Checklist and Account of the lophogastrida (Crustacea: Peracarida) of the Canary Islands, with notes of taxonomy and biogeography of the species

              Rev. Acad. Canar. Cienc. Vol. XXIV, 63-80 (diciembre dc 2012)
CHECKLIST AND ACCOUNT OF THE LOPHOGASTRIDA
(CRUSTACEA, PERACARIDA) OF THE CANARY ISLANDS,
W ITH NOTES ON TAXONOMY
AND BIOGEOGRAPHY OF THE SPECIES
Wittmann, K.J.' & R. Riera *
Abteilung tiir Okotoxikologie, Zentrum fiir Public Health, Medizinische Uni\ersitat W ien
Kinderspitalgasse 15, A- 1090 Vienna, Austria
- Centre de Investigaciones Medioambientales del Atlantico (CIMA SL)
Arzobispo Elias Yanes. 44. 38206 La Laguna, Tenerife. islas Canarias
*corresponding author: rodrigo(a cimacanarias.com
RESUMEN
Las aguas que rodean el archipielago canario constituyen las de mayor riqueza de espe-cies
de lofogastridos en el Oceano Atlantico, con doce especies meso y batipelagicas de este
grupo. Esta di\ersidad es debida a la ele\ada intensidad de nuestros conjuntamente con las fa-vorables
condiciones marinas. Todas las especies citadas son propias de esta zona, aunque el
grado de endemicidad es muy bajo. Diez especies son consideradas cosmopolitas y dos presen-tan
una distribucion restringida al area Atlantica, encontrandose limitada una de ellas al Atlantico
Este. Ademas. se analizan a ni\'el biogeografico las areas de distribucion de las especies canarias
de lofogastridos con zonas adyacentes (Macaronesia, Mediterraneo y Costa Atlantica Africana).
Palabras clave: Crustacea. Lophogastrida, plancton. mesopelagico, batipelagico, islas
Canarias. Oceano Atlantico
ABSTRACT
The species numbers of lophogastrids in open waters off E. Atlantic islands (Azores,
Madeira. Canaries, and Cape Verdes) exceed those in waters off N.W. African continental
coasts and in the Mediterranean Sea. With twelve currently known, mainly meso- to bathy-pelagic
species, the waters off the Canary Islands appear to be the most species-rich in the At-lantic
Ocean. This comparatively large number may reflect high sampling intensity in
combination with favourable environmental conditions. All the here documented species ap-pear
to be indigenous, but the degree of endemism is very low: ten Canarian species are cos-mopolitans,
two are endemic to the Atlantic, only one of the latter two is restricted to the E.
Atlantic. Biogeographical data are provided for the lophogastrids from the Canary Islands
and surrounding areas (Macaronesian region, Mediterranean and Atlantic African coasts).
Key words: Crustacea, Lophogastrida, plankton, mesopelagic, bathypelagic, Canary
Islands, Atlantic Ocean
63
INTRODUCTION
The present contribution critically evaluates the literature data along with original
records of lophogastrid species in the framework of large-scale surveys on the marine biodi-versity
of the Canary archipelago. Consideration is also given to the species diversity in open
waters off the Canaries versus that off other E. Atlantic archipelagos, off W. African coasts,
and in the Mediterranean Sea. The literature data for the Gulf of Mexico is provided for com-parison.
Finally, the importance of potential endemics as opposed to pan-thalassic cos-mopolitans
for local faunas is evaluated.
The lophogastrids are shrimp-like marine crustaceans with stalked eyes. They are char-acterized
by an elongate carapace fused with anterior thoracic segments but not with some of
the terminal ones. Eight pairs of biramous thoracic legs are present, all or only some equipped
with gills. Thoracic exopods are feebly to moderately developed, supporting the respiratory
current and swimming; the endopods are variously specialized, such as for filter feeding and
grasping prey. Females carry eggs and larvae in a brood pouch below the thorax. A marsupium
is formed by seven pairs of oostegites. The five pairs of pleopods act as strong swimming
legs; all are biramous and well developed in both sexes, without sexual dimorphism. No sta-tocyst
is present in the tail fan.
Members of this order occur in all oceans except the Arctic Ocean. They are generally
meso- to bathypelagic (mostly > 1 ,000 m depth), but a few—mostly bentho-pelagic—species
are common in shallower waters (< 200 m depth, e.g. many populations of Lophogaster typ-icus
M. Sars, 1857). Interestingly, this group shows more species in the Pacific and Indian
Oceans than in the Atlantic Ocean. Currently, a total of 50 living species plus one non-nomino-typical
subspecies, belonging to eight genera and three families (Lophogastridae,
Gnathophausiidae, and Eucopiidae), are recognized for the Lophogastrida (Anderson [2]).
MATERIALS AND METHODS
Body length was measured from tip ofrostrum to terminal margin oftelson without spines.
Most material studied was from pelagic and also epibenthic samples taken by the expeditions
listed below. For additional sampling campaigns see Wittmann et al. [71]. From this extensive ma-terial,
only those species are treated here in detail that were (also) confirmed for the Canaries.
Expeditions CANCAP-I (Madeira, 1976), CANCAP-II (Canaries, Morocco, 1977), CAN-CAP-
III (Madeira, Mauritania, 1978), CANCAP-V (Azores, 1981), and CANCAP-VII
(Cape Verde, 1986), and TYRO Mauritania-II (1988). Expeditions organized and material
deposited in the National Natural History Museum (Rijksmuseum) Leiden.
Expedition ANTARKTIS-XXIII/2 organized by the Alfred-Wegener-Institut fur Polar- und
Meeresforschung, Bremerhaven: Pelagic samples taken in the S. Atlantic and Antarctic
during southern summer 2005/06 with the vessel PES Polarstem, leg. Ute Miihlenhardt-
Siegel. Material at the Zoological Museum Hamburg.
Expeditions Kana Koeki, Norpax Equat. Himb. (tropical Pacific, 1977-1979), Dominator
Cruise 021 (tropical Pacific, 1982), and Sargasso Eel & Maine, Cruise 83 (N.W. Atlantic,
1983). Material determined for the Smithsonian Oceanographic Sorting Center.
Cruise 113 by FS "Poseidon" (off Portugal, 1984). Material at the Senckenberg Museum,
Frankftirt am Main; coll. n° 39206-39212.
64
RESULTS
Order Lophogastrida G.O. Sars, 1870
Family Eucopiidae G.O. Sars, 1885
Eucopia australis Dana, 1852
Eiicopia australis Dana [13]: 609-611, PI. 40 (lOa-m): Caiman [5]: 15: Illig [35]: 404-405:
Nouvel [45]: 26-27, PI. 1(17-19); Tattersall [60]: 48-49, Fig. 4C-D; Wittmann [70]: 130,
Fig. 257; Wittmann & Wirtz [73]; Haroun & Garrido [30]; Wittmann et al. [71]; Fukuoka
[23]: 407-408; Price £^r^//. [52].
Material studied: S. Atlantic and Antarctic (all samples from expedition ANTARKTIS-XXIII/2): 1
1
spec, N of Bouvet Island, sta. 30-1, combined rectangular midwater trawls (RMT 1 and RMT 8), tow
from 51.01 17S 6.1283E to 51.1783S 6.3800E, 3.130-1,000 m, bottom 3,738 m, 25 Nov. 2005.- 2 spec,
off Bouvet Island, sta. 34-1, RMT 8, tow from 55.0367S 2.98 1 7E to 54.9 150S 2.83 17E, 1,000-2.429 m,
bottom 2.860 m. 26 Nov. 2005.- 6 spec, Lazarev Sea. sta. 93-5. combined RMT 1 and RMT 8. tow
from64.9950S2.9867Wto64.8083S2.8300\V. 1.000-3.003 m. bottom 3.270 m. 23 Dec 2005.- 1 spec.
Antarctic, NNW of Atka bay. sta. 127-1, RMT 8, tow from 69.4600S 8.9567W to 69.4083S 9.21 17W.
1,000 m, bottom 2,850 m. 2 Jan. 2006.- 2 F ad.. 1 M ad.. 1 F subad.. 3 imm.. 3 juv.. Antarctic. NNW of
Atka bay, sta. 127-2. RMT 8. tow from 69.4083S 9.2183W to 69.2883S 9.6917W, 1,000-2,500 m, bot-tom
3.400 m. 2 Jan. 2006.
Distribution.- Depth range: 600-6,000 m. Cosmopolitan, but not found in Arctic waters. Well
known from Antarctic, S. Atlantic, and Indian Oceans according to Ortmann [47, 48], Hansen
[27], Zimmer [76], and Illig [35]. These samples confirm previous records from the S. At-lantic
and Antarctic. In addition, this species was recorded by Caiman [5] for deep waters SW
off Ireland; by Nouvel [45] for the Azores, based on two juveniles only; and by Tattersall [60]
for waters off Cape Verde Islands. Given in faunal lists by Haroun &. Garrido [30] for the Ca-naries
and by Price et al. [52] for the Gulf of Mexico.
Eucopia grimaldii H. Nouvel, 1942
Eucopia Grimaldii Nouvel [44]: 5-6, Figs. 5-8; [45]: 40-43. PI. II (36-40).
Eucopia grimaldii - Tattersall & Tattersall [63]: 106-108. Fig. 11a; Tattersall [60]: 54-56. Fig.
5; [61]: 146; Casanova [7]: 20-22. 385. Fig. 6; Lagardere [37]: 812-813; Casanova £^/ ^/. [8^:
60-61, Figs. 2-3; Wittmann & Wirtz [73]; Hargreaves [29]: 12, Tabs. 2, 4-6; Haroun & Gar-rido
[30]; Wittmann et al. [71]; Fukuoka [23]: 408-409; Price et al. [52]; San Vicente [53].
Eucopia grimaldi - Vereshchaka [65]: 118.
Material studied: N.W. Atlantic- 2 F ad., body length 25.2-27.9 mm. 1 imm.. NW of Bemiuda. 32.41 7N
64. 983 W. 1.000-2.000 m. 13 Sept. 1989. leg. Teddy Tucker: Bemiuda Aquarium. Natural Histor\ Mu-seum
and Zoo.
Distribution.- Depth range: 300-3.000 m. Cosmopolitan, but not found in the Arctic and
Antarctic Oceans. Recorded by Nouvel [45] from Azores, Canaries, and off Brittany. Reported
65
by Tattersall & Tattersall [63] for waters offAzores and Madeira, and by Tattersall [60] for wa-ters
off Cape Verde Islands.
Eucopia major Hansen, 1910
Eucopia w^yor Hansen [26]: 21, PL I (4a-b); Illig [35]: 405; Nouvel [45]: 28-29, PI. I (20-25);
Tattersall [62]: 33; Springer & Bullis [59]; Fumestin [24]: 1 8 1 ; Lagardere [37]: 812-813;
Wittmann & Wirtz [73]; Haroun & Garrido [30]; Wittmann et al. [71]; San Vicente [53].
Material studied: Canary Islands.- 1 M ad., body length 66.8 mm, SW of Fuerteventura, Punta de Mor-rojable,
27.967N 14.200W, CANCAP-II, sta. 2.68, ring-trawl, 1,700 m, bottom 1,810 m, 29 Aug. 1977,
22.20-00.55 hrs.- 1 F imm. 37. 1 mm, SW of El Hierro, off Punta de Orchilla, 27. 600N 1 8.267W, CAN-CAP-
II, sta. 2.140, ring-trawl with weight, 2,000 m, bottom > 2,800 m, 9 Sept. 1977, 3.15-6.15 hrs.
Distribution.- Depth range 1000-5000 m. Boreal to tropical waters of the Atlantic, Indian, and
Pacific Oceans. Recorded by Nouvel [45] from Golfe de Gascogne, Canaries, and off the
coast of Morocco. Indicated by Furnestin [24] with query for the Gulf of Lion (Mediter-ranean).
The present resuhs confirm the previous records from the Canaries.
Eucopia sculpticauda Faxon, 1893
Eucopia sculpticauda Faxon [21]: 218; [22]: 219-221, Pis. K (2, 2d), LIII (1-ld); Hansen
[25]: 7, Fig. 4; Illig [35]: 400-403, Figs. 1-6; Fage [20]: 56-60, Figs. 40-42; Nouvel [45]:
22-26, PI. I (14-16); Tattersall & Tattersall [63]: 109-112, Figs. 12-13; Tattersall [60]: 52-
54; [61]: 146; Casanova [7]: 300-302, 387-389, Fig. 86, Tab. 52; Lagardere [37]: 813-
814; Wittmann & Wirtz [73]; Haroun & Garrido [30]; Wittmann et al [71]: 1262, Tab. I;
Price era/. [52].
Eucopia intermedia Hansen [25]: 5-7, Figs. 2-3.
Material studied: Madeira.- 1 F ad., body length 42.8 mm, S of Porto Santo, 32.883N 16.367W, CAN-CAP-
III, sta. 3.021, ring-trawl with weight, wire 2,500 m, bottom 2,580-3,220 m, 16 Oct. 1978.- 1 imm.
29.6 mm, S of Madeira, 32.500N 16.817W, CANCAP-I, sta. 116, ring trawl, wire 500 m, bottom 2,320-
2,420 m, 18 Mar. 1976.- 1 M ad. 40.3 mm, S of Madeira, 32.367N 16.917W, CANCAP-III, sta. 3.046,
ring-trawl with stone, wire 2,000 m, bottom 3,160-3,200 m, 19 Oct. 1978.- 1 F ad. 47.1 mm, SE of
Madeira, 32.450N 16.883W, CANCAP-I, sta. 1.105, ring-trawl, wire 1250 m, bottom 2,500-2,650 m, 17
Mar. 1976.- 1 F ad. 42.8 mm, S of Porto Santo, 32.883N 16.367W, CANCAP-III, sta. 3.021, ring-trawl
with weight, wire 2,500 m, bottom 2,580-3,220 m, 16 Oct. 1978.
Morocco.- 1 M ad. 40.0 mm, W of Cape Yubi, 28.017N 13.367W, CANCAP-II, sta. 2.52, ring-trawl
with weight, 400 m, bottom 570 m, 00.30-01.40 hrs, 28 Aug. 1977.
Canary Islands.- 1 imm. 12.5 mm, SW of Fuerteventura, Punta de Morrojable, 27.967N 14.200W, CAN-CAP-
II, sta. 2.68, ring-trawl, 1,700 m, bottom 1,810 m, 22.20-00.55 hrs, 29 Aug. 1977.- 1 juv. 11.3 mm,
sampling data as before, 8 Sept. 1994.- 1 F ad. 48.0 mm, SW of El Hierro, off Punta de Orchilla, 27.667N
18.167W, CANCAP-II, sta. 2.116, ring-trawl with weight, 1,000 m, bottom 2,500 m, 2.20-3.40 hrs, 5
Sept. 1977.- 2 F ad. 48.2-53.4 mm, 1 juv. with head missing, SW of El Hierro, off Punta de Orchilla,
27.600N 18.267W, CANCAP-II, sta. 2.140, ring-trawl with weight, 2,000 m, bottom > 2,800 m, 3.15-
6.15 hrs, 8 Sept. 1977.- For additional materials see Wittmann et al. [71].
Mauritania.- 1 F ad. 50.0 mm in 2 parts, off Banc d'Arguin, 19.717N 17.500W, TYRO Maurita-nia-
II, sta. MAU.104, 3.5 m Agassiz trawl in 1,500 m, 17 June 1988.
66
Cape Verde Islands.- 1 F ad. 56.8 mm. S of Santiago, 14.85N 23.25 W, CANCAP-VII, sta. 7.001,
midwater trawl, 0-750 m, 1 8 Aug. 1 986.- 1 F ad. 32.2 mm, 2 M ad. 30.0-30.6 mm, S of Branco, 1 6.5 1 7N
24.81 7W, CANCAP-VII, sta. 7.148, midwater trawl. 0-1,100 m. 4/5 Sept. 1986.
Tropical Pacific.- 1 F ad. 52 mm, tow from 4.0 17N 1 50.1 83W to 4.183N 150.216W, 1.100 m,
Norpax Equat. Himb.. sta. 77-12-16, 1 1 Dec. 1977.
Distribution.- Depth range: 400-6,000 m. Cosmopolitan, but not found in the Arctic and
Antarctic Oceans. Recorded by Illig [35] and Tattersall [60] for waters off Cape Verde Is-lands;
by Page [20] and Nouvel [45] from Azores, Madeira, Canaries, and off the coasts all
along western Africa, from waters near Iceland, Hebrides, off Mexico and the Cape of Good
Hope. The present results fit well with this distributional scheme.
Eiicopia ungiiiculata (Willemoes-Suhm, 1875)
Chalaraspis imguiciilata Willemoes-Suhm [67]: 37-43, PI. VIII.
Eiicopia imguiciilata - Colosi [11]: 2; Illig [35]: 403-405; Tattersall & Tattersall [63]: 101-106.
Figs. 9-1 1; Tattersall [60]: 50-52, Fig. 4A-B; Springer & Bullis [59]; Wittmann & Wirtz
[73]; Haroun & Garrido [30]; Nikoforos [40]; Wittmann et al. [71]: 1263. Tab. I; Price et
al. [52]; San Vicente [53].
Eiicopia Hanseni Nouvel [44]: 3, Figs. 1-4; Fage [20]: 47-56, Fig. 36; Nouvel [45]: 30-40, PI.
I (26-35).
Eiicopia hanseni - Fumestin [24]: 181; Hoenigman [31]: 606-607, Fig. 3; Casanova [7]: 20-
38, 201, Figs. 5-8, 48, 53, 55, 58, 70; Lagardere & Nouvel [38]: 384; Lagardere [37]: 811-
812; Cartes & Sorbe [6]: 191, 194; Tabs. 1-2; San Vicente [53].
Material studied: N.W. Atlantic- 1 M ad., body length 27.7 mm, NW of Bemiuda, 32.41 7N 64.983W.
1000-2000 m, 13 Sept. 1989, leg. Teddy Tucker; Bermuda Aquarium, Natural History Museum and Zoo.
Mediterranean.- 1 fragmented subad. ca. 20 mm, Levantine Sea, S of Crete, tow with trawl from
34.383N 26.033E to 34.383N 26.087E, 4 1 78-4390 m, FS. Meteor cmise 5. section 1 , sta. M5-22Ku. 1
8
Jan. 1987, leg. Senckenberg Institut Wilhelmshafen.
Madeira.- 1 F ad. 24.0 mm, 2 F subad., 2 F num., S of Porto Santo, 32.883N 16.367W, CANCAP-III,
sta. 3.021, ring-trawl with weight, wire 2,500 m, bottom 2,580-3,220 m, 16 Oct. 1978.- 6 imm. ju\., S
of Porto Santo, 32.883N 16.400W, CANCAP-III, sta. 3.035, ring-trawl with weight, wire 1,000 m, bottom
3,024-3,196 m, 17 Oct. 1978.- 2 F imm. 21.5-25.1 mm, SE of Madeira, 32.583N 16.733W, CANCAP-1.
sta. 1.035, ring-trawl, wire 1,250 m, bottom about 2,000 m, 10 Mar. 1976.- 1 imm. 13.1 mm. S of Madeira.
32.567N 16.850W, CANCAP-I, sta. 88, ring-trawl, wire 1,500 m, bottom 1,920-2,060 m, 16 Mar. 1976.-
1 M29.3mm, 1 imm., S of Madeira, 32.51 7N 16.883W, CANCAP-I, sta. 1.1 15, ring-trawl, wire 1,000 m,
bottom 2,350-2.400 m, 18 Mar. 1976, 22.20-00.55 hrs.- 1 F subad. 27.7 mm. 1 M subad. 27.0 mm. S of
Madeira, 32.5N 16.9W, CANCAP-I, sta. 89, ring-trawl, wire 2.000 m. bottom 2.200-2.370 m. 16 Mar.
1976.- 4 F ad. 25.3-28.8 mm, S of Madeira, 32.500N 16.817W, CANCAP-I, sta. 116, ring trawl, wire
500 m, bottom 2,320-2,420 m, 1 8 Mar. 1 976.- 1 M ad. 30.0 mm, 2 F ad. 27.6-28.5 mm, 1 F subad.. 1 imm..
SE of Madeira, 32.450N 16.883W, CANCAP-I, sta. 1.105, ring-trawl, wire 1,250 m, bottom 2,500-2,650 m.
17Mar. 1976.- 1 Fad. 25.1 mm, 8 F subad.,4 imm., 6juv., S of Madeira. 32.367N 1 6.91 7W, CANCAP-III,
sta. 3.046, ring-trawl with stone, wire 2,000 m, bottom 3,160-3,200 m, 19 Oct. 1978.
Selvagens Islands.- Samples listed in Wittmann et al. [71].
Morocco.- 7 imm. 15.3-20.4 mm. 10 juv.. W of Cape Yubi. 28.033N 13.450W, CANCAP-II. sta.
2.42, ring-trawl with weight, 750 m, bottom 1.000 m, 27 Aug. 1977, 03.50-05.30 hrs.- 3 F ad. 20.0-26.2
mm, 1 Mad. 21.3 mm, 1 F subad., 2 imm., W of Cape Yubi, 28.01 7N 13.367W, CANCAP-II. sta. 2.52.
ring-trawl with weight, 400 m, bottom 570 m, 28 Aus. 1977. 00.30-01.40 hrs.
67
Canary Islands.- 1 F ad. 28.7 mm, 2 M ad. 27.2-31.1 mm, SW of Fuerteventura, Punta de Mor-rojable,
27.967N 14.200W, CANCAP-II, sta. 2.68, ring-trawl, 1,700 m, bottom 1,810 m, 29 Aug.
1977, 22.20-00.55 hrs.- 1 F ad. 28.6 mm, S of Fuerteventura, Punta Jandia, 27.783N 14.400W, CAN-CAP-
II, sta. 2.06, abyssal plain at 2,050 m depth, 2.4 m Agassiz trawl, 23 Aug. 1977.- 1 M ad. 30.
1
mm, 5 F ad. 25.5-31.2 mm, 1 F subad., 2 imm., 2 juv., SW of El Hierro, off Punta de Orchilla,
27.600N 18.267W, CANCAP-II, sta. 2.140, ring-trawl with weight, 2,000 m, bottom > 2,800 m, 8
Sept. 1977, 3.15-6.15 hrs.- 1 F imm. 25.6 mm, 1 imm. 25.5 mm, SW of El Hierro, off Punta de Or-chilla,
27. 550N 18.167W, CANCAP-II, sta. 2.151, ring-trawl with weight, 2,500 m, bottom 3,000
m, 10 Sept. 1977, 2.15-6.00 hrs.- 1 M ad. 26.2 mm, SW of El Hierro, off Punta de Orchilla, 27.
350N 18.0I7W, CANCAP-II, sta. 2.164, ring-trawl, 0-2,000 m, bottom 2,700 m, 10 Sept. 1977,
21.14-00.45 hrs.
Mauritania.- Fragments of 1 F ad. and 1 F subad., off Mauritania, 18.850N 1 6.933W, TYRO Mau-ritania-
II, sta. MAU.041, 3.5 m Agassiz trawl, 800-840 m, 10 June 1988.
Cape Verde Islands.- 1 F ad. 27.6 mm, 1 F imm., S of Branco, 16.517N 24.817W, CANCAP-VII,
sta. 7.148, midwater trawl, 0-1,100 m, 4/5 Sept. 1986.- 2 F ad. 25.0-28.2 mm, N of Ilheu Grande,
15.133N 24.583W, CANCAP-VII, sta. 7.035, midwater trawl, 0-2,000 m, 23/24 Aug. 1986.- 1 M ad. 32.8
mm, 4 F subad. 26.2-29.3 mm, S of Santiago, 14.85N 23.25W, CANCAP-VII, sta. 7.001, midwater
trawl, 0-750 m, 18 Aug. 1986.
S. Atlantic- 3 spec, N of Bouvet Island, ANTARKTIS-XXIII/2, sta. 30-1, combined rectangular
midwater trawls (RMT 1 and RMT 8), tow from 5 1.0 11 7S 6.1283E to 5 1.1 783 S 6.3800E, 1,000-3,130
m, bottom 3,738 m, 25 Nov. 2005.
Distribution.- Depth range: 100-6,500 m. Mostly in tropical to temperate waters of the Pa-cific,
Indian, and Atlantic Oceans, also in boreal and subarctic waters; absent in the Arctic
and Antarctic Oceans. Recorded by Colosi [10, 11] from the Bosporus (Mamiora Sea) and the
Gulf of Naples (Mediterranean), and by Illig [35] from waters off Madeira, Mauritania, and
Senegal. Reported by Fage [20] and Nouvel [45] as E. Hanseni from many stations in the E.
Atlantic between 37°S and 48°N, including Azores, Madeira, Canaries, and waters off the
coast of Morocco; and from inany stations in the W. Mediterranean between Gibraltar and
Sardinia. Reported by Tattersall [60] as E. unguiculata for waters off Cape Verde Islands. The
present results confirm previous records from the Atlantic.
Fam. Gnathophausiidae Udrescu, 1984
Gnathophausia gracilis Willemoes-Suhm, 1875
Gnathophausia gracilis W\\\QmoQS-S\x\m\ [67]: 33-37, PL IX (1-15); Illig [35]: 409-410, Figs.
15-17; Fage [19]: 27-34, Figs. 27-30; Vereshchaka [65]: 118; Wittmann & Wirtz [73];
Haroun & Garrido [30]; Wittmann et al. [71]; Price ct al [52].
Gnathophausia gracilis var. brevispinis Wood-Mason & Alcock [74]: 188.
Gnathophausia brevispinis - Wood-Mason & Alcock [75]: 269; Faxon [22]: 216-218, 257-258,
262, PL J.
Gnathophausia dentata Faxon [21]: 217.
Gnathophausia bidentata Illig [34]: 229-230, Fig. 2.
Material studied: Canary Islands.- 1 M ad., body length 69.8 mm, SW of El Hierro. off Punta de Or-chilla,
27.600N 18.267W, CANCAP-II, sta. 2.140, ring-trawl with weight, 2,000 m. bottom > 2,800 m,
8 Sept. 1977, 3.15-6.15 hrs.
68
Cape Verde Islands.- 2 imm. 76.8-79.3 mm. 1 juv.. N of Ilheu Grande. 15.133N 24.583W, CAN-CAP-
VII, sta. 7.035, midwater trawl, 0-2,000 m, 23/24 Aug. 1986.
N-Pacific (all samples by Norpax Equat. Himb.).- 2 F subad. 46-57 mm; 1 M ad. 68 mm, tow
from 4.0 17N 150.183W to 4.183N 150.216W. 1.100 m, sta. 77-12-16. 11 Dec. 1977.- 1 F subad. 50
mm, tow from 13.5N 150.0Wto 13.317N 150.000\V. 950 m. sta. 78-1-2. 7 Jan. 1978.- 2 F subad. 56-66
mm, 1 M subad. 48 mm, 2 JUV.. tow from 20.0 17N 150.200Wto 19.967N 1 50.383 W, 900 m. sta. 77-1 2-
34, 17 Dec. 1977.- 2 juv. 39-44 mm. tow tVom 21.33N 1 58.33W to 21.35N 1 58.50W. 1.100 m. sta. 78-
05-6, 2 May 1978.
Distribution.- Depth range: 900-5,000 m. Circumtropical, so far not recorded from boreal to arc-tic
waters. Reported by Fage [19] from (sub)tropical waters of all oceans, including waters off
the Canary and Cape Verde Islands. The present results tit well w ith this distributional scheme.
Gnathophaiisia zoea W illemoes-Suhm, 1873
Gnathophaiisia zoea WiUemoes-Suhm [66]: 401: [67]: 32-33, 37, PI. X (4): lUig [35]: 408-
409, Figs. 13-14: Nouvel [45]: 15-19, PI. 1(12), Tab. I; Tattersall & TattersalU63]: 82-88,
Figs. 3-^5.
Gnathophaiisia ^villemoesii Sars [56]: 38, PI. V (1-6); Faxon [22]: 215, PI. K (1).
Gnathophaiisia zoea var. sarsi Wood-Mason & Alcock [74]: 187.
Gnathophaiisia cristata lUig [34]: 319-321, Figs. A-B.
Gnathophaiisia zoea - Fage [19]: 34-39, Figs. 31-36: Lagardere & Nouvel [38]: 377-382,
Figs. 1-10: Lagardere [37]: 810; Vereshchaka [65]: 118; Escobar-Briones & Soto [17];
Diirr [15]; Wittmann & Wirtz [73]; Duir & Gonzalez [16]; Haroun & Garrido [30];
Wittmann et al. [71]: 1261; Price et al [52]; San Vicente [53].
Material studied: N.E. Atlantic off Portugal (all samples from FS "Poseidon" cruises).- 1 F ad., body
length 54.4 mm, 41.155N9.333\V. sta. Pos 113-9kul 1002. 800-900 m. 20 Nov. 1984.- 1 M ad. 42.9 mm.
40.120N9.838W, sta. Pos 113-11 ZD 1010. 1.160-1.340 m. 21 Nov. 1984.- 1 M ad. 44.9 mm, 39.21 7N
10.360W, sta. Pos 113-16 ZD 1017, 1,380-1,450 m, 22 Nov. 1984.- 2 imm. 24.9-35.8 mm. 39.217N
1 0.2 1 OW, sta. Po- 1 1 3 1 1 7 ZD, 740-920 m, 22 Nov. 1 984.
Canary Islands.- 2 spec. 31.7-35.2 mm, SW of Fuerteventura. Punta de Morrojable. 27.967N
14.200W, CANCAP-II, sta. 2.68, ring-trawl, 1,700 m, bottom 1,810 m, 29 Aug. 1977, 22.20-00.55 hrs.-
1 spec. ca. 47 mm, SW of El Hierro, off Punta de Orchilla, 27.65N 18.20W, CANCAP-II, sta. 2.123, ring-trawl
with weight. 1.500 m. bottom > 2.000 m. 2.25-4.50 hrs. 6 Sept. 1977.- 1 F subad. ca. 48.9 mm.
SW of El Hierro. off Punta de Orchilla. 27.667N 18.167W. CANCAP-II. sta. 2.130. 1.2 m Agassiz trawl.
1,500-1,800 m, 8 Sept. 1977.
Mauritania.- 3 F ad. 57.7-62.3 mm. 2 M ad. 52.5-56.7 mm. 1 imm., off Mauritania. 20.367N
17.900W. CANCAP-III, sta. 3. 119, 5 m beam trawl, 1,000 m, 28 Oct. 1978.- 1 M ad. 58.4 mm. 1 F subad.
57.5 mm, 3 imm.. off Banc d'Arguin. 19.717N 17.500\V. TYRO Mauritania-ll. sta. MAU.104. 3.5 m
Agassiz trawl in 1,500 m, 17 June 1988.- 2 F ad. 62.3-63.8 mm, 4 M ad. 53.2-64.9 mm, 4 subad., 5 imm..
off Mauritania, 19.150N 16.867W, TYRO Mauritania-II, sta. MAU.062, 2.4 m Agassiz trawl, 800-1,200
m. steep slope. 12 June 1988.- 3 F ad. 49.6-59.7 mm. 2 M ad. 58.4-60.0 mm, 1 M subad.. 4 imm.. 2 juv..
otTMauritania, 19.050N 16.967W. TYRO Mauritania-II. sta. MAU.042. 3.5 m Agassiz trawl in 820-990
m, 10 June 1988.- 1 Fad. 48.5 mm, 4 M ad. 51.1-64.0 mm, 6 F subad., off Mauritania, 18.850N 16.933W.
TYRO Mauritania-II. sta. MAU.041, 3.5 m Agassiz trawl in 800-840 m. 10 June 1988.
Cape Verde Islands.- 1 spec. ca. 47 mm in 2 parts. N of Ilheu Grande, 16.567N 24.683 W, CAN-CAP-
VIl. sta. 7.147. 1.2 m Agassiz trawl in 1,500-1.627 m. 4 Sept. 1986.- 1 F imm. ca. 51 mm. SE of
Sao Nicolau, off Preguica, 16.533N 24.233W, CANCAP-VII. sta. 7.137, 1.2 m Agassiz trawl in 715 m,
69
3 Sept. 1986.- 1 spec. 26.8 mm, S of Santiago, Pta Temerosa, 14.883N 23.533W, CANCAP-VII, sta.
7.023, 1 .2 m Agassiz trawl in 525 m, 22 Aug. 1 986.
N. Pacific (all samples by Norpax Equat. Himb.).- 1 M ad. 103 mm, 2 M imm., 2 juv., tow from
21.33N 158.33Wto21.35N 158.50W, 1,200 m, sta. 78-05-5, 2 May 1978.- 4 juv. 62-73 mm, tow from
21.33N 158.33Wto21.35N 158.50W, 1,100 m, sta. 78-05-6, 2 May 1978.- 1 Fad. 113 mm, 1 F imm.,
towfrom21.33N 158.33W to 21.50N 158.50W, 1,100 m, sta. 77-11-5, 1 Nov. 1977.
Distribution.- Depth range: 400-6,000 m. Cosmopolitan, mainly (sub)tropical, also in boreal
to subarctic waters, but not in the Arctic Ocean. Recorded by Fage [19] and Nouvel [45] from
the Azores. Cited by Nouvel [45] for Atlantic waters between Greenland and the Cape of
Good Hope. First record for the Canaries by Wittmann et al. [71]. Cited by San Vicente [53]
for the Mediterranean. The present results fit well with the previously known distribution.
Neognathophausia gigas Willemoes-Suhm, 1873
Gnathophausia gigas Willemoes-Suhm [66]: 400; [67]: 28-31, 37, Pis. IX (16-17), X (2-3);
Fage [19]: 24-27, Fig. 26; Nouvel [45]: 12-15, PI. I (5-11); Tattersall & Tattersall [63]:
77-82, Figs. 1-2; Lagardere [37]: 810.
Gnathophausia drepanephora Hoh & Tattersall [32]: 113, 142, Fig. I, PI. XVIII; Ortmann
[48]: 38; Illig 1930:408.
Neognathophausia gigas - Petryashev [49]: 47-48, PI. 1 (1-5); [50]: 959, 968-969, Fig. 2;
[51]: Tab. 2; Wittmann & Wirtz [73]; Haroun & Garrido [30]; Wittmann et al. [71]: 1261,
Tab. I; Fukuoka [23]: 405-406.
Material studied: N. Pacific- 1 F subad., body length 107 mm, 1 M subad. 99 mm, 4 F imm., 3 juv.,
Bering Sea, 53.227N 163.838W, Dominator Cruise 021, sta. 75, midwater trawl, 681 m, 1 Aug. 1982.-
1 juv. 52 mm, tow from 21.33N 1 58.33W to 21.35N 158.50W, 1,100 m, Norpax Equat. Himb., sta. 78-
05-6, 2 May 1978.
Distribution.- Depth range: 300-4,000 m. Cosmopolitan, but not found in the Arctic Ocean.
Recorded by Fage [19] for temperate to tropical waters of the Pacific and the N. Atlantic, in-cluding
the Cape Verde Islands; by Nouvel [45] from the Azores, Brittany, and Gibraltar; by
Haroun & Garrido [30] for the Canaries. The E. Atlantic distribution ranges from Ireland to
Dakar according to Nouvel [45].
Neognathophausia ingens (Dohrn, 1870)
Lophogaster ingens Dohrn [14]: 610, PL 31 (12-14).
