Papers Oceanogr. lnstit., 2 :
111-156.
KTJMA,E. F. 1959. Aspects of the biology and the fishery for Spanibh inackerel Sco~nbero~rcorus
rtraculatus (Mitchill), off Southern Florida. Florida Bd. C'oweru. Tech. Ser., 27 : 1-39.
KLIMA,E. F. & TABB,D. C. 1959. A contribution to the biology of the spotted weakfish,
Oynoscion webulouus (Cuvirr) from northwest Florida, with a description of the fishery.
Florida Board C'oiuerv. Tech. Ser., 30 : 1-26.
LONGLEY,
W. H. & RILDERRAND,
S. F. 1940. New genera and species of fish from Tortugas,
Florida. T'up. Tortugas Lab., G x n e g i e Inst. Washington, 32 (No. 517) : 22S-285.
_ _ __ 1941. Systomatic catalogue of the fishes of Tortugas, Florida, with observations on
color, habit and local distribution. Pap. Toitugns Lab., Carwegic Inst. Washington, 34
(535) : 1-331.
LOWE(MCCONNELL),It. H.(in press). Notes on the fiahes found in Georgetown fish inarkcts
and their seasonal fluctuations. Brit. Gzciana a'ish. Uiv. Bull. No. 4.
IIEEK,S. E. & HILUEBRAND,
8. F. 1923-1928. The marine fiHhex of Panama. Field iMus. Nut.
Hist. (Chicago) zool. Scries. 15 (Pts. 1-111): 1-1045.
MI:TZELAAR,J. 1919. Over tropisch AtZant.lachc Vkscheii. Part I : 1-179.
Kruyt-Uitgrwr
Amstcrdam.
MITCHELL,W. G. & MCCONNELL,It. H. 1959. The trawl survey carried out by tho Cape S t Mary
off British Guiana, 1957-1959. Brit. Guiana Fish. Div. Bull., No. 2 : 1-53.
NICHOLS,
J. T. 1929-1930. Thc fishes of Porto Rico and the Virgin Islands. N.1-. Acad. Sci.
Surv. Porto Rico and thc Virgin Is., 10 (2-3) : 161-399.
NORMAN,
J. R. 1934. A syRteinatic inonograph of the j7atjshes (Heterosoinatu) Vol. 1 . 459 pp.
British Museum (Nat. Hist.) London,
-1935. A revision of the lizard-fishes of the genera Synodus, .?'rachi)ioceplkulii~,
and SaiLrida.
Proc. ZooZ. Sac. Lond. 1935 (1) : 99-135.
NOTA,D. J. G. 1958. Sediments of the Western Guiana shelf. Rept. Orhnoco Shelf E'xpedition,
2 : 1-98, Vconman & Zonon Wageningen, Holland.
PALMER,
G. 1950. A new specics of electric ray of the genus Diplobatis from British Guiana.
Ann. Mag. nut. Hist., Ser. 12, 3 : 480-484.
Prryo, J . 1949. Poissorls dc la Guyane Francaise. 180 pp. Faiirie do I'Einpire E'rancais 12,
Ofice Recherche Sci. Outre-Mer, Paris.
REID,G. K., Jr. 1954. An ecological study of tha Gulf of Mexico fishes, in the vicinity of Cedar
f Caribbean, 4 ( 1 ) : 1-94.
Keg, Florida. Bull. Mar. Sci. G ~ l &
KIBEIRO,
M. 1915. F a u m Brasiliensis-Peixes.
Archivos Museu Kiacional Xi0 de Janerio, 17.
RIVAS,L. R. 1954. The origin, relationships and geographical distribution of the marinr
fishes of the Gulf of Mexico. [J.S. Fish. Wildlife Fish. B d l . , 89 : 503-505.
ROBINS,
C. R. 1958. Checklist of the Florida game and commercial marine fishes including
those of the Gulf of Mexico and the West Indies. Florida Board C'onreru. Idducat. Ser., 12 :
1-44.
NCHTJLTZ, L. P. 1944. The catfishes of Venezuela, with descriptions of thirtyeight new forms.
Proc. U.S. Nat. Mus., 94 (3172) : 173-338.
-~ 1949. A furthei contribution to the ichthj ology of Venezuela. i'ror.
U S . N a t . M u s . , 99
(3235) : 1-211.
SPRINGER,
8. 1939. Two new Atlantic species of Dog sharks, with a koy t o the species of
ihfustelus. Proc. U.S. N u t . Mus., 86 (3058) : 461-468.
-1960. A revision of North Amoricaii sharks allied to the genus Carcharhinus. Amer. M u x .
Nov&Am, NO. 1451 : 1-13.
SPRINGER,
S. & IAOWI,
R. H. (in press). A new smooth dogshark of the genus M ~ f s t e l ufrom
s
the
Northeastcwi .itlantic coaPt of South Amerwa.
[ J .L. s
Ci92
.z.
STARKY,
E . U. 1913. The j i s h e ~of the Stunford Expedition to Brazil. Stanford Univ. Pub. :
1L77. [Thalaasophrync branneri.]
STOREY,
M. 1938. West Indian Clnpeid fixlins o f t h o genus Harengulu. Atat?ford Ichthyol Uldll.,
1 (1) : 3-56.
.
rmt. H i s t . ,
TATE: REGAN. 1902. A rovirsion of the fishos of t h u faniily Stromatoidae. ~ I L ~Tfllaq.
set. 7, 10 : 115-131, 194-207.
