Abstract

Phyllosoma larvae taken in the Cape Verde Islands in 1998 are compared with earlier descriptions and with specimens from the Canary Islands. Previously unknown stages of Panulirus sp. are illustrated.

There is a commercial fishery for Palinuridea in the Cape Verde Islands. The species known from the archipelago are Palinurus charlestoni Forest & Postel, 1964, Panulirus echinatus S. I. Smith, 1869, Panulirus regius De Brito Capello, 1865, Scyllarides latus (Latreille, 1803) and Scyllarus pygmaeus (Bate, 1888). Palinurus mauritanicus Gruvel 1911, Scyllarides herklotsii (Herklots, 1851), Scyllarus caparti (Holthuis, 1952), Scyllarus paradoxus, Miers, 1881 and Scyllarus posteli Forest 1963 are recorded from nearby West African waters. The zoea phase of development in Palinuridea is the phyllosoma, which is readily recognizable because of dorso-ventral compression and the presence of a large cephalic shield.

Gurney referred three types of phyllosoma to Panulirus, one to Scyllarides and at least one to Scyllarus from samples in the vicinity of the Cape Verde Islands (Gurney, 1936). He compiled a key for the identification of these and other phyllosomas from the ‘Discovery’ samples. Lebour recorded phyllosomas of Panulirus sp. and Scyllarus sp. from the region of the Cape Verde Islands, but without further description or illustration (Lebour, 1959). Ribeiro described phyllosomas that he referred to Scyllarus pygmaeus from samples taken in the archipelago (Ribeiro, 1973). Robertson (Robertson, 1969) showed that the Scyllarides of Gurney (Gurney, 1936) and Stephensen (Stephensen, 1923) were actually Scyllarus, whereas specimens previously referred to as Thenus sp. were Scyllarides. Lewis (Lewis, 1951) and Berry (Berry, 1974) demonstrated intraspecific variability during development in the numbers of coxal spines, a character used in Gurney’s key. Baisre has summarized generic characteristics of phyllosomas (Baisre, 1994).

Plankton samples were taken in the Cape Verde archipelago from 23 to 28 September 1998 as part of the Macaronesia 2000 project of the Museo de Ciencias Naturales, Santa Cruz de Tenerife. Phyllosoma larvae have previously been identified as Scyllaridae or Palinuridae (Lindley et al., 2002). These specimens and specimens attributed to Scyllarus arctus from samples taken around the Canary Islands have been re-examined and their characteristics noted to provide further information on the phyllosomas of the Cape Verde Islands. Early stages of Panulirus sp., which have not previously been described for the species occurring in the Cape Verde Islands, are illustrated here (Figures 1 and 2). The identity of these specimens and those of Gurney (Gurney, 1936) is discussed.

Fig. 1.

Panulirus sp. phyllosomas from the Cape Verde Islands. (a) Stage I. (b) Stage II. (c) Stage III. (d) Second maxilla of stage I. The 1.0 mm scale bar refers to (a), (b) and (c); the 0.1 mm bar to (d) only.

Fig. 1.

Panulirus sp. phyllosomas from the Cape Verde Islands. (a) Stage I. (b) Stage II. (c) Stage III. (d) Second maxilla of stage I. The 1.0 mm scale bar refers to (a), (b) and (c); the 0.1 mm bar to (d) only.

Fig. 2.

Panulirus sp. phyllosoma stage IV from the Cape Verde Islands.

Fig. 2.

Panulirus sp. phyllosoma stage IV from the Cape Verde Islands.

Samples were taken with a triple WP-2 net with 200 μm mesh. Those containing phyllosomas are listed in Table I. Specimens were re-examined without dissection under stereomicroscopes and micrographs were taken with a digital camera. Drawings were made using the digital images and direct reference to the specimens, which are preserved intact in the collection of the Museo de Ciencias Naturales, Tenerife, for future reference.

Table I:

Samples from the Cape Verde Islands and the Canary Islands in which phyllosoma larvae were taken and examined

Sample Position Date/time Specimens 
24C 98N 16°38′59′N, 24°49′07′ 24/09/98    12:01 Scyllaridae 1 
24C 98T 16°38′90′N, 24° 49′36′W 24/9/98    15:55 Palinuridae 11 
   Scyllaridae 3 
27C 98T 16°42′492N, 24°42′49′W 27/9/98    17:13 Palinuridae 1 
28B 98T 16°31′47′N, 24°21′22′W 28/9/98    17:00 Palinuridae 1 
8A 90D 28°01′06′N, 16°44′18′W 03/08/90    11:30 Scyllarus arctus
7C 95D 28°00′24′N, 14°21′45′W 07/09/95    9:40 Scyllarus arctus
9C 95D 28°00′24′N, 14°21′45′W 09/09/95    9:40 Scyllarus arctus
10C 95D 28°00′24′N, 14°21′45′W 10/09/95    9:40 Scyllarus arctus
Sample Position Date/time Specimens 
24C 98N 16°38′59′N, 24°49′07′ 24/09/98    12:01 Scyllaridae 1 
24C 98T 16°38′90′N, 24° 49′36′W 24/9/98    15:55 Palinuridae 11 
   Scyllaridae 3 
27C 98T 16°42′492N, 24°42′49′W 27/9/98    17:13 Palinuridae 1 
28B 98T 16°31′47′N, 24°21′22′W 28/9/98    17:00 Palinuridae 1 
8A 90D 28°01′06′N, 16°44′18′W 03/08/90    11:30 Scyllarus arctus
7C 95D 28°00′24′N, 14°21′45′W 07/09/95    9:40 Scyllarus arctus
9C 95D 28°00′24′N, 14°21′45′W 09/09/95    9:40 Scyllarus arctus
10C 95D 28°00′24′N, 14°21′45′W 10/09/95    9:40 Scyllarus arctus

The 13 specimens of Palinuridae were three stage I, one stage II, four stage III, four stage IV and one later stage provisionally referred to stage VIII. All were taken in sample 24C 98T, except for two specimens of stage I which occurred in samples 27C 98T and 28B 98T (Table I). Some characteristics of these phyllosomas are listed in Table II. As Gurney’s specimens of Palinuridae from the area around the Cape Verde Islands did not include stages I–IV, these are illustrated here, in Figures 1 and 2.

Table II:

Some characteristics of palinurid phyllