1
Cephalopods caught on the outer Patagonian shelf and its upper and medium slope in relation to the main oceanographic features
2
3
Ángel Guerraa, *, Julio M. Portelab, José Luis del Ríob
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
b
a
Instituto de Investigaciones Marinas (CSIC), Eduardo Cabello 6, 36208 Vigo, Spain
Instituto Español de Oceanografía (IEO) , C. O. de Vigo. P. O. Box 1552, 36200 Vigo, Spain.
Abstract
Ninety cephalopod specimens were collected in 71 of 132 hauls (54%) during the bottom trawl survey ATLANTIS 2009 undertaken between 24 February and 1 April
2009. The surveyed area was the zone between parallels 44° and 48° South, east of the Argentinean Exclusive Economic Zone down to the 1500 m depth contour on the high
seas of the Southwest Atlantic. The collection was composed of 16 species of squids and 5 of octopods. The best represented groups were Histioteuthidae (5 species) and
Octopodidae (5). The most abundant species were Gonatus antarcticus (25.5%), Histioteuthis atlantica (11.1%), and Muusoctopus eureka (8.9%), which were also the most
widely encountered. The geographic and/or bathymetric distribution ranges of 9 species are extended, and this is the first record of Galiteuthis glacialis outside circumpolar
Antarctic waters. Our data show that several species, mainly of octopuses, penetrate the area studied as the plume of cold sub-Antarctic waters is pushed far into the South
Atlantic by the Falkland (Malvinas) Current.
Keywords: Cephalopods, Biogeography, Patagonian slope, Southwest Atlantic.
Running title: Cephalopods from the Patagonian slope
25
26
27
28
29
* Corresponding author at: Instituto de Investigaciones Marinas (CSIC), Eduardo Cabello 6, 36208 Vigo, Spain. Tel. +34 986231939 ext. 180; Fax: +34 986 292762
E-mail address: angelguerra@iim.csic.es (A. Guerra).
1
30
1. Introduction
31
32
The Southwest Atlantic Ocean (SWA) supports two major cephalopod fisheries. One is focused on the Patagonian long-finned squid Loligo gahi and the other on the
33
Argentine short-finned squid Illex argentinus. Although there is considerable information on these 2 species along the continental shelf within the Argentinean and Uruguayan
34
Exclusive Economic Zones (EEZ) (Nigmatullin, 1989; Basson et al., 1996; Agnew et al., 1998; Haimovici et al.,1998; Bunetti et al., 1999; Agnew et al., 2005; Portela et al.,
35
2010), mainly around the Falkland (Malvinas) Islands (e.g. Arkhipkin et al., 2004; Bazino et al., 2005; Sacau et al., 2005), very little research has been carried out on the High
36
Seas (HS) region. The recently developed Falkland (Malvinas) fishery regularly catches octopuses during survey trawls as does the Falkland Islands Government Fisheries
37
Department (FIGFD) through observers attending commercial trawling operations on the South American continental shelf in the SWA, and research received a further boost
38
when the FIGFD funded a study to identify the octopuses in their fisheries collection (Gleadall et al., 2010). Recently, L. gahi, I. argentinus, Onykia ingens, and Gonatus
39
antarcticus have also been shown to occupy the ecological niche of epipelagic fish on the Falkland (Malvinas) shelf and slope (Laptikhovsky & Arkhipkin, 2010).
40
Many papers have been devoted to discovering knowledge of the teuthofauna from the Antarctic waters (see, for example, Rodhouse, 1989, and Gleadall et al., 2010, and
41
the literature cited therein). However, little attention has been paid to the cephalopod species from the Argentinean shelf and even less to the biodiversity present in the
42
Patagonian upper and medium slope. Practically the only data available came from the cruises carried out by Rodhouse et al. (1992) devoted to the early life cycle of
43
cephalopods, which surveyed a wide area using RMT8 and Bongo nets, and associated the captures with the major oceanographic features of the SWA.
44
This paper examines the collection of cephalopod gathered in the cruise ATLANTIS 2009 carried out in the HS of the SWA, including the outer Patagonian shelf and the
45
upper and medium slope, in relation to the main oceanographic features of the SWA. It also aims to contribute to the knowledge of the teuthofauna in the region, the
46
taxonomy of the material collected, as well as its geographical and bathymetric distribution.
47
48
2. Study area
2
49
The area surveyed was the zone between parallels 44° and 48° South, east of the Argentinean EEZ down to the 1500 m depth contour (Fig. 1), and the depth of the trawls
50
ranged from 107 to 1527 m. The physical oceanography of this area—virtually opposite the Gulf of San Jorge on the Argentinean coast—is complex. We therefore present a
51
schematic scenario, which will allow us to understand the main features that occur in the study area. The major oceanographic features of the SWA are the Brazil Current
52
(BC), the Falkland (Malvinas) Current (FMC), and their confluence region. The BC originates as a branch of the South Equatorial Current (around 08° South) and runs
53
poleward almost parallel to the shelf break. It is tropical in origin and is therefore warm, saline, and relatively oligotrophic. The southern limit of the warm water associated
54
with the BC fluctuates between 36°-38° and 46° South, and is accompanied by the intermittent formation of warm-core, anti-cyclonic eddies (Campos et al., 1995). The
55
encounter between the warm southward flowing BC and the cold northward flow of the FMC, at approximately 38° South, generates a strong thermohaline front: the
56
Subtropical Convergence or Subtropical Zone (STZ). The cool, nutrient-rich FMC branches off the Antarctic Circumpolar Current and flows northward along the Patagonian
57
continental slope. This current flows round both sides of the Falkland Islands before turning northward once again as a single flow, approximately 100 km wide. The FMC is
58
the dominant circulation feature along the Falkland Shelf Break. The sub-Antarctic waters from the FMC flow along the shelf as far north as 32° South before returning south
59
as the Falkland (Malvinas) Return Current and flowing offshore at the confluence zone. The surface water temperatures in the northerly flowing coastal current, between 24°
60
and 33° South, are several degrees lower than the Brazil Current and consist of a mixture of cool water from the FMC and warm water being recirculated from the BC. Brazil
61
Current waters have a temperature of >20.0 °C and salinity of >36.0, while FMC waters have a large temperature range between 4.0 and 15.0 °C, but a narrow salinity signal
62
between 33.7 and 34.1. Because of the large differential between these salinity ranges specified for the Brazil and FM Currents, a large proportion of the water sampled can be
63
expected to have mixed characteristics (Wilson and Rees, 2000 and references therein).
64
65
Using the nomenclature for the fronts and zones adopted by Peterson and Whitworth (1989), the three zones south of the STZ associated with the Antarctic Circumpolar
Current are, from north to south, the Sub-Antarctic Zone (SAZ), the Polar Frontal Zone (PFZ), and the Antarctic Zone (AZ). These zones are associated with fronts (Fig 2).
66
3
67
3. Material and methods
68
A multidisciplinary research cruise was conducted by the Spanish Institute of Oceanography (IEO) to assess the biomass of the main commercial fish stocks on the HS of
69
the SWA and to identify Vulnerable Marine Ecosystems (VME). The multidisciplinary bottom trawl survey (ATLANTIS 2009), undertaken between 24 February and 1 April
70
2009 on board the RV Miguel Oliver, included 132 trawl stations (Fig. 3 Table 1). The survey used a stratified random design with strata boundaries defined by latitude and
71
depth ranges. Scheduled fishing stations (hauls of 30 min) were performed using a LOFOTEN type net fitted with a “Rockhopper” mix train with bobbins and rubber
72
separators, suitable for deepwater fishing over irregular bottoms. The headline height averaged 6.1 m, with a wing spread of 20 m. Mesh size ranged from 160 mm in the
73
upper wings to 40 mm in the cod end liner. Trawling was restricted to daylight hours. The positions of the hauls were randomly chosen prior to starting each stratum survey.
74
75
Thirteen depth strata were defined in the study area (Table 1) and were further subdivided into 2571 grids of around 5 square nautical miles (nm2). Finally, 147 hauls were
randomly allocated according to the following criteria:
76
i) The number of hauls in each stratum was proportional to its surface, with a minimum of 2 hauls per stratum; ii) For each stratum, hauls were randomly allocated among
77
all possible grids, excluding those in adjacent squares; iii) When fishing was not possible in a selected grid due to bottom characteristics, the haul was put in the nearest square
78
allowing for bottom trawling.
79
Only 132 hauls were performed out of the 149 planned, as characteristics of the seafloor hindered bottom trawling, mainly in stratum 7 (1001-1500 m depth). Moreover, 5
80
of the hauls were considered null for different reasons, such as net breakage or net entanglement. Table 1 shows the scheme of fishing stations by depth stratum and
81
summarizes their main characteristics: depth range (m), surface (nm2), and number of grids per stratum. Hydrographical parameters (sea surface temperature [SST], sea
82
bottom temperature [SBT], and salinity) were collected during the cruise using a CTD-probe.
83
84
Catches were sorted immediately after capture. Cephalopods were stored in labelled plastic boxes and frozen at -20 ºC on board. Specimens were taken by the vessel to
Vigo (Spain) where the samples were transported to the Instituto de Investigaciones Marinas (IIM, CSIC) marine research laboratory for detailed examination.
4
85
After defrosting at room temperature, cephalopods were identified from published guides, and old (e.g. Nesis, 1987; G. Voss, 1978; N. Voss et al., 1998) and recent papers
86
(e.g. Guerra et al., 2000; Lipinski, 2001; Allcock et al., 2004; Vecchione et al., 2005; Gleadall et al., 2010). Total length (TL), dorsal mantle length (ML), and wet weight
87
were measured immediately, and the sex was also determined.
88
89
All the material caught on this cruise, except Batoteuthis skolops, which is in the National Museum of Natural History (Washington D.C., USA), has been deposited at the
Museo do Mar de Galicia (Vigo, Spain-www.museodomar.com).
90
91
92
93
4. Results
Locations of trawls conducted during the ATLANTIS 2009 survey are shown in Fig. 3. The typical range of SST, SBT, and salinity at the different zones, or the
oceanographic features cited in the text and Fig. 2 are shown in Table 2.
94
During the cruise, 90 cephalopod specimens, comprising 16 squids and 5 octopods, were collected. Depth ranges of capture and numbers of hauls with cephalopods
95
catches are shown in Table 3. Cephalopods were present in 71 of the 127 valid hauls (56%) carried out during the cruise. The family of squids best represented was
96
Histioteuthidae with 5 species, while the Octopodidae was represented with 5 species (Table 3). The most abundant squids were of the Gonatus antarcticus Lönngerg, 1898,
97
species (25.5%), followed by Histioteuthis atlantica (Hoyle, 1885) (11.1%). The most abundant octopus species (8.9%) was Muusoctopus eureka (Robson, 1929). G.
98
antarcticus also appears to be the most diverse spatially and was captured at 14 stations (Table 3). However, the distribution of H. atlantica and M. eureka, which appear in 8
99
stations, respectively, also have a wide geographical area of distribution within the limits of the total area explored.
100
Size and some other biological data of captured specimens and families (10) of each species are detailed in Table 4.
101
Table 5 shows the cephalopod distribution caught during this cruise in relation to the major oceanographic features of the study area.
102
5
103
5. Discussion
104
105
106
Six specimens of Brachioteuthidae share the diagnostic characters of Slosarczykovia circumantarctica Lipinski, 2001. This is one of the most common squids in Antarctic
waters (Lipinski and Young, http://tolweb.org/Slosarczykovia/24102).
107
Until now, very few specimens of Batoteuthis skolops Young and Roper, 1968 have been collected (Young & Roper, 1968; Rodhouse et al., 1992; Young and Roper
108
http://tolweb.org/Batoteuthidae/19452). The species is found only in Antarctic and sub-Antarctic waters and little is known of its biology. Due to the interest in our specimen,
109
we plan to present a detailed description and discussion elsewhere.
110
The taxonomy of the octopuses is still undergoing major revision (see, for example, Norman & Hochberg, 2005) and to date incirrate octopuses have yet to be the subject
111
of a comprehensive study using nucleotide sequence data. Such data are now being collected and research is also under way to provide clear species re-descriptions based
112
upon morphometric data, leading to a much better understanding of the major groups and biodiversity of shallow water octopuses (Gleadall et al., 2010). These authors have
113
described three inkless octopodids from the continental shelf off south-eastern South America. These octopuses are a non-commercial by-catch in the Falkland (Malvinas)
114
fishery. Muusoctopus eureka (Robson, 1929) is one of two common inkless octopuses, is of medium size, with orange-pink skin, and a distinctive pattern of irregular dark
115
markings, interspersed with white spots visible only in living or freshly dead specimens. Eight specimens caught in our cruise belong to this species. Another 4 specimens also
116
captured during our cruise belong to Muusoctopus longibrachus akambei Gleadall et al., 2010.
117
Two specimens (21-30mm ML) of the Patagonian bobtail squid Semirossia patagonica (Smith, 1881) were caught on the Falkland (Malvinas) shelf by an RMT8 net from
118
the surface to the bottom (Rodhouse et al., 1992). The species have also been collected on the Argentinean shelf (Reid & Jereb, 2005). The records presented here represent an
119
expansion of the geographical range of the species eastwards. According to depth, SST, and SBT (Table 2), our specimens were caught in the STZ, usually located between
120
35° and 45° South, where the average SST changes from about 12 °C to 7-8 °C (Campos et al., 1995). One specimen of the bobtail Austrorossia mastigophora (Chun, 1915)
121
was also captured in this STZ (45° 46' 30'' South - 60° 27' 29'' West). The known geographical distribution of this bobtail squid was western, southern, and eastern Africa
6
122
(from Guinea and Somalia to the Cape of Good Hope), but doubtful in Chilean waters (Reid and Jereb, 2005). Therefore, this is the first time that the species is caught in the
123
western part of the South Atlantic Ocean. The Carol bobtail squid Neorossia caroli (Joubin, 1902) was caught south more frequently than the previous bobtail squids (47° 32'
124
60'' South - 60° 14' 19'' West) and in the deeper, cooler waters (Table 2) of the FMC. This species is represented on the Patagonian slope north of the Falkland (Malvinas)
125
Islands by the subspecies N. caroli jeannae Nesis et al., 2001, while our specimen was recorded much further north than any other until now.
126
The jewel squids Histioteuthis reversa (Verrill, 1880), H. atlantica (Hoyle, 1885), H. bonnelli (Férussac, 1834), and H. eltaninae N. Voss, 1969 caught are common in the
127
area explored by our cruise, showing an already well-known vertical and geographic distribution inside the FMC (Voss et al., 1998). However, this is the most southern record
128
(45° 13' 52'' South - 59° 24' 04'' West) of Histioteuthis arcturi (Robson, 1948), which had previously only been registered between 28° North and 25° South (Voss et al.,
129
1998).
130
To date, the known distribution of the squid Slosarczykonia circumantarctica was off Wilkes Land (Australian Antarctic Territory) and in circumpolar Antarctic waters
131
(Lipinski and Young http://tolweb.org/Slosarczykovia/24102). Our findings extend the geographical distribution area of the species towards the north and east of the SWA,
132
occupying the western border of the FMC (from approximately 46° 54' South - 59° 49' West to 45° 17' South - 59° 33' West), at depths ranging from 881 to 1224 m, and SST
133
and SBT between 11.9 and 2.3 ºC, respectively (Table 2).