TEAG~JE,
G. W. & MYERS,G. 8. 1945. A New Qurnard (Friomtua alipionis) froin tho coast, of
Brazil. Bol. Mus. Nac. llio de .Tnnw?iro. ~ L O J J acirw
I
Zool., 31 : 1-18.
'rEAUTW, G . w. 1951. Tho sea robinH of Arrierioa, a revision of tho 'rrigiid f i S h C S of t h e g e l l U S
P ~ ~ O T L OCOIIL.
~ U XZool.
.
MUY.Hist. Nut. Montcviclao, 3 (61) : 1-58.
TORTONESE,
E. 1948. Agggiuntu e rottifiche ctl catalog0 dei pesci mariui del Brasib. 61011.
Muse0 Zool. Urnin. I'orino, 1 (1943-48) (8): 3-7.
... - 1950.
A note on the Htrrnrnerhoad shark S p h y m a tudea Val., after a study of the typos.
A m . Mug. nat. El&., XBP. 12,3 : 1030-1033.
TRAVASSOS,
H. & PAIVA,M. P. 1957. I,ista do8 Scitronidae rnariiihiw Uraxiloiros oontctndo
chave de ident.ifacao H proposta da ' nomes vulgares oficittis '. Hol. In.&. Oceanoyr.
Univ. Sao Paulo, 8 (1 and 2) : 139--165.
WATTS,J. C. D. 1958. Tliermoclini: ofT thr, coast of ttoj)ical West. Africa. Nuture, 181 :
863-864.
WHITELEATHEH, xi. T. & I % U O W N , H'. H. 1$h?.
1 1 1 ~esperiitiu?nbl $ShWy S U r l J C y i l l [ I ' T i r b i d U d ,
Tobugo, and Rritiah (;uintm. 130 1 ~ 1 1 . Aiifilo-Aim~rioaiiCaribbean Uotmn., Washington.
\YOO-OS, L. P. 1958. Wetitern Atlantic specien of the genus Holoce.rLtricx. (!hfrrzgo Nat. His.
Mus. Fiieldiana. 25001. 37 : 9lLI 19.
APPENDIXI.-The naa&bejfiuhes caught qft' British G u i a m , mainly i i the
~ truwl, showirq
the zone in which each species I I most
~
frequenthy caught and its distribution
in neighbouring irwieru.
(Name of species in brai*kutadenotes that this was not caught in & / V C a p St !Wary'\
trawl : x donotcw pr(wont. ( x ) denotes occasional.)
British Guiana
Waters
Zones (see p. 675)
7-7
T
...
TI IT1 T V
*:
... x
...
x
......
(X)
(X)
...
...
......... x
...... x . . .
.< . . . . . . . . .
... x ( > < ) ...
............
... r: x ...
... j: :. Y ...
... x X ? ...
......
......
. . . ? ., . . . . . .
...
...
,L'
Y
.....
x
x
:<
.....
......
......
Species
TRIAKIDAE---8nlOotll dogfinherj
MzcRteZua nov. sp.
CARCITAREIINIDAE-~QClll bin sharks
Scoliodon terrae-novae Richardson
Scoliodon sp.
Carcharhinus acrotmtus (Poey)
Camharlciwua 1rmcu1ipi~nrc.i~
(Poey)
Carcharhinus oxyrhynchtcs Muller & Honk:
Carcharhima porosu8 Rttnzttni
(Corchnrhir~~t.s
leiicus (h.lullei~
8c Herile))
Eulaimia obscicrus (Lo Sucur)
Oarcharhinus limbatus (Muller & Heiilt,)
Aprionodon isodort. (Bluller & Hcdv)
SPIIYI~NIUAE-Hammor shark6
SphyrrLn lib uro (Liiin.)
S p h y m a tudes (Val.) (S.biyelowi Springer)
"5'ph.yrnalewini Griffith (8.diplcimz Springer)
PI4IYTTDAE-%W
. . . . . . . . . . . . . . . ...
x
......
x
>:
x
. . . . . . . . . . . . ...
. . . . . . . . . . . . . . . ...
i
:
.
............
.........
...
...
\'
. . . . . . . . . . . . ...
x . . . . . . . . . ...
x
.........
. . . . . . . . . . . . ...
. . . . . . . ...
fiSheS
Prist<apectinntus Latiiuin
Pristw perotteti Mullor & Honk
x
.......
............
...
ItIIINOBATIDAR~--QUitttrI'aJ'Y
..
Hhtnobatos percellem (Wttlbauiii)
TOIWEDINIDAEElectric: rays
Narcine brmilieJi8is (Olfern)
.........
. . . . . . . . . . Jjiplobotis pictics l'alnwr
DASYATIUAE-Sting rayY
Ilasyatis geijslcasi Bocnomon
... ,_. . . . . .
i.
;
i
. . . . . . Jlasyulis guttatus (Bloch & Schiicider)
GY.llr;uRID.~-Butturay rays
CJymnwa niicrura (Rloch & Schnoidor)
>
:x . . . . . .
~ J R O L O ~ H I ~ A E - ~ E O sting
U ~ ~ rays
Ilrotrygoia ,Ji,icrophfholt)iu t i t Delsmaii
.......
i
...
.........
Y
. . . . . ,' ...
...............
...
i
,<
. . . . ...
...
.i
.....
...
...............
...
Kritish Chianti
Waters
Zones (see p. 675)
--r
11 111 IV
Specioa
...
(X)
......
.........
...
......
.........