Gnathophausia ingens - Sars [56]: 30, PI. II; Illig [35]: 407-408, Figs. 11-12; Fage [19]; 15-
24, Figs. 20, 24-25; Nouvel [45]: 9-12, PI. I (4); Tattersall [60]: 31-35; Casanova [7]: 328-
330; Escobar-Briones & Soto [17]: Tab. 1.
Gnathophausia calcarata Sars [56]: 35, PI. IV; Ortmann [48]: 30, PI. I (2a-f).
Gnathophausia bengalensis Wood-Mason & Alcock [75]: 269.
Gnathophausia doiyphora Illig [34]: 227, Fig. 1 A-D.
Neognathophausia ingens - Petryashev [49]: 47-48; [50]: 959, 968-969, Fig. 3; [51]: Tab. 2;
Diirr [15]; Wittmann & Wirtz [73]; Diirr & Gonzalez [16]; Haroun & Garrido [30]:
Wittmann e? fl/. [71]: 1261, Tab. I.
70
Material studied: Madeira.- 1 ju\., body length 36.8 mm, SE of Madeira, 32.583N 16.733\V. CANC.'VP-I,
sta. 1.035, ring-trawl, wire 1,250 m, bottom about 2,000 m, 10 Mar. 1976.
Canary Islands.- Samples listed in Wittmann et al. [71].
Cape Verde Islands.- 1 spec. 90.3 mm. S\V of Sal, 16.700N 23.683W, CANCAP-VII, sta. 7.1 14.
0-330 m, midwater trawl, 31 Aug. 1986.- 1 spec. 49.4 mm, S of Branco, 16.517N 24.8 17\V. CANCAP-VII.
sta. 7.148. 0-1.100 m. midwater trawl. 4-5 Sept. 1986.
Tropical Pacific (all samples by Norpa.x Equat. Himb.).- 2 M ad. 11 2- 122 mm. 2 F imm.. 4 juv.,
tow from 21.33N 1 58.33W to 21.50N 158.50W, 1.200 m, sta. 78-05-5, 2 May 1978.- 1 F subad.. tow
from21.33N 158.33Wto2l.50N 158.50W, 310 m, sta. 77-1 1-2, 31 Oct. 1977.- 1 F imm. 84 mm, Ijuv.,
towfrom2l.33N 158.33Wto 21.50N 158.50W, 1,100 m, sta. 77-11-5, I Nov. 1977.- 3 juv. 36-40 mm,
tow from 15.083N 1 50.1 83W to 15.217N 1 50.1 83 W. 295 m. sta. 77-12-30. 15 Dec. 1977.- I F subad.
88 mm. tow from 14.983N 153.000W to 15.083N 153.000W, 400 m, sta. 79-02-8. 19 Feb. 1978.- 3 juv.
38-54 mm. tow from 14.733N 157.983Wto 14.617N 1 57.983W, 330 m, sta. 79-5-3. 12 May 1979.- 1 F
subad. 181 mm. tow from 9.850N 150.000W to 9.833N 150.017W, 1,500 m. sta. 79-2-1. 8 Feb. 1979.-
2juv. 58-72 mm. 0.867N 153.033W to 0.967N 153.033W. 360 m, sta. 79-5-12, 23 May 1979.- 2 juv. 58-
63 mm, tow from 1.367S 150.167\Vto 1.483S 1 50. 167\V. 290 m, sta. 77-12-10, 9 Dec. 1977.- 1 juv. 70
mm,towfrom3.067S 157.950Wto3.067S 1 57.833 W, 320 m. sta. 78-5-11. 20 May 1978.- 1 juv. 70 mm.
tow from 6.967S 158.1 I7W to 6.833S 1 58.1 33W, 335 m, sta. 77-12-6. 6 Dec. 1977.
Distribution.- Depth range: 200-4,000 m. Circumtropical, ranging from temperate to tropi-cal
latitudes. Recorded by Page [19] from the Pacific, Indian, and Atlantic Oceans, including
waters off Azores. Madeira, Canary; and Cape Verde archipelagos, and off the continental
coasts of Morocco and West Sahara. Recorded by Nou\ el [45] from the Azores, Canaries,
and Gibraltar. Details for the Canaries in Wittmann et al. [71]. The present results fit well
with the already know n distribution.
Fam. Lophogastridae G.O. Sars. 1870
Chalaraspidiim alatiim (\\ illemoes-Suhm, 1876)
Chalaraspis alata Willemoes-Suhm [68]: 592: Page [18]: 68-75. Figs. I-\TI: [19]: 4.
Figs. 1-10.
Chalaraspidiim alarum - Willemoes-Suhm [69]: 521; Tattersall [62]: 14-15: Tattersall [60]: 28-
30, Fig. 1; Springer & Bullis [59]; Haroun & Garrido [30]; Price et al. [52].
Eclytaspis alata - Faxon [22]: 219.
Distribution.- Depth range: 1.000-3.000 m. Cosmopolitan, but so far not recorded from the
Arctic and Antarctic Oceans. It was listed by Haroun & Garrido [30] for the fauna of the Ca-naries,
by Springer & Bullis [59] and Price et al. [52] for the Gulf of Mexico. A damaged
specimen of this genus was recorded without detemiination at species le\el by Zimmer [76]
from the Azores.
Lophogaster spinosits Ortmann, 1907
Lophogaster spinosus Ortmann [48]: 26-27. PI. I (la. b); Page [20]: 23-29. Pigs. 13-15, 16f
17g; Casanova [7]: 352. Pig. 82: Diirr [15]: Diirr & Gonzalez [16]: Wittmann et al. [71]:
1261-1262. Tab. I.
71
Material studied: Canary Islands.- Samples listed in Wittmann et al. [71].
NW-Atlantic- 1 F subad., body length 26 mm, 2 juv., tow from 24.772N 70.4 lOW to 24.745N
70.440W, Sargasso Eel & Maine, Cruise 83, sta. 1014, 280 m, 17 Feb. 1983.- 1 F subad. 24 mm, 2 juv.,
tow from 24.747N 70.385W to 24.7 18N 70.405W, Sargasso Eel & Maine, Cruise 83, sta. 1055; ALQ
1/16, 304 m, 24 Feb. 1983.
Distribution.- Depth range: 200-5,000 m. According to Fage [20] restricted to tropical and
subtropical waters of the W.- and E. Atlantic. The present results fit well with this distribu-tional
scheme.
Lophogaster typicus M. Sars, 1857
Lophogaster typicus Sars [57]: 160; [58]: 1-37, Tabs. I-III; Norman [42]: 459; [43]: 10; Colosi
[10]: 6-7, Figs. 7, 7a; [11]: 2; lUig [35]: 405; Fage [20]: 7-13, figs. 2-4, 16b, 17b; Nouvel
[45]: 7-8, PL I (1-3); Tattersall & Tattersall [62]: 90-97, Figs. 6-8; Tattersall [61]: 145;
Hoenigman [31]: 605-606, Fig. 2; Lagardere & Nouvel [38]: 382-383; Katagan [36]: 288;
Wittmann & Stagl [72]: 160; Diirr [15]; Diirr & Gonzalez [16]; Nikoforos [40]; Wittmann
et al [71]: 1262, Tab. II; San Vicente [53].
Ctenomysis alata Norman [41]: 151.
Lophogaster serratus Bjorck [4]: 6-8, Figs. 1-5.
Material studied: Boreal to temperate N.E. Atlantic.- 3 F ad., body length 18.0-20.5 mm, 3 M ad. 17.7-
20.0 mm, Norway, HjelteQord, 61.583N 4.9 17E, 260 m, 4 July 1978, leg. Torleiv Brattegard.- 1 F ad.
24.4 mm, 1 Fad. with pleon missing, Norway, W of Stavanger 59.11 2N 3.083E, 153-156 m, 7 Aug. 1984,
Senckenberg Museum, Frankfurt am Main.- 1 F ad. 18.7 mm, 4 M ad. 18.2-23.5 mm, 1 F subad., off Por-tugal,
41.9217N 9.3167W, FS 'Toseidon", sta. Po-113-7 ZD2 999, 6 F ad. 25.5-32.1 mm, 4 M ad.
25.5-27.1 mm, 3 juv., off N-Portugal, 41.917N 9.333 W, FS "Poseidon", sta. Pos 113-7 ZDl
998, 175-184 m, 18 Nov. 1984.- For sample off Ireland see Wittmann & Stagl [72].
Azores.- 1 F ad. 21.8 mm, S of Sao Miguel, 37.650N 25.517W, CANCAP-V, sta. 5.012, van Veen
grab in 480 m, 26 May 1981.