134
135
According to the map of the records of B. skolops provided by Young and Roper (http://tolweb.org/Batoteuthidae/19452), our record is the most northern for this species
(44° 30' 22'' - 44° 29' 54'' South).
136
The glacial cranch squid Galiteuthis glacialis (Chun, 1906) is found throughout the circumpolar Antarctic waters where it is one of the most abundant (Nesis, 1987).
137
According to the map given by Young and Mangold (http://tolweb.org/Galiteuthis_glacialis/19572), our records are the first outside circumpolar Antarctic waters, thus
138
expanding the geographical distribution of the species towards the north, at least until 44° 02' 95'' South. SST and SBT seem to indicate that this species was caught along the
139
western border of the FMC.
7
140
Type locality of Graneledone antarctica Voss, 1976 is the Ross Sea (74° 05' 06'' South, 175° 05' 02'' West) (Voss, 1976). However, Kubodera and Okutani (1994)
141
captured 1 specimen at 45° 44' South, 59° 46' West at 858 m depth, which is close to our area of study and within the bathymetric range of our specimens (Table 3). Despite
142
the resemblance of the Kubodera and Okutani specimen to G. antarctica, these authors did not risk identifying it as such. Their argument was that it was captured near the
143
Crozet Islands at such a considerable distance that they hesitate to conclude that the material was conspecific. Nevertheless, the coordinates of capture that the authors give
144
twice (their Table I and on page 214) do not correspond to these islands but to the Patagonian shelf or south-eastern Argentinean EEZ waters, where they also obtained some
145
samples. Our specimens (1 juvenile and 2 immature female, Table 4) share the diagnostic characters of the species. Considering this information, we therefore believe that the
146
species is present off the Patagonian shelf at depths of 1156-1461(Table 3) and from 44º to 47º South. The species was found in the western boundary (59º 25' West) of the
147
FMC, where the SST was 9.7 ºC and the SBT 2.4 ºC (Table 2).
148
The large-tuberculate octopus Graneledone macrotyla Voss, 1976 was for many years only known by type-locality (54° 43' South, 55° 30' West in 1647-2044 m) (Voss,
149
1976, 1988). However, the collection studied by Kubodera and Okutani (1994) extended the known distribution of the species to the north of 45° South along south-eastern
150
Argentina, at depths ranging from 858 to 2044 m, which agrees with the latitudinal distribution shown by the present sample (from 45° to 47° South). The western range
151
where this species was caught (59° 32' - 60° 01' West) and the SST and SBT (Table 2) suggest that it inhabits the northern part of the FMC.
152
Our 4 mature males of Thaumeledone gunteri Robson, 1930 share the main characters given by Allcock et al. (2004). These authors indicated that the distribution of this
153
species is probably restricted to South Georgia and Shang Rocks at a depth of 366-964 m and probably deeper. However, we found this inkless octopus further north, at
154
depths ranging from 855 to 1190 m, with a sea surface temperature of 9.6 ºC, and a sea bottom temperature of 2.6 ºC (Table 3).
155
Laptikhovsky (2001) reported 32 females of Benthoctopus eureka collected on the Falkland (Malvinas) shelf (48° 40' - 52° 59' South, 56° 51' - 60° 15' West, depth range
156
125-344 m) during expeditions on board the RV Dorada. Gleadall et al. (2010) are currently proposing a new taxonomic combination for eureka, which is Muusoctopus
157
eureka (Robson, 1929). The species was caught in the same area explored by our cruise. However, the specimens were mainly on the 200 m depth contour (Gleadall et al.,
8
158
2010), whereas the bathymetric range of the 8 specimens included in this paper was 111 to 1156 m. Therefore, it seems that this species is adapted to a relatively wide range
159
of temperature, possibly from 8 ºC to -0.8 ºC (Tables 2 and 5).
160
Previously known specimens of M. longibrachus akambei (holotype and paratypes) were caught on different cruises around 49° 22' South and between 59° 10' - 66° 70'
161
West from 147 to 377 m depth off the coastal waters of Argentina (Gleadall et al., 2010). The specimens presented in this paper were captured about 140 nautical miles to the
162
north (46° 53' 98'' South), at a similar western longitude (59° 48' - 60° 39' West), and in deeper waters (428-921 m) (see Table 3).
163
From a biogeographic point of view our data show that several species, mainly benthic octopuses (Table 5), penetrate the area studied with the plume of cold sub-Antarctic
164
waters and are pushed far into the South Atlantic by the FMC (Fig. 4). This concurs with observations by Strugnell et al. (2008), who showed that the deep-sea lineage had
165
their evolutionary origins in Antarctica. We suggest that sampling in the near future should be attempted on either side of this plume, in regions where there is no Antarctic
166
water, in order to test the presence of cold water-adapted species.
167
168
169
170
171
Acknowledgements
172
vessel. We are also very grateful to Dr Ian Cleadall for his useful comments on the original manuscript and to María Teresa Fernández Álvarez (IIM, CSIC) for her technical
173
assistance. We would also like to thank the crew of the vessel, the scientific staff, and Raúl Vilela for his assistance with GIS tools.
We would like to extend our warmest thanks to authorities of the Spanish Secretaría General del Mar (SGM, General Secretary for the Sea) who owned the research
174
9
175
References
176
177
178
179
180
181
182
183
184
185
186
187
188
189
190
191
192
193
194
195
Allcock, A.L., Collins, M.A., Piatkowski, U. Vecchione, M., 2004. Thaumeledone and other deep water octopodids from the Southern Ocean. Deep-Sea Res. Part II, 51,
1883-1901.
Agnew, D.J., Baranowski, R., Beddington, J.R., des Clers, S., Nolan, C.P., 1998. Approaches to assessing stocks of Loligo gahi around the Falkland Islands. Fish. Res. 35,
155-169.
Agnew, D., Hill, S.L., Beddington, J.R., Purchase, L.V., Wakeford, R.C., 2005. Sustainability and management of Southwest Atlantic squid fisheries. Bull. Mar. Sci. 76, 579593.
Arkhipkin, A., Grzebielec, R., Sirota, A.M., Remeslo, A.V., Polishchuk, I.A., Middleton D.A.J., 2004. The influence of seasonal environmental changes on ontogenetic
migrations of the squid Loligo gahi on the Falkland shelf. Fish. Oceanogr. 13, 1-9.
Basson, M., Beddington, J.R., Crombie, J.A., Holden, S.J., Purchase, L.V., Tingley, G.A., 1996. Assessment and management techniques for migratory annual squid stocks:
the Illex argentinus fishery in the Southwest Atlantic as an example. Fish. Res. 28, 3-27.
Bazzino, G., Quińones, R.A., Norbis, W., 2005. Environmental associations of shortfin squid Illex argentinus (Cephalopoda: Ommastrephidae) in the Northern Patagonian
Shelf. Fish. Res. 76, 401-416.
Boebel, O., Davis, R.E., Ollitrault , M., Peterson, R.G., Richardson, P.L., Schimid, C., Zenk, W., 1999. The Intermediate Depth Circulation of the Western South Atlantic.
Geophys. Res Let. 26, 3329-3332.
Brunetti, N.E., Ivanovic, M.L., Sakai, M., 1999. Calamares de importancia comercial en la Argentina. Biología, distribución, pesquerías, muestreo biológico. Instituto
Nacional de Investigación y Desarrollo Pesquero (INIDEP), 45 p. Mar del Plata, Argentina.
196
Cooperative Institute for Marine and Atmospheric Studies (CIMAS), University of Miami. http.//oceancurrents.rmas.miami.edu.
197
Campos E.J.D., Jerry, L., Miller, J.L., Müller, T.J. Peterson, R.G., 1995. Physical oceanography of the Southwest Atlantic Ocean. Oceanography 8, 87-91.
198
Deacon, G., 1984. The Antarctic Circumpolar Ocean. Cambridge University Press.
10
199
200
Gleadall, I. G., Guerrero-Kommritz, J., Hochberg Jr, F.G., Laptikhovsky, V.V., 2010. The inkless octopuses (Cephalopoda: Octopodidae) of the Southwest Atlantic. Zool.
Sci. 27, 528–553.
201
Glorioso, P.D., 1987. Temperature distribution related to shelf-sea fronts on the Patagonian shelf. Continental Shelf Research, 7: 27-34.
202
Guerra, A., González, A.F., Cherel, Y., 2000. Graneledone gonzalezi sp. nov. (Mollusca: Cephalopoda): a new octopod from the Îlles Kerguelen. Antarc. Sci. 12, 33-40.
203
Haimovici, M., Brunetti, N.E., Rodhouse, P.G. Csirke, J. Leta H.R., 1998. Illex argentinus.. FAO Fish. Tech. Pap. 376, 27-58.
204
Kubodera, T., Okutani, T., 1994. Eledonine octopods from the Southern Ocean: systematics and distribution. Antarc. Sci. 6, 205-214.
205
Laptikhovsky, V.V., 2001. Fecundity, egg masses and hatchlings of Benthoctopus spp. (Octopodidae) in Falkland waters. J. Mar. Biol. Ass. 81, 267-270.
206
Laptikhovsky, V.V., Arkhipkin , A., 2010. Squid as fish prey and human harvest on the southern part of Patagonian shelf (Falkland Islands). ICES J. Mar. Sci., 106, 151-155.
207
Lipinski, M.R., 2001. Preliminary description of two new species of Cephalopods (Cephalopoda: Brachiteuthidae) from South Atlantic and Antarctic waters. Bull. Sea Fish.
208
Inst. 152, 3-14.
209
Nesis, K.N., 1987. Cephalopods of the world. T. F. H. Publications, Inc., Ltd. Neptune City, USA.
210
Nesis, K.N., 2003. Distribution of recent cephalopoda and implications for Plio-Pleistocene events. Berliner Paläobiol. Abh. 3, 199-224.