.........
i
...
rays
Aetobatzis narirmri (Euphrasen)
RHINOPTERIDAE-$OWOSt? rRV8
(Rhinoptera sp.)
MOBULIDAE--hf&nta TI%).&
Xobzcla sp.
ELOPIDAIC-TB~~OU
(Elops sauTza Linn.)
(Tarpon atlaiitici~s(Cuvier Sr Vdencirnne~)
fish
ALBUI~IDAE-BOI~~
Albula vulpes (Linn.)
CcbTEm AE-Herrings
Harengzila pensacolae Goode & Bean
flislia argentata Meek & Hildebrand
Opiathonet~zoglinum (Le Rueur)
Pellorn sp.
~lYLIOBATIIJAE--&h&?
( X )
... x
... :<
...
x
......
,<~
i
( x ) <;
...
...
...
.........
%:N~l~AULIIJAE-~llC~lOVit)S
x
x:
...
.........
.........
.........
h
......
...........
...
A
...
.........
...
h
...
x
,<
...
...
x
<,
x
X
x
':
......
......
......
......
......
.........
.........
/
.........
.........
>,
.........
+
.........
..........
......... /
......... x
............
.........
i'
/.
.........
...........
..........
. . . . . . ( X ) ...
......
(XY)
( X )
...
.........
......... x
.........
JOURN. Id".
x
ELou.--zooLoBY: VOL. XLIV.
.........
i'
................
...............
>'
x
i
,
:.:
i
I..
...
x
A
Y
:,
_, . . . . . . . . . . . .
................
... >.
...........
...
... :.\
.................
i
>r
i
:
. . . . . . . . . . ..'
..................
\
. . . . . . . ,.
/ /
...
+
>'
.........
..................
..............
1
...........
2
A mhouielfu (StoEephofwa)s p h j e r (Cuv. & Val.)
Nolephorus suriiramensis Bleeker
Pterengraulh atheTimides (Linn.) & ot'hwn
MUR~NESOCIDAE-SaV&nll& edn
Muraenesor savanna (Cuvier)
001s
ECHILIDAE-WOI~~~
Ahlia sp.
OPHICHTHYIDAIZ
- -Snake eel
Aplutopliis sh a uliodus Biihlkt
>rT'RAENIDAE --MOr&y 001s
Qymnoth.oru.cocellatws Agasnix
r~ltIIDAE---8etl CtttfiSileS
Bagre (Pelickthys) w m ~ i n a(Mitchill)
Ragre (Pelichthys) bagre (Linn.)
Brim stvicticassis Cuvier & Valencionneti
pariiwcassis Cuvicr & Valenciennes
Arius grattriicassis Cuvier & Valenciennccl
rlrius parkeri (Traill)
Arius fisstis Cuvier & Valenoicnnrs
(Arius tipixi (Agassiz))
(Seleriaspis hertbergi (Bloch) )
( ~ s e u c l a c i c l ~ e 7 l t etod
r ~do
s sits (Bloch))
(Hexan,cirntichthp-viryispiiiis (Cuviur 6
Valenciennes))
(Aspredo aspredo (Linn.))
SuNouoNTInAE--'Lizard fishes
7'rachinocephaluu myops (Forstor)
SyLodm foeteiw (Linn.)
SyiLodus poeyi .Jordan
Synodus iiitwmedim (Spix)
Sauridu sp. (nearest t%L7nbil and
udosquat)i.iY)
AN A R ~ , E I ~ I ~ , ~ E - ~ o u ~ - ~ ~ B U
(AiLablepsanablcps Linu.)
(Ana5lcp.q itiicroleppis Mullur K. Trt)sc+ic:ll
I ~ E L O N I I L + - -Needle fishes
(Stromgylum sp.)
Ablei~tteshiatla (Cuvier & Va1encieiuic:s)
Er;oc:oi~TrUAE-.Jl'lying fish
(Pure.cocoet.usbrachypterus littoralis Hreder
BOTHIDaE--FlOund0rS
(Paralichthys sp.)
Bothua near ocellatw Agassiz
Symiunt papillonurn (Linn.)
i
:
h
........
...
.....
......
.....
. . . . . . ... ... <,
. . . . . . ... ... x
. . . . . . ... ... x
. . . . . . ... . .
. . . . . . ... ...
......
......
......
......
,'\
.:\ "
...
...
...
. . ...
.....
...
.....
...
x
... . . . .
.........
...
...
...
..............
...
x
........
,.\
.
............
.........
?~
.................
............
. . . . . . > ...
...
...
..................
.............
...
( X )
.........
............
...
s
(2.)
( X ) (X)
............
41
ROSEMARY H. LOWE
( MCCONNELL)
[J.L.s.z.
APFENDIXI-cont.
British Cuiana
Waters
Zones (see p. 676)
&
T
I1 I11 Iv
Species
B o ~ ~ ~ ~ u ~ ~ + P l o u n dcont.)
ers-(
...... ( x ) x
Cyclopsetta jimbriata (Coode & Bean)
...... x ... Cyclopsetta chittcndewi B. A. Bean
AcHIaIDAE-soles
......... x
Gymmchirus near fasciatus Gunther
(P syn. zebrinus Ribeiro)
... x
Achirus near fusciatus Lacepede
x .........
Achirus lineatus (Linn.)