Mediterranean.- 1 F ad. 17.5 mm, Adriatic Sea, off Zirje, 43.5847N 15.6333E, Ockelmann
epibenthic sledge in 199 m, mud, 12 Sept. 1971, leg. Peter Cate.- 1 F ad. 15.8 mm, Adriatic Sea, E of
Ancona, 42.4633N 17.1667E, Ockelmann epibenthic sledge in 400 m, 4 July 1973, leg. Peter Cate.- 1
Fad. 20.7 mm, Sardinia, Gulf of Cagliari, 539-560 m, 20 July 1984, 07:48-09:25 hrs.- 1 Fad.21.5mm,
1 Mimm., Sardinia, Gulf of Cagliari, 329-402 m, 20 July 1984, 16:00-1 7:30 hrs.- For sample off Messina
see Wittmann & Stagl [72].
Canary Islands.- 1 M 23.8 mm, S of Fuerteventura, Punta Jandia, 28.050N 14.483W, CANCAP-II,
sta. 2.03, van Veen grab in 140-200 m, 23 Aug. 1977, 22.20-00.55 hrs.
Mauritania.- 3 M ad. 19.0-23.4 mm, off Mauritania, 18.833N 16.467W, TYRO Mauritania-II,
sta. MAU.022, 1.2 m Agassiz trawl in 60-66 m, 8 June 1988.
Distribution.- Mainly bentho-pelagic in 30-500 m. Fage [20] provided a number of records
from the Mediterranean and concluded that this species may be restricted to the N.E. Atlantic
and Mediterranean. According to Nouvel [45], it occurs in the E. Atlantic from Norway to
Mauritania, throughout the Mediterranean Sea, and as far east as the Bosporus. In line with this,
Tattersall & Tattersall [63] and Tattersall [61] argued that previous records from the Pacific may
be attributed to other species and concluded that L. typicus may be restricted to the (N&S) E.
Atlantic and Mediterranean. The present results fit well with this distributional scheme.
72
DISCUSSION
Taxonomic notes
The following discussions may help to reduce certain confusions and misunderstand-ings
in the literature about lophogastrids:
Status of £. hanseni Nouvel, 1942, versus Eucopia ungiiiciilata Willemoes-Suhm,
1875.- Willemoes-Suhm [67] described E. ungiiiciilata based on material of the "Challenger"
expedition. Hansen [26] re-examined this material and concluded that it actually belongs to
three different species, E. ungiiiciilata, E. australis, and E. sculpticaiida. Nouvel [44] noted
much confusion about the identity of E. ungiiiciilata and proposed to suppress this taxon in
favour of two new taxa, established by him in the same publication as E. hanseni and E.
ghmaldii. According to the nomenclatorial code of that time, still valid today (ICZN [33]), he
ought not to have suppressed but to have revised the taxon ungiiiciilata and to have described
only one new species (probably E. grimaldii based on longer spines on the telson). Therefore
we agree with Tattersall [60], who considered E. hanseni di]vin\ov synonym of £". ungiiiculata.
Status of the genus Neognathophausia Petryashev, 1992, versus Gnathophausia Wille-moes-
Suhm, 1873.- Page [19] established three morphologically distinct groups within the
genus Gnathophausia: group I with G. ingens and G. gigas, group II w ith G. gracilis only, and
group III with all remaining species of the genus. Later, Tattersall [60] essentially confirmed
this grouping, but she argued that it would not be practical for taxonomic purposes. Based on
the structure of antennal scale, maxillipeds, and abdominal pleura, Petryashev [49] defined a
new genus, named Neognathophausia. to include all members of group I. Casanova et al. [8],
however, rejected the definition of the new genus. Their conclusion was based on rRNA gene
sequences and moiphological data, in their opinion pointing to a basal position of G. gracilis
with respect to the t\\ o remaining groups. Howe\ er, they examined only one species per group,
which appears quantitati\ely insufficient to estimate genetic distance between multi-species
groups. Unless further evidence is presented, Neognathophausia may be maintained, as al-ready
practiced (without detailed reasoning) by Wittmann et al. [71], Petryashev [50, 51],
Fukuoka [23], and Anderson [2].
Doubtful Mediterranean records of Lophogaster afflnis Colosi, 1930.- This species is
well known from the Red Sea and Indian Ocean (Colosi [12], Coifmann [9], Nouvel [46],
Almeida Prado-Por [1]). Previous reports for the Mediterranean were found only in second-ary
literature (Miiller [39], Nikoforos [40], San Vicente [53]). These later reports may be based
merely on erroneous interpretation of a puzzling fonnulation by Page [20] on p. 20: "Mais il
existe, de I 'autre cote de la Mediterranee, dans la moitie septentrionale de la Mer Rouge, line
forme decrite par Colosi (1929) sous le nome de L. affinis ...
" to our translation "But on the
other side of the Mediterranean, in the northern half of the Red Sea, there is a form described
by Colosi (1929) under the name L. affinis ...". From this and additional data, we conclude
that this species has so far never been found in the Mediterranean.
Diversity and distribution of the Canarian lophogastrids
Most species show a widespread geographical distribution, typical of oceanic, meso-to
bathypelagic organisms. Tab. 1 compiles the so far known distribution of the di\'erse species
in mostly subtropical waters of the N.E. Atlantic, for comparison also including the Gulf of
Mexico. Ten out of the twelve species from the Canary Islands have a world-wide distribu-tion.
The remaining tvvo species are endemic for the Atlantic Ocean, i.e. Lophogaster spinosiis
73
on both sides of the Atlantic, and L. typicus only in the E. Atlantic. A similar pattern is found
in the Gulf of Mexico: nine out of the eleven species are cosmopolitan, only L. americamis
and L. longirostris are endemic for the western Atlantic (species list compiled from Springer
& Bullis [59], Escobar-Briones & Soto [17], and Price et al. [52]).
The Canaries apparently represent the most species-rich area in the Atlantic due to high
sampling intensity in combination with favourable natural conditions: comparably high plank-ton
and micronekton densities representing good feeding grounds influenced by the Saharan
upwelling along the African Atlantic coast, < 90 km from the Canary archipelago (Barton et
al. [3]), and strong depth gradients allowing deep-water organisms to approach closely to the
islands. Compared with the four N.E. Atlantic archipelagos considered in table 1 , fewer species
are reported along the continental coasts of Morocco and Mauritania compared to the Canary
Islands. The same pattern is found in the oligotrophic Mediterranean basin, characterized by
a 'shallow' entrance through the Strait of Gibraltar and by warmer, more saline bottom water.
ACKNOWLEDGEMENTS
We are indebted to the Viceconsejeria de Medioambiente del Gobiemo de Canarias
for coordinating the BIOTA MARINO project and for supporting the bibliographic compila-tion
of this crustacean group.
REFERENCES
[1] ALMEIDA PRADO-POR, M.S. 1980. Mysidacea from the Gulf of Elat (Gulf of 'Aqaba).
Israel Journal ofZoology^, 29: 1 88- 1 9 1
[2] ANDERSON, G. 2010. Lophogastrida Classification, January 20, 2010. http://per-acarida.
usm.edu/LophogastridaTaxa.pdf. 1-2. (Instant Web Publishing, USA)
[21.1.2010].
[3] BARTON, E.D., J. ARISTEGUI, R TETT, M. CANTON, J. GARCIA-BRAUN, S.
HERNANDEZ-LEON, L. NYKJAER, C. ALMEIDA, J, ALMUNIA, S. BALLES-TEROS,
G. BASTERRETXEA, J. ESCANEZ, L. GARCIA-WEILL, A. HERNANDEZ-GUERRA,
F. LOPEZ-LAATZEN, R. MOLINA, M.F. MONTERO, E.
NAVARRO-PEREZ, J.M. RODRIGUEZ, K. VAN LENNING, H. VELEZ & K. WILD.
1998. The transition zone of the Canary Current upwelling region. Progress in Oceanog-raphy,
41:455-504.
[4] BJORCK, W. 1916. Svenska krafdjur i Goteborgs Museum 1 . Schizopoda. Meddelanden
frdn Goteborgs Musei zoolgiska Afdelning, 7: 1-17.
[5] CALMAN, W.T. 1 896. On the deep-sea Crustacea from the southwest of Ireland. Trans-actions
ofthe Royal Irish Academy, 3 1 : 1-20, pis. I-II.
[6] CARTES, J.E. & J.C. SORBE. 1995. Deep-water mysids of the Catalan Sea: species
composition, bathymetric and near-bottom distribution. Journal of the Marine Biologi-cal
Association ofthe United Kingdom, 75(1): 187-197.
[7] CASANOVA J. -P. 1977. Lafaune pelagique profonde (zooplancton et micronecton) de
la province atlanto-mediterraneenne. Aspects taxonomique, biologique et zoogeo-graphique.
These, Universite de Provence (Aix - Marseille I): i-ix, 1-500.
74
[8] CASANOVA, J. P., L. DE JONG & E. FAURE. 1998. Interrelationships ot the two fam-ilies
constituting the Lophogastrida (Crustacea: Mysidacea) inferred from morphologi-cal
and molecular data. Marine Biolog\\ 132( 1 ): 59-65.
[9] COIFMANN, I. 1 937. 1 misidacei del Mar Rosso. Studio del materiale raccolto dal Prof.