211
Nesis, K.N., Arkhipkin, A.I., Nikitina, I.V., Middleton, D.A.J., Brickle, P., 2001. A new subspecies of the bathyal sepiolid cephalopod Neorossia caroli (Joubin, 1902) from
212
213
214
the southwestern Atlantic off the Falkland Islands. Ruthenica 11, 51-56.
Nigmatullin, Ch. N., 1989. Las especies de calamar más abundantes del Atlántico Sudoeste y sinopsis sobre la ecología del calamar (Illex argentinus). Publ. Com. Téc. Mix.
Frente Mar., 5, Sec. A, 71 – 81.
215
Norman, M.D., Hochberg, F.G., 2005. The current state of octopus taxonomy. Phuket Mar. Biol. Center Res. Bull., 66, 127-154.
216
Peterson, R.G., Whitworth III, T., 1989. The Subantarctic and Polar Fronts in relation to deep water masses through the southwestern Atlantic. J. Geophy. Res. 94, 10817-
217
218
219
10838.
Portela, J. M., Pierce, G. J., del Río, J.L., Sacau,M., Patrocinio, T. Vilela, R., 2010. Preliminary analysis of the stock status of Illex argentinus and Loligo gahi on the High
Seas of the SW Atlantic and possible interactions of fishing activities with VMEs in this area. Fish. Res. 106, 229-238.
220
Reid, A., Jereb P., 2005. Family Sepiolidae Leach. In: Jereb,P., Roper, C.F.E. (Eds.), Cephalopods of the world. An annotated and illustrated catalogue of cephalopod species
221
known to date, n. 4, vol. 1, pp. 153-212. Chambered nautiluses and sepioids (Nautilidae, Sepiidae, Sepiadaridae, Idiosepiidae and Sepiolidae. FAO Species Catalogue for
222
Fisheries Purposes. FAO, Rome.
11
223
Reid, J.L., 1989. On the total geostrophic circulation of the South Atlantic Ocean: flow patterns, tracers and transports. Progr. Oceanogr. 23, 149-244.
224
Rodhouse, P.G., 1989. Pelagic cephalopods caught by nets during the Antarctic research cruises of the Polarstern and Walther Herwig, 1985-1987. Archiv. Fisch. Wiss. 39,
225
226
227
228
229
230
231
232
233
111-121.
Rodhouse, P.G., Symon, C., Hatfield, E.M.C., 1992. Early life cycle of cephalopods in relation to the major oceanographic features of the southwest Atlantic Ocean. Mar.
Ecol. Progr. Ser. 89, 183-195.
Sacau, M., Pierce, G.J., Wang, J., Arkhipkin , A.I., Portela, J., Brickle, P., Santos, M.B., Zuur, A.F., Cardoso, X., 2005. The spatio-temporal pattern of Argentine shortfin
squid Illex argentinus abundance in the southwest Atlantic. Aquat. Living Resour. 18, 361-372.
Strugnell, J.M., Rogers, A.D., Prodöhl, P.A., Collins, M.A., Allcock, A.L., 2008. The thermohaline expressway: the Southern Ocean as a centre of origin for deep-sea
octopuses. Cladistics 24, 853–860.
Vecchione, M., Allcock, A. L., Piatkowski, U., 2005. Unusual incirrate octopods from the South Shetland Islands, Antarctica, including Bathypurpurata profunda, a newly
discovered genus and species of deepwater pygmy octopod (Cephalopoda). Phuket Mar. Biol. Center Res. Bull. 66, 109-115.
234
Voss, G.L., 1976. Two new species of octopods of the genus Graneledone (Mollusca: Cephalopoda) from the Southern Ocean. Proc. Biol. Soc. Wash. 88, 447-458.
235
Voss, G.L., 1988. The Biogeography of the Deep-Sea Octopoda. Malacologia, 29, 295-307.
236
Voss, N.A., 1980. A generic revision of the cranchiidae (Cephalopoda: Oegopsida). Bull. Mar. Sci. 30, 365-412.
237
Voss, N.A., Nesis, K.N., Rodhouse, P.G., 1998. The cephalopod family Histioteuthidae (Oegopsida): systematics, biology and biogeography. Smithson. Cont. Zool. 586, 293-
238
372.
239
Wilson, H.E., Rees, N.W., 2000. Classification of mesoscale features in the Brazil-Falkland Current confluence zone. Prog. Oceanogr. 45, 415-426.
240
Young, R.E., Roper, C.F.E., 1968. The Batoteuthidae, a new family of squid (Cephalopoda: Oegopsida) from Antarctic waters. Antarc. Res. Ser. 11, 185-202.
241
242
243
244
12
245
246
247
248
249
250
Table 1
Characteristics of depth strata and number of hauls per stratum in ATLANTIS 2009 cruise. Meters (m); nautical miles (nm), square nautical miles (nm2)
Stra
tum
251
252
1
2
3
4
5
6
7
8
9
10
11
12
13
TOT
AL
Depth
range
(m)
<200
201-300
301-400
401-500
501-700
701-1000
1001<200
1500
201-300
301-400
401-500
501-1000
10011500
Surface
(nm2)
No of grids
(~5 nm2)
Scheduled
hauls
1,144
279
366
538
1,483
1,964
2,037
1,395
111
123
74
977
2,547
13,038
229
56
73
108
297
393
407
279
22
25
15
195
509
2,608
13
3
4
6
17
22
22
16
2
2
2
11
29
149
13
Valid
Null
13
3
4
1
Hauls made
6
17
2
21
1
18
1
2
2
2
11
28
127
5
253
Table 2
254
255
256
257
258
ATLANTIS 2009 cruise typical range of Sea Surface Temperature (SST), Sea Bottom Temperature (SBT), and salinity at the different zones or currents mentioned in the text
and Figure 2. PS: Patagonian shelf; STZ: Subtropical Zone or Subtropical Front; FMC: Falkland (Malvinas) Current (at 52-53.3ºS-58-61º W); SAZ: Subantartic Zone; PFZ:
Polar Frontal Zone; AZ: Antarctic Zone. S: Summer; W: Winter. Data sources: CIMAS (Cooperative Institute for Marine and Atmospheric Studies, University of Miami;
Arkhipkin et al., 2004; Deacon, 1984; Glorioso, 1987; and Peterson and Whitworth, 1989). These data vary with time of the year and weather conditions, when there is no
indication the information correspond to March-April months.
259
260
261
Zone
SST°C
PS
7.5 - 18.0
STZ
7.0-14.0
FMC
9.5-14.6
SAZ
4.0-14.0 S
4.0-8.0 W
PFZ
2.0-4.0
AZ
2.0-3.0
SBT
13.0-14.0 (450 m)
4.0-5.0 (200 m)
4.5-8.0 (200 m)
2.2-3.8 (1,000 m)
2.0-3.0
2.0 - 0.0
(1,000 m)
-0.2-0.4
(2,000 m)
Salinity
34.57 - 35.70
34.90-34.60
33.8-34.2
33.90 - 34.90 W
>33.0 S
33.80-34.10 W
>33.0 s
33.50 - 34.70 W
>33.0 S
14
262
263
264
265
266
Table 3
Cephalopod species caught during the ATLANTIS 2009 survey in March 2009, with number of hauls (NH) where they were caught, depth range (DR), Maximum Sea Surface
Temperature (SST) and Sea Bottom temperature (SBT), and number of specimens caught (NC).
267
Species
NH
DR (m)
Semirossia patagonica
2
114-144
SST-SBT (º
C)
13.4-4.1
N
C
2
Austrorossia
mastigophora
1
116-119
13.5-5.9
1
Neorossia caroli
1
601-603
12.0-3.8
1
Gonatus antarcticus
14
6901502
10.5-2.2
Histioteuthis arcturi
1
11761187
Histioteuthis atlantica
8
Histioteuthis bonnellii
Histioteuthis eltaninae
Species
Onykia ingens
Bathyteuthis
abyssicola
N
H
1
DR (m)
10431050
SST-BT
(ºC)
9.8-2.9
N
C
1
1
14971502
9.9-2.2
1
Batoteuthis skolops
1
11981221
9.8.2.8
1
23
Galiteuthis glacialis
2
11031365
10.5-2.6
3
9.5-2.7
1
Taonis pavo
1
13591365
10.2-2.5
1
135-998
9.8-2.8
10
Graneledone
antractica
3
11561461
9.7-2.4
3
5
6251291
11.0-2.3
5
Graneledone
macrotyla
2
774-870
10.3-3.1
2
1
717-720
12.9-3.6
4
Thaumeledone
gunteri
4
8551190
9.6-2.6
4
15
268
Histioteuthis reversa
3
618-846
13.1-3.0
3
Chiroteuthis veranyii
3
766-1187
12.0-2.7
3
Slosarczykovia
circumantarctica
6
881-1224
11.9-2.3
9
Muusoctopus eureka
8
1111156
14.6-2.8
8
3
428-921
12.0-2.9
4
269
M.
akambei
Totals
16
longibrachus
71
90
270
271
272
273
274
275
Table 4. Cephalopod species caught during ATLANTIS 2009 cruise. Family; size range (ML: dorsal mantle length in millimeters); BD: biological data; M: male; F: female;
SA: subadult; J: juvenile); NC: number of specimens caught; Ma: mature; Mat: maturating; IM: immature; Sfm: stomach full of myctophids; Sfe: stomach full of euphausids.
Species
Semirossia patagonica
Family
Sepiolidae
ML
21-30
BD
Ma M+F
NC
2
Austrorossia mastigophora
Sepiolidae
21
Ma F
1
Neorossia caroli
Sepiolidae
47
Ma F
Gonatus antarcticus
Gonatidae
50-226
Histioteuthis arcturi
Histioteuthidae
Histioteuthis atlantica
Family
Onychoteuthidae
ML
480
NC
1
52
BD
MA F
Sme
M
Bathyteuthis abyssicola
Bathyteuthidae
1
Batoteuthis skolops
Batoteuthidae
125
J
1
23
21
9 Ma M
10 Mat F, 4J
Sfm
J
Galiteuthis glacialis
Cranchiidae
350-372
J and SA
3
1
Taonis pavo
Cranchiidae
323
SA
1
Histioteuthidae
43-79
6M +4F
10
Graneledone antractica
Octopodidae
21-130
3
58-82
2M+3F
5
Graneledone macrotyla
Octopodidae
27-60
Histioteuthidae
66-75
1M+3F
4
Thaumeledone gunteri
Octopodidae
35-51
2 In F
1J
1 Ma M
1J
Ma M
Histioteuthis bonnellii
Histioteuthidae
Histioteuthis eltaninae
Histioteuthis reversa
Histioteuthidae
21-83
2M+1F
3
Muusoctopus eureka
Octopodidae
49-80
8
Chiroteuthis veranyii
Chiroteuthidae
84-297
1F+2J
3
Muusoctopus longibrachus akambei
Octopodidae
42-78
5 Ma M
3 In F
2 Ma M
2 Mat F
Brachioteuthidae
105-182
4 Ma M
2 Ma F, 3 J
9
Total
Slosarczykovia circumantarctica
276
277
278
279
280
17
Species
Onykia ingens
1
2
4
4
90
281
282
283
284
285
286
Species
P
S
287
288
289
290
291
292
293
294
295
296
297
298
299
300
301
302
303
304
305
306
307
STZ
FMC
SAZ
PF
Z
AZ
Authority
PS
ST
FM
Z
C
SAZ
PFZ
AZ
Authority
Table 5
Cephalopod distribution in relation to Subtropical Zone (STZ), Patagonian shelf (PS), Falkland/Malvinas Current (FMC), Subantartic Zone (SAZ), Polar Frontal Zone (PFZ),
Antarctic Zone (AZ). X: previous papers: P: present paper.
18
Semirossia
patagonica
X
Austrorossia
mastigophora
P
X P
X
XP
Histioteuthis arcturi
X
X
X
X
X
P
X
X
X
P
X
X
X
Nesis et al., 2001
Batoteuthis skolops
X P
X
X
X
Rodhouse et al.
1999; Nesis, 2003
Galiteuthis glacialis
P
X
X
X
XP
Histioteuthis
atlantica
Onykia ingens
Bathyteuthis
abyssicola
P
Neorossia caroli
jeannae
Gonatus antarcticus
Rodhouse et al.
1999; Reid and
Jereb, 2005
X
Taonis pavo
P
X
X
X
N. Voss, 1980
X?
X
X
Voss,
1976;
1988;
Kubodera
&
Okutani, 1994
Voss,
1976;
1988;
Kubodera
&
Okutani, 1994
Allcock et al.,
Rodhouse et al.
1999
Graneledone
antractica
P
XP
N. Voss, et al.,
1998; Nesis, 2003
Graneledone
macrotyla
P
Histioteuthis
eltaninae
XP
Rodhouse et al.
1999
Thaumeledone
gunteri
P
Histioteuthis reversa
XP
N. Voss et al.,
1998
Muusoctopus eureka
Nesis, 1987
Muusoctopus
longibrachus
akambei
Histioteuthis
bonnellii
Chiroteuthis veranyii
X
X
X
P
X
19
Young
&
Roper, 1968;
Rodhouse
et
al.,
1999;
Nesis, 2003
Nesis,
1987;
2003
XP
X
Nesis,
1987;
Vecchione et
al., tolweb
Nesis, 1987
X
X?
X?
X
2004
X P
X
X
X
Geadall et al.,
2010
X P
Geadall et al.,
2010
Slosarczykovia
circumantarctica
P
X
X
Lipinski, 2001;
Lipinski & Young
tolweb
308
309
310
311
312
313
314
315
316
317
318
20
319
320
321
Fig. 1. Study area comprised between 44º and 48º south and to the east of the Argentinean Exclusive Economic Zone (EEZ or ZEE in Spanish). FOCZ: Falkland Islands Outer
Conservation Zone; FICZ: Falkland Islands Conservation Zone.
21
322
323
324
325
326
327
Fig. 2. Location of the cephalopod trawled species in ATLANTIS 2009 cruise. Semirossia patagonica (Sp); Austrorossia mastigophora (Am); Neorossia caroli (Nc); Gonatus
antracticus (Ga); Histioteuthis arcturi (Ha); H. atlantica (Hat); H. bonnellii (Hb); H. eltaninae (He); H. reversa (Hr); Chiroteuthis veranyii (Cv); Slosarczykovia
circumantarctica (Sc); Onykia ingens (Oi); Bathytheuthis abyssicola (Ba); Batoteuthis skolops (Bs); Galiteuthis glacialis (Gg); Taonius pavo (Tp); Graneledona antractica
(Gan); Granleledone macrotyla (Gm); Taumeledone gunteri (Tg); Muusoctopus eureka (Me); Muusoctopus longibrachus akambei (Mla).
22
328
329
330
Fig. 3. Major oceanographic features of the southwest Atlantic Ocean (from Rodhouse et al., 1992). The limits of the fronts and zones change with time of the year and
climatic conditions.
331
332
23
333
334
Fig. 4. Falkland/ Malvinas Current in the January- March period (black arrows enclosed in white). Source: CIMAS, University of Miami, Rosenstiel School of Marine and
Atmospheric Science, USA.
335
336
337
338
24