Achirus sp. (near ?nazatlanus)
CYNOGLoSSIDAE-TOngUO fishott
... x . . . . . . Symphurus plagusia (Sohneider)
HOLOCENTRIDAE-Soldier or Squirrel fishes
......... x
Myripristus jacobus Cuvier & Valenciennes
......... x
Hobcentrus ascensionis (Oshcck)
. . . . . . . . . x Holocentrus bullisi Woods
8YNGNATJiIDAE’-l-’ipe fish and Seahorties
......... x
Hippocampw, longiroatris Cuvier
FISTULARIIDAE-Cornet fishos
......... x
Fistularia petirnba Lacrpcde
MUGIILIDAE-MU~~~~S
x .........
(Mugil lizu Cuvier & Valenciennes
(basiliensis Agassiz))
x .........
(Mugil curema Cuvier & Valenciennox)
x .........
(Mugil incilis Hancock)
8P~YItAENIDAE-Barracudas
... x x ... Sphyraena guachaiicho Cuvier & Valeneietlnos
POLYNEauDaE-ThreadfinS
x x ......
Polynemus virginicus Linn.
CYBIIDAE-Spanish mackerel
. . . . . . x ... Scomberomorus cavalh (Cuvier)
... x x
Scomberomorus maculatus (Mitchill)
TRICHIURIDAE-CUtlass fishes
x x ......
Trichiums lepturus Linn.
CORYPIIAENXDAE-Dolphin
. . . . . . . . . x ? (Coryphaena hippurus Linn.)
8TROMATEIDAE-H&tr‘VeSt fishes
... x x ... Peprilus paru (Linn.)
......... x
Cubiceps sp. (nearest nigriargenteus Ginsburg)
......... x
1 Peenes sp. (nearest P. maculatus)
CARANOIDAE-Jacks, PoInpanos, Scads
......... x
Dccapterus punctatus (Agassiz)
......... x
Decapterus macarellus (Cuvier & Valonciennes)
......... x
T’rachurus lathami Nichols
x
Selar crumelzopiithalmus (Bloch)
... x x
Hemicaranx amblyrhynclws (Cuvier &
Valenciennes)
. . . . . . x ... Caranz hippos (Linn.)
. . . . . . x ... Carunx crysos (Mitehill)
. . . . . . x ... Alectis crinitus (Mitchill)
... x x ... Vorne~cretupinnis (Mitchill)
...... x ... Selene vorner (Linn.)
x A
Chloroscombrus chrysurus (Linn.)
. . . . . . A ... l’rachinotus carolinus (Linn.)
x .........
o.%goplitcs 8 a u w (Uloch)
... x x ... Oligoplites salicns (Bluah)
......
...
.........
...
...
...
x
,,
...
...
...
x x ...
.........
.........
x
x
..................
x . . . . . . . . . x ...
...
...
(x)
x
...
...
x
x
x
............
......
X
?
x
...
x
x
x
x
......
......
x
x
x
...............
............... x
... x . . . . . . . . . . . .
hNTROPomDAE-8nook
Centtopomus undecimalis (Bloch)
Centropomus ensiferus Poey
... x
x
x
x
x
x
x
x
x
x
x
x
... x
x
x
<;
.........
... x
x
x
x
x
x
x
x
x
x
x
...
...
x
x
x
x
x
x
x
... x
x
x
x
x
x
x
x
x
x
......
... x
...............
x
x
x
x
x
............ x
...............
x x ...... x
x ... x ... x
x
......... x
x
x
x
x
x
x
x
x
x
x
x
......
......
x
... x
x
x
x
x
x
x
x
x
x
x
...... x
............
x ...... x
x x x x
x x ... x x
x
...
x
x
x
x
x
x
x
......
x
x
x
x
......
x
X
x
x
x
x
x
X
x
x
x
POMATO&ZIDAE-BlUefiShC38
Pomatomus saltatrix (Linn.)
RACIrYCENTsIDAE-Cobis
Rachycentron canadus (Linn.)
x
x
x
Y
A
x
x
British Guiaiiu,
Waters
Zones (see p. 675)
------I
r
II
......
rrr
IV
...
......
... % . . . . . .
...........
......... x
. . . . . . .< ,:
. . . . . . x <;
...
j ,
I.
x
/
.........
,<
.x
.........
......... x
.........
Spocius
8EssANibAi~;--8ea basses, Groupers
Epinepld us ,tnystucinus (Poey )
(:arrupa riigritu (Holbrook)
Afyceberoperca sp. probably fulcata (Posy)
Hypticus sp. probably bwtripinus (Mitchill)
Iliplectruna rudiale (Quoy & Gaimard)
I)iplcctrum fortnosu:tti. (Linn.)
Uiplectrum sp. ('! nov. sp.)
Prionodes phoebe (Poey)
(Promicrops ituiara (Lichenstt:iu))
~'I~rAcaNTHIDAE-Bigeyes
Priacanthus arenutus Cuvier & Valencienntu
LOJJOl'IDAg-T~i~Jletails
(Lobotes s w i m i n e m i s (Bloch))
I~UTJANIDAE--fhapper
Lut.jwzus a p (Bloch)
......... x
... x :,' x
Lutjunus synagris (Linn.)
... ( X ) . . . . . . Lutjunus analis (Cuvier & Valenciennea)
Lzctjanus spp. (2 or 3 spp.)
......... x
Ocyurw near chrysurus (Bloch)
......... x
Rhoinboplites aurorubens (Cuvior
......... x
Valenciennes)
POMAI>ASYIDAE-G~tS
Haemdon sci'urus (Shaw)
. . . . . . . . . <,
Haemulon steindachneri (Jordan & Gilbert)
......... x
... ( X ? ) . . . . . . Haetnulon inclunuruni (Linn.)