L. Sanzo durante la campagna idrografica della R. Na\e Ammiraglio Magnaghi ( 1923-
1924). Memohe del R. Comitate Talassografico Italiano, 233: 1-52, Pis. I-XXV.
[10] COLOSI, G. 1922. Eufausiacei e Misidacei raccolti nella campagna del 1920. Memorie
del R. Comitato Talassografico Italiano, 96: 1-12, Tab. 1.
[11] COLOSI, G. 1929. I Misidacei del Golfo di Napoli. Puhhlicazioni della Stazione Zoo-logica
di Napoli, 9(3): 405-439.
[12] COLOSI, G. 1930. Lofogastridi nuovi. Bolletino di Zoologia, 1(4): 1 19-125.
[13] DANA, J.D. 1852. Crustacea. Part I. In: United States Exploring Expedition during the
years 1 838-1842 under the command of Charles Wilkes, U.S.N. , 13: i-viii. 1-685. (C.
Shemian, Philadelphia).
[14] DOHRN, A. 1870. Untersuchungen iiber Bau und Entwicklung der Arthropoden. 10.
Beitrage zur Kenntnis der Malacostraken und ihrer Larven. Zeitschrift fiir %vis-senschaftliche
Zoologie, 20: 607-625, 3 pis.
[15] DURR, J. 1997. Nahrungsokologie von Beryx splendens und Benx decadactyius fOste-ichthyes,
Betycidae) im Bereich der Kanarischen Inseln. In: Diplomarbeit an der Mathe-matisch-
Naturwissenschaftlichen Fakultat: 1-85. (Christian-Albrechts-Universitat, Kiel).
[16] DURR. J. & J. A. GONZALEZ. 2002. Feeding habits of Beiyx splendens and ^m.v
decadactylus (Berycidae) off the Canary Islands. Fisheries Research, 54: 363-374.
[17] ESCOBAR-PRIONES. E. & L.A. SOTO. 1991. Biogeografia de los Misidaceos (Crus-tacea:
Peracarida) del Golfo de Mexico. Caribbean Journal ofScience, 27(1-2): 80-89.
[18] FAGE, L. 1939. Apropos d'un mysidace bathypelagique peu connu: Chalaraspis alata
G.O. Sars (Willemoes-Suhm in lit.). Archives de Zoologie experimentale et generale,
80: 68-76.
[19] FAGE, L. 1941. Mysidacea Lophogastrida - I. The Carlsberg Foundation's oceano-graphical
expedition round the world 1928-1930 and previous Dana-expeditions under
the leadership of Prof. Johannes Schmidt. Dana Reports, 4(19): 1-52.
[20] FAGE, L. 1942. Mysidacea Lophogastrida - II. The Carlsberg Foundation's oceano-graphical
expedition round the world 1928-1930 and previous Dana-expeditions under
the leadership of Prof. Johannes Schmidt. Dana Reports, 4(23): 1-67.
[21] FAXON, W. 1893. VI. Preliminary descriptions of new species of Crustacea. In: Reports
on the dredging operations off the west coast of Central America to the Galapagos, to the
west coast of Mexico, and the Gulf of California; in charge of Alexander Agassiz, car-ried
on by the U.S. Fish Commission Steamer Albatross during 1 89 1 . Bulletin ofthe Mu-seum
of Comparative Zoology- at Hanard College, 24 (7): 149-220.
[22] FAXON, W. 1895. XV. The stalk-eyed Crustacea. In: Reports on an exploration off
the west coasts of Mexico, Central and South America, and off the Galapagos Islands,
in charge of Alexander Agassiz, by the U.S. Fish Commission Steamer Albatross
during 1891. Memoirs of the Museum of Comparative Zoology at Harvard College,
18: 1-292, 67 pis.
[23] FUKUOKA. K. 2009. Deep-sea mysidaceans (Crustacea: Lophogastrida and Mysida)
from the northwestern North Pacific off Japan, with descriptions of six new species. In:
75
Fujita, T. (ed.), Deep-sea fauna and pollutants off Pacific coast of northern Japan. A^a-tional
Museum ofNature and Science Monographs, 39: 405-446.
[24] FURNESTIN, M.-L. 1960. Zooplancton du Golfe du Lion et de la cote orientale de
Corse. Revue des Travaux de I'lnstitut des Peches Maritimes, 24 (2): 153-252.
[25] HANSEN, H.J. 1905. Preliminary report on the Schizopoda collected by H.S.H. Prince
Albert of Monaco during the cruise of the Princesse-Alice in the year 1901. Bulletin du
Musee Oceanographique de Monaco, 30: 1-32.
[26] HANSEN, H.J. 1910. The Schizopoda of the Siboga Expedition. Siboga Expeditie,
Monographic, 37: 1-123, pis. 1-16. (Leyden).
[27] HANSEN, H.J. 1913. Report on the Crustacea Schizopoda collected by the Swedish
Antarctic Expedition 1901-1903, under the charge of Baron Dr. Otto Nordenskjold: 1-56.
(G.E.C. Gad, Copenhagen).
[28] HANSEN, H.J. 1927. Sergestides et Schizopodes. In: Expedition scientifique du "Tra-vailleur"
et du "Talisman" 1880-3: 1-26, pi. I. (G. Mason, Paris).
[29] HARGREAVES, RM. 1999. The vertical distribution of micronektonic decapod and
mysid crustaceans across the Goban Spur of the Porcupine Seabight. Sarsia, 84: 1-18.
[30] HAROUN, R. & M.J. GARRIDO. 2003. Orden Mysidacea. R 68, 69 in: L. Moro, J.L.
Martin, M.J. Garrido & I. Izquierdo (eds.), Lista de especies marinas de Canarias (algas,
hongos, plantas y animales) 2003: 248 pp. (Consejeria de Politica Territorial y Medio
Ambient, Tenerife).
[31] HOENIGMAN, J. 1963. Mysidacea de Texpedition Hvar (1948-49) dans I'Adriatique.
Rapportes de la Commission internationale pour I'Exploration scientifique de la Mer
Mediterranee, 17: 603-616.
[32] HOLT, E.W.L. & W.M. TATTERSALL. 1905. Schizopodous Crustacea from the north-east
Atlantic slope. Scientific Investigations. Fisheries Branch, Department ofAgricul-ture
for Ireland, Dublin - Annual Report, 1902-1903, pt. II, app. IV: 99-152, pis.
xv-xxv.
[33] ICZN (International Commission on Zoological Nomenclature), 1999. International Code
of Zoological Nomenclature, 4^'' edition: 1-306. (International Trust for Zoological
Nomenclature, London). Also available at http://www.iczn.org/iczn/index.jsp
[34] ILLIG, G. 1906. 4. Ein weiterer Bericht iiber die Schizopoden der Deutschen Tiefsee
Expedition 1898-1899. II. Gnathophausien (Fortsetzung). Zoologischer Anzeiger, 30(10):
319-322.
[35] ILLIG, G. 1930. Die Schizopoden der Deutschen Tiefsee-Expedition. In: C. Chun (ed.),
Wissenschaftliche Ergebnisse der Deutschen Tiefsee-Expedition auf dem Dampfer „Val-divia"
1898-1899, 22(6): 397-620.
[36] KATAGAN, T. 1985. Mysidaces et cumaces des cotes egeennes de Turquie. Rapportes
et proces-verbaux des Reunions de la Commission internationale pour I Exploration sci-entifique
de la Mer Mediterranee, 29(5): 287-288.
[37] LAGARDERE, J.P. 1983. Les mysidaces de la plaine abyssale du Golfe de Gascogne 1.
Families des Lophogastridae, Eucopiidae et Petalophthalmidae. Bulletin du Museum na-tional
d'Histoire naturelle, 4^""' serie, section A (Zoologie, Biologic, Ecologie animale),
5(3): 809-843.
[38] LAGARDERE, J.R & H. NOUVEL. 1980. Les Mysidaces du talus continental du golfe
de Gascogne. II. Families des Lophogastridae, Eucopiidae et Mysidae (tribu des Ery-
76
thropini exceptee). Bulletin dii Museum national d'Histoire naturelle. 4''""'' serie, section
A (Zoologie, Biologie, Ecologie animate), 2(2): 375-412.
[39] MULLER, H.-G. 1993. World catalogue and bibliography of the recent Mysidacea. 1-
491. (Wissenschaftlicher Verlag H.-G. Muller, Wetzlar).
[40] NIKOFOROS, G. 2002. Fauna del Mcditcrraneo. 1-366. (Giunti, Firenze).
[41] NORMAN, A.M. 1862. On the Crustacea, Echinodermata, and zoophytes obtained in
deep-sea dredging off the Shetland Isles in 1861. Reports of the British Association for
the Advancement ofScience, 3 1 ( 1 86 1 ): 151-152.
[42] NORMAN, A.M. 1892. British Schizopoda of the families Lophogastridae and Eu-phausiidae.
Annals and Magazine ofnatural History?, Sen VI, vol. 9: 454-464.