......... x
Bath!/sto,rin h i u t o r (Jordan 8G Gilbert,)
Hnthystoniu aurolil~eutum(Cuvior B
......... x
Valencionncs)
Buthystotna striatura (Linn.)
. . . . . . . . . '*.
Anisotrem u s viryinicus (Linii.)
......... x
... ( x ) . . . . . . AnisotrenLus surinanimsis (Bloch)
( X ) x
. . . . . . Conodou nolriliu (Linn.)
... x . . . . . . I'oniudasys corvinaefonnis (Steind~clmer)
... x . . . . . . Pottiudasys sp. (near crocro (C. & V.))
... N . . . . . . Orthopristis sp.
Ue~~yutrcni~zrs
luteus (Bloch)
,< x . . . . . .
SPARIDAE--POPgit%
Calanius proridem Jordan & Gilbwt
. . . . . . . . . h'
GmrtIuAE-i'vfojarras
... ( X ) x >: Eucinostoinus gula (Cuvier & Valoncietiiies)
. . . . . . x . . . Eucimstonws argenteus Baird & Girard
( X ) x
. . . . . . Diupterus rhonibcus (Cuvier)
RfumruA~--Red mullets or Goatlislies
M u l l u s sp. (near surtnicletus Linn.)
............
. . . . . . . . . i: (Jpeneus tnuculatus (Bloch)
Uperieus yurc.us Poey
. . . . . . . . . b':
BcImNmm-Croalrers, Wealifislies
Larirnus
breviceps Cuvier & Va1enciernit.s
... s . . . . . .
... x . . . . . . Corvula sartctae-lzwkze Jordan
Stellifer rustrqer Jordan & Eigenmanii
x >< . . . . . .
Stellqer tnicrops (Steindnaehner)
X
x ......
-Wicropnyoii furnieri (Desmarest)
Y
x ......
... x . . . . . . Urnbrina yracilicirriius Metzelaar
Menticirrhus aniericuiius (Linu.)
... ( X ) x ...
1Men&irrhus martG&ei~sia (Cuvier B
... ( x ) x ...
Valenciennes)
Purulnnchzmrs sp. (nearest rathbrori ( 1 ordan
... Y . . . . . .
& Bollinan))
Lonchurus lanceolatus (Linn.)
i: . . . . . . . . .
x x . . . . . . Isopisthus paruipinnis (Cuvier & Valenoieiium)
Webris niicrnps Cuvinr & Va1rw:iennt.s
x >: . . . . . .
\
,
~
,':
<,
...
...
... ... ... X
. . . . . . . . . ... ...
. . . . . . . . . ... ,\. ...
i
.I.
x
X
Y
>'.
...
... ...
A.
. . . . . . . . ... ... ...
x
i
,
...
, ... ...
...
/:
x
x
.x
\I
*~
x
... X
... ...
i
.I'
... ...
x
Y.
x
x
x
:
x
x
,'.
...
x
.,
i
:
x
.....
\I
x
Y
...
...
...
,.. ...
... ...
... ... ...
,.
J
...
h
...
... ...
........
...
1..
x
... ...
... ...
... ...
?<
j
:
x
... ... ...
x :.: x
x ... X
...
x
... ...
.........
.
,
.,.,,
...
:i
... ... ...
... ... ?
... ... ...
... , .
P
/I
N
...
.*
... ...
...
...
x
i:
x
x
... ...
Y
X
Y
...
...
,'j
......
......
j j
... ,~
...
. . . . . . ... ...
><
x
x
... ... ...
... x
x
... ...
... ( x ) ... ...
.........
...
...
i(
x
...
...
,
\
...
...
X
696
ROHXMARY €1. LOWIG (M~CONNBLL)
[J.L.Y.z.
APPENDIX1-coat.
British Guiana
Waters
Zones (see p. 675)
0
E
Q
T
I1 I11 IV
Species
SCIAENIDAE-Croakers, Weakfishes (COnt.)
Macrodon ancylodon (Bloch & Schneider)
x
x ......
Cynoscion acoupa (Lacopede)
x .........
... x . . . . . . Cynoscion jamaicemis (Vaillant & Bocoiirt)
... x ? ... Cynoscion leiarchus (Cuvier C Valenciennes)
... x ......
Cynoscion vireacens (Cuvier & Valenciennes)
x . . . . . . . . . Cymscion steindachneri (Jordan & Eigenmann)
(Cymseion microkpidotus (Cuvier &.
x .........
Valenciennes))
x .........
(Plagioscion aquamosiusinavs Heckel)
Equea lanccolatus (Linn.)
......... x
BRANCHrosTEGIDAl?-~~anquil~os,Tilefishes
......... x
Caulolatitus guppyi Beebe & Tor. Van
ANTIaoNIDAE-Boar fishes
......... x
Antigonia capros Lowc
ErHIPPIDAE-spade fishes
... x . . . . . . Chaetodipterua faber (Broussonet)
CHAETODONTInAE--HUttOrflY
fishes, Angel fishes
Chaetodon biinnculatus Blvch
......... x
Chaetodon sp.
......... x
......... x
Pomacanthus arcuatus (Linn.)
......... x
Pornacanthus paru (Bloch)
......... x
Holacanthus tricolor (Blooh)
ACANTHURIDAE-SUrgeOn fishes
. . . . . . . . . x Acanthumu, h e p t u s (Linn.)
SCORPAENIDD-SCOrpiOIl fishes
... x . . . . . . Scorpa.e.tm pbumieri Rloch
Scorpaena braailiensis C & V
......... x
......... x
Scorpaena agasaizii Goode & Bean
. . . . . . . . . x Scorpaena sp. (near bergii)
Bcorpaena sp.
......... x
T ~ r a ~ ~ ~ ~ ~ - S e a r ogurnards
bins,
... x ...... Prionotus alipionis Teague & Myers
......... x
Prionotus sp. (near stcrnsi Jordan & Swain)
......... x
Prionotus sp. (nearest carolinus (Linn.))
......... x
Bellator militaris (Govde & Bean)
DACTYLOPTERIDAE-Flying Curnard
Dactylopterus volituns (Linn.)
...... x x
......... x
.........
.........
x
x
......... x
......... x
x
...
. . . . . . /':
...
...
...
x
x
......
...........
...... x . . .
x ? ......
.........
.........
x
x
"
!a?z
v w
7
-
... x
......
... x
... x
...
x
>.
X
X
x
X
x
x
x
x
x
x
x
x
... ...
X
x
x
>:
x
x
x
. . . . . . X ...
. . . . . . . . . ... x
......
:*
>:
... ...
x
... ... ... x
x
x
:
. . . . . . ... . . . . . .
. . . . . . . . . ... ...
...
x
x
x
. . . . . . .*. . . . . . .
Y,
x
x
x
x
x
x
x
,<
x
x
... ...
x
... ...
x
... ...
x
. . . . . . . . ... *.. x
,,
,,
. ' ~
x
><
><
x:
... ...
x
... ... x x
. . . . . . ... . . . . . .
x
. . . . . . . . . ... ...
x
. . . . . . ...
X
x
x
...
x
:..:,
i
>:
POMACeNTRID~-DeinOiRell~S
Abudefduf sp.
LABRIDAIE-Wr&RH05
Bodinnus sp. (near rujm (Linn.))
( X ) ( X ) ( X )
Halichoerea sp. (nearest rndiatzts Jordau)
(x) (x) (x)
kkARIDAE-ParrOt fiSheS
Cryptotomus sp. (nearest w t u s (C. C V . ) )
... ( x ) ...
S p a r i ~ o m aaqualid.um (Poey)or
... x ( X )
flavescens (Schneidor)
Ecamarum-Reruoras
Echeneia naucratee Linn.
x ... >:
Remora albescens (Temminck & Schlegel)
.........
OPHIDIIDAE-GUBk d H
Lepophidiuni brcvibarbe (Cuvior)
...
...
BATRACHOIDIDAE-Toad fiShe8
Batrachoides sp. probably s ~ ~ r i ~ n u ~ ~ i e t i s i s ... x
<
(Schneider)
Porichthys poroaksi?nus (Valenoienneb)
x x ...
Thalassophryne 6rameri Sttlrks
.........
BALISTIDAE-Trigger fidleS
Balistes carolinemis Gmelin
Ba.listes oetula Linn.
... ... ( X )
... ... ( x )
... ...
... ...
(X)
(X)
X
x
<;
...
<;
h
... ... x
x.
...
i
:
x
x
... ...
x
... ...
A
... ...
;
I
_I
XLIV
1
(197
THE FISHES OF THE: BRITISH OUIANA CONTINENTAL SHELF
APPENDIXI - c o n t .
British Guiana
Waters
Zones (see p. 676)
*
T TI 111
1x7
......... x
......... x
......... x
......... x
......... x
......... x
......... x
......... x
... x . . . . . .
x
x
......
...
...
x
x
......
......
......... x
......... x
... x . . . . . .
... x ......
.........
x
Species
MONACAiVTHIDAE-Fil8 fishes
Monacanthus ciliatus (Mitchill)
Monacnnthus hi8pidu.s (Linn.)
Alutem schoep$ (Walhaum)
Blutem ventralis Longloy
Alutera morwceros (Linn.)
OSTACIIDAE-Trunk fish08
Lactqphrys tricornis (Linn.)
TETRAODONTIDAE-Puffer fishes
Sphoeroides dorsalis Longley
Sphoeroides marmratus (Ranzani)
Sphoeroides testudinew (Linn.)
Colomesus psittacwr (Bloch t Schneider)
DIODONTIDAE-PO~CUpiIle fishes
Chilomycterus apinosue (Linn.)
Chilomycterus sp. ( P atinga var. (Linn.)
AmENNARIIDAE-Fishing frogs
Antennarius scaber (&vier)
OaCOCEPHALIDAE-Batfish
Ogcocephalus vwpertilio (Linn.)
Ogcocephalus truncatus (Cuvier k
Valenciennes)
Ogcocephalua sp. ( ? nasutus (c.& v.))
Halieutichlhys sp. (acukatus (Mitchill)
or caribbaeirs Garmen)
x
......
x
x
x
x
x
x
x
x
...............
......... x
......... x
x ...............
............... x
x x x ... x x
...
x
x
x
......
...... x . . .
x ...... x
x . . . x x
x ......... x
............... (x)
... x ...... x x
x x x ... x ...
............... x
...
(X)(X)
......
(X)
...
dddendum :
Since the preparation of Appendix I the following additional species of nematognaths have becn taken from the estuaries and from the coastal waters in the wet
seasons (see Lowe (McConnell)(in press) ; Xciadeichthys jlavescens (C. & V.), S. emphysetus (Muller & Trotschel), S. poops (C. & V.), S. (Selenaspis) passany (C. & V.), S .
(Selenaspis) sp. Brachyplatqdoma vnillanti (C. & V.), and Brachyplaaty,?tom,a spp.
(2 species).
~
APPENDIX11.-Additional
notes on certain little known ,fish species found off
British Guiana
It is hoped to publish elsewhere a t a later date an annotated list of fish species
caught off British Guiana. Meanwhile some comment should be made on the
occurrence of the rarer species listed in Appendix I and certain other species.
Before this survey was made, the following species appeared to be known only
from type specimens (or from types and very few others) : Diplobatis pictus, Dasyatis
geijskesi, Urotrygon microphthulmum, Aplatophis chauliodus, Cyclopsetta chittendeni,
Holocentrus bullisi, and Caulolatilm guppyi.
Other species recorded about which special comment should be made include :
Remora albescens, Nobula sp., Ogcocephulus sp., Halieutichthys sp., Eucimstomus gula
and E . argentem, Isopisthus sp., and Arius of the stricticassis, parmocassis, grandimsis
group.
A. Little Known Species
Diplobatis pictus Palmer, 1950.
Holotype collected during Arthur Rogers trawl survey off British Guiana and known
only from this and three specimens taken in Rosaura trawl from 10 and 43 fathoms off
698
~ ~ O S E M A I ~ii.
Y LOWE (MOCONBELL)
[J.L.Y.z.
Orinoco mouth (Tucker, 1954)". .During the Cape 8t Mary survcy it was caught
in ones or twos on numerous occasions, in February, March, April, August and
Septcmber, from Boxes I-VI and from 11-32 fat,homs over soft mud and sandy mud
with shells. All specimens were small, ranging from 9-15 cm. long. Both sexes
were caught but no females with embryos were noted ; as all spccimens were preserved for later study, no stomach contents or gonad stateR were examined in the
field.
Dmyatis geijslcefii Roeseman, 1948.
This stingray with its characteristic elongated snout and distinctive ' butterfly
shaped ' pelvic fins, previously known only from the type (a juvenile male, disc36 cm. long) taken off Surinam (Bocseman, 1948), was noticed in the R/V Cape St
Mary catch on one occasion when three large specimens were caught. This was in
Box I in February 1959, from 11-12 fathoms over mud sand and shell, where i t
occurrcd together with the relatively common D . guttatus. The colour in life in
black-grey (contrasting with the brown-grey of D.guttatus) and the sting spine is
noticeably shorter than in D. guttatus. The stomach of a female, (disc 44 cm.,
total length 100 cm.), was full of small shrimps.
iJrotrygon microphthalnzuin Delsman, 1941
Previously known from two specimens taken in 20-30 fathoms from the Amazon
mouth, this small ray was taken in numbers (together with Narcine brasiliensis and
Crymnura micrura) in February 1959, when strong onshore winds were blowing and the
fish had moved out, in Boxes 1-111 in 12-14 fathoms over ' mud, sand and shell'.
Both sexes were taken together. The sizes ranged from eight em. (disc length)
/17 cm. (total length) to 15/25 cm. ; the dark brown males were smaller than the
blacker females (males 8117-11/21 cm. and females 13/24-16/25 cm.). Many of
these females contained young nearly ready for birth. Two females each contained
three young, which were born, tailfirst with pectoral fins nearly rolled around the
body, when 4-5/74 cm. long. Unpigmented embryos had a high spiracular structure
over the eye ; the yolk ribbon was occluded while the embryo was still unpigmented.
The stomachs of two adult males and three females were full of small shrimps.
Aplatophis chauliodus Rohlke, 1956
A splendid specimen of Aplatophis chnuliodus, a name signifying with good reason
the unapproachable tuaky one, 87 cm. long with hugh teeth, was taken in Box
VI from 18 fathoms over soft mud in January 1959. Two other specimens between
60 and 70 cm. long were seen from shrimp trawl catches off Surinam ; one was a 67 cm.
female full of ripening eggs. This species, previously known only from the types,
was described from three much smaller specimens (the largest 466 mm. long) taken
near Puerto Rico. The extraordinary dentition of these larger specimens suggests
that this species, for which a new gemis u'as created by Bohlke, may one day bt>
comidered to deserve higher status.
Cyclopsetta chittedeni B. A. Bean, 1894
This species appeared to be known only from the type collected from Trinidad,
but has recently been recorded from French Guiantt (nurand, 1961). During the
R/V Cape St Mary survey it was taken in Box V in 24 fathoms over mud in January
1959. It is also occasionally to be seen in Georgetown fish market, together with its
near relative C. jimbriata (Goode and Bean).
"
.
/ l ~ r / /Brit.
,
iM!~.v. (.Vtlt. Hixt ). ZOO/ 2 ( 0 ) : 163- 314,
XLIV)
THE PIYHES OP THE BRITISH GUIANA CONTINICNTAL SHELF
699
Holocentrus bullisi Woods, 1955
Described from the Campeche Banks Mexico, Bermuda and Cape San Blas,
Florida, this species was occasionally taken, a few a t a time, from the offshore deeper
waters over hard bottoms, for example from 45 fathoms in Box 111 in April 1958.
Caulolatilus guppyi Beebe & Tee-Van, 1937
This species, described from Trinidad, was taken on two occasions, both in deeper
water with the ' island arc ' fauna, from 50 fathoms in Box I over coral and from 35
fathoms in Box V over sand, shells and mud. Several specimens all about the same
size (about 20123 cm.) were taken each time.
B. Other Comments on the RIV Cape Xt Mary Material
Remora albescens (Temminck & Schlegel)
Two wrinkled albino remora appear to be referable to Remora albescens (an
' Indo-Pacific ' species rarely recorded from the Atlantic Ocean), although they both
had only twelve doral laminae. These two fish, both 18 cm. long, were found on
deck with the large Mobula sp. (see below) caught in Box VI from 22 fathoms in
February 1959. They were completely pigmentless except for the pale yellow iris
and black pupil, and are said to live in the gill cavities of manta rays. When caught
they produced faecel pellets of shrimp remains (the Mobula was packed with shrimps).
These two remora were kept alive until the end of the cruise (four days) feeding on
bits of shrimp. They lived, and fed, attached to bottom or sides of a n enamel basin,
upside down or sideways apparently indifferently, staying in the same position for
days on end. When the water was changed they would make a flying leap a t the
side of the bowl and hang on with th9 dorsal sucker immediately on contact, and
often remain above the water for considerable periods, gulping air. The short
ventral pelvic fins with their stout but well padded, spiny skeleton were shot out
ventrally, suddenly and with great force, if the ventral surface of the fish was touched,
presumably a n adaptation t o ensure that the fish is not crushed by the host.
Mobula sp.
Manta rays were seen on several occasions swimming at the surface near the ship.
One caught in the trawl over 22 fathoms in Box VI in February 1959 was a male
Mobula sp. three metres (ten ft) wide. This seems very large for the western Atlantic
species M . hypostoma (Bancroft), which according to Bigelow & Schroeder (1953)
rarely exceeds four feet in width. The possibility was considered that this might be
a specimen of the larger-growing Mobuh mobula (Bonnaterre), the eastern Atlantic
form which sometimes strays west and grows to 3 4 metres wide, but this was very
unlikely as photographs show no tail spines or prickles. The stomach contained
half a bucketful of small white sergestid shrimps, two small jelly fish and one small
Trichiurus lepturus. The two Remora albescens came from this fish.
Ogcocephulus sp.
The systematics of the batfishes are known to need revision. In British Guiana
the common batfish over mud in less than 20 fathoms appears to be that figured by
Ribeiro (1915) from Brazil under the name 0. truncatus (Cuvier & Valenciennes), a
species now considered synonymous with 0. radiatus (Mitchill) Jordan, Evermann &
Clark (1930), Fowler (1941)). The Cape St Mar9 material when compared with
specimens of 0. radiatus in the British Museum (sent from Texas by Baughman,
1948) have shorter rostra1 processes and are not spotted, and suggest that the species
0, truncatus should be resurrected. It is noted that Durand (1961) now lists the
inshore species of batfish from French Guiana as Ogcocephalus sp., and not radiatus
as in Puyo (1949).
700
ttosmimT 1 1 .
mwb;
(M(CWVNNI,L)
1.1 . t . S . %
Hdieutichthys sli
The round hatfish caught offshore in 25-35 fathoms on several occasions seems
to be closer to a form of H . aculeatus (Mitchill), recorded from Florida, than to H .
cnribbaeus Garman, though the latter is the species found in the Caribbean and French
( h i a m (Jordan A Evcrmann. 1900 : h r n n d . 1961). This needs further investigation.
Eucinostowiuu g z h (Cuviw & Valenciennes) and E . argenteus Baird & Girard.
These two species are given as synonymous in Jordan Evermann & Clark (1930).
hut from Cape St Mory material, available for furt,her study, i t seems t8hatthey nit'
distinct species.
Isopisthu,~sp.
Some of the fsopisthux cwmrnon in catchrs in thc iuore inshore waters and estuaries,
from Chinese seines as well as trawl, and included (with Cynoscion jamaicensis)
tinder the name ' table bashaw ', differed from I . paroipinnis (Cuvier & Valenciennes)
previously recorded from Panama to Brazil, including Guiana. mainly by their
xhorter anal fin and the longer gap between the two doma1 fins.
.4P ~ I L Sstricticassis (Cuvier & Valenciennes), A . parmocmsis (Cuvier & Valenciennes).
A . grandicussis (Cuvier & Valenciennes)
Data collected during the R/V Cape St. Mary survey suggested that these three
Arius may repreeent three forms of a polymorphic species and not three distinct'
species. Field evidence for this was :
(n) these three ' species ' form a series from stricticassis with narrowest occipital
process and least tooth development (none on vomer and only two palatine patches),
via parmocmsis (wider occipital process and vomerine teeth present in adult but only
t,wo palatine patches), to grandicmsis, with widest occipital process and greatest,
tooth development (vomerine teeth in adult and four palatine patches). On occaions when any appeared in the catches, all three were generally taken togcther and intermediate stages were also caught a t the same time, between both stricticassis and
parmocassis and between parmocassis and grandicassis. Moreover there was considerable variability, tooth development not always agreeing with occipital process
form (for example a ' strieticussis ' occipital process form fish having four patches
of palatine teeth) ; sometimes tooth patches were better developed on one side of
the head than the other. It seemed in fact, that there was a more or less graded series
of increasing ' boniness ', and i t was interesting that the weight for length increasc.d with the boniness, being greatest in grandicassis.
(b) Ripening fish of d l three forms were caught a t the same timt:. Zn all forms
ripening females were ahout 50 cm. long.
(c) Stomach conten& showed that small rrabs (leucosids) and small shrimps are
(.aten by all forms.
Ma,terialis available for further study of this problem.