[43] NORMAN, A.M. 1905. Museum Normanianum, or a catalogue of the Invertebrata of
the Arctic and north Atlantic temperate ocean and Palaearctic region. III. Crustacea. 2"*^
ed.: i-vi, 1-47. (Thos. Caldcleugh & Son. Printers. Durham).
[44] NOUVEL, H. 1942. Sur la systematique des especes du genre Eucopia Dana 1852 (Crust.
Mysidacea). Bulletin de I'lnstitut Oceanographique, 818: 1-8. (Monaco).
[45] NOUVEL, H. 1943. Mysidaces provenant des campagnes du Prince-Albert ler de
Monaco. In : J. Richard (ed.), Resultats des campagnes scientifiques accomplies sur son
yacht par Albert ler Fasc. 105. 1-128, pis. I-V. (Imprimerie de Monaco).
[46] NOUVEL, H. 1978. Mysidaces recoltes par S. Frontier a Nosy Be. VII. Lophogaster
affinis Colosi et Mesopodopsis africana madagascariensis n. ssp. Bulletin de la Societe
d'Histoire Naturelle de Toulouse, 113(3-4): 396-403.
[47] ORTMANN, A.E. 1906. Schizopods of the Hawaiian Islands collected by the steamer Al-batross
in 1902. Bulletin ofthe United States Fish Commission, 23. pt. 3(1903): 961-973.
[48] ORTMANN, A.E. 1907. Schizopod crustaceans in the U.S. National Museum. The fam-ilies
Lophogastridae and Eucopiidae. Proceedings of the United States National Mu-seum,
31: 23-54, pis. 1, 11.
[49] PETRYASHEV, VV 1992. Notes on mysid systematics (Crustacea, Mysidacea) of Arc-tic
and the North-Westem Pacific. Zoologicheskij Zhurnal, 71(10): 47-58.
[50] PETRYASHEV. VV. 2005. Mysids (Crustacea, Mysidacea) collected by Soviet and Russ-ian
Antarctic expeditions. Lophogastrida, Petalophthalmida, and Mysida: Boreomysi-dae.
Zoologicheskij Zhurnal, 84(8): 957-973.
[51] PETRYASHEV, VV 2007. Biogeographical division of Antarctic and Subantarctic by
Mysid (Crustacea: Mysidacea) fauna. Russian Journal ofmarine Biology, 33(1): 1-16.
[52] PRICE, W.W., R.W. HEARD, R AAS & K. MELAND. 2009. Lophogastrida (Crustacea)
ofthe GulfofMexico, 923-927 in D.L. Felder & D.K. Camp (eds.). Gulf of Mexico-Ori-gins,
Waters, and Biota. Biodiversity. (Texas A&M Press. College Station. Texas).
[53] SAN VICENTE, C. 2010. Mysidaceans. In: M. Coll. C. Piroddi. .1. Steenbeek, K.
Kaschner. F. Ben Rais Lasram. J. Aguzzi. E. Ballesteros. C.N. Bianchi et al. (eds.). Bio-diversity
of the Mediterranean Sea: estimates, patterns & threats. PloS ONE, 5(8):
el 1842: 254-275.
[54] SARS. G.O. 1870. Carcinologiske Bidrag til Norges Fauna. I. Monographi over de ved
Norges Kyster forekommende Mysider. Pt. 1. 1-64, 5 pis. (K. Norske Viidenskab. Trond-hjem,
Christiania).
[55] SARS. G.O. 1884. Preliminary notice on the Schizopoda of H. M.S. Challenger expedi-tion.
Forhandlinger i Videnskahs-Selskahet, 7(1883): 1-43. (Christiania).
77
[56] SARS, G.O. 1 885. Report on the Schizopoda collected by H.M.S. Challenger during the
years 1873-1876. In: G.S. Nartes (ed.), Report on the scientific results ofthe Voyage of
H.M.S. Challenger during the years 1873-76, 13(37): 1-228, 38 pis. (Longmans & Co.,
London).
[57] SARS, M. 1 857. Om 3 nye norske Krebsdyr. Forhandlinger ved det Skandinavske Naturf
Mode i Christiania, 7: 160-175.
[58] SARS, M. 1862. Beskrivelse over Lophogaster typicus, en maerkvaerdig form af de la-vere
tifoddede krebsdyr. In: Universitetsprogram for andet Halvaar 1862, i-iv, 1-37,
Tabs. I-III. (Det Kongl. Norske Universitet, Christiania).
[59] SPRINGER, S. & H.R. BULLIS, Jr. 1956. Collections by the Oregon in the Gulf of Mex-ico.
List of crustaceans, molluscs and fishes identified from the collections made by the
exploratory fishing vessel Oregon in the Gulf of Mexico and adjacent seas 1950 through
1955. Special Scientific Report: Fisheries, 196: 1-134. (United States Fish and Wildlife
Service, Washington, D. C).
[60] TATTERSALL, O.S. \955.MysidacQSL. Discoveiy Reports, 2S: 1-190.
[61] TATTERSALL, O.S. 1961. Mysidacea from the coasts of tropical West Africa. Atlantide
Report, 6: 143-159.
[62] TATTERSALL, W.M. 1951. A review of the Mysidacea of the United States National
Museum. Bulletin ofthe United States National Museum, 201: 1-292.
[63] TATTERSALL, W.M. & O.S. TATTERSALL. 1951. The British Mysidacea. Ray Soci-ety,
Monograph, no. 136: 1-460. (The Ray Society, London).
[64] UDRESCU, A. 1984. Transspecific-evolution (family level) within Lophogastrida. A
new family - Gnathophausiidae (Lophogastrida, Mysidacea). Travaux du Museum na-tional
d'Histoire naturelle «Grigore Antipa», 25: 59-11
[65] VERESHCHAKA, A.L. 1990. Mysids from seamounts of Nasca and Sala-y-Gomez
Ridges. Trudy Instituta Okeanologii Akademija Nauk SSSR, 124: 118-128.
[66] WILLEMOES-SUHM, R. von. 1873. Notes from the Challenger, VII. Nature, 8: 400-403.
[67] WILLEMOES-SUHM R. von. 1875. II. On some Atlantic Crustacea from the Challenger
expedition. Transactions oftheLinnean Society ofLondon, Zoology^, Ser. II, vol. 1(1): 23-
59, Pis. VI-XIII.
[68] WILLEMOES-SUHM R. von. 1876. Preliminary report to Prof Wyville Thomson, on
Crustacea observed during the cruise of H.M.S. Challenger in the southern seas. Pro-ceedings
ofthe Royal Society ofLondon, 24: 585-592.
[69] WILLEMOES-SUHM, R. von. 1895. In: J. Murray (ed.), A summary of the scientific re-sults
obtained at the sounding, dredging, and trawling stations of H.M.S. Challenger. Re-port
on the scientific results of the voyage of H.M.S. Challenger during the years
1872-76, 1: i-vii, 1-796. (Eyre and Spottiswoode, London).
[70] WITTMANN, K.J. 1990. Mysidacea. In: J. Sieg & J.W. Wagele (eds.). Fauna der An-tarktis.
Paul Parey, Berlin: 130-133.
[71] WITTMANN, K.J., F. HERNANEZ, J. DURR, E. TEJERA, J.A. GONZALEZ & S.
JIMENEZ. 2004. The epi- to bathypelagic Mysidacea (Peracarida) off the Selvagens,
Canary, and Cape Verde Islands (NE Atlantic), with first description of the male of Lon-githoraxaliceiU.
Nouvel, 1942. Crustaceana, 76(10): 1257-1280.
[72] WITTMANN, K.J. & V. STAGE. 1996. Die Mysidaceen-Sammlung am Naturhis-torischen
Museum in Wien: eine kritische Sichtung im Spiegel der Sammlungs-geschichte.
Annalen des Naturhistorischen Museums in Wien, 98B: 157-191.
78
[73] WITTMANN, K.J. & P. WIRTZ. 1998. A first inventory of the mysid fauna (Crustacea:
Mysidacea) in coastal waters of the Madeira and Canar\' archipelagos. Boletim do Miiseii
Municipal do Funchal, (Sup.) 5: 5 11 -533.
[74] WOOD-MASON, J. & A. ALCOCK. 1891a. Natural history- notes from H.M. Indian
Marine Steamer In\estigator. Commander R.F. Hoskyn. R.N.. commanding. - No. 12.
Note on the results of the last season's deep-sea dredging. Annals and Magazine of nat-ural
Histoiy, sen 6, vol. 7: 186-202.
[75] WOOD-MASON, J. & A. ALCOCK. 1891b. Natural history notes from H.M. Indian
Marine Steamer Investigator. Commander R.F. Hoskyn, R.N., commanding. On the re-sults
of deep-sea dredging during the season 1890-91. Annals and Magazine ofnatural
History, ser. 6, vol. 8: 268-286.
[76] ZIMMER, C. 1914. Die Schizopoden der deutschen Sudpolar-Expedition 1901-1903.
In: E. von Drygalski (ed.), Deutsche Siidpolar-Expedition 1901-1903, XJ'. Zoologie. 7:
377-445. Pis. XXIII-XXVI.
79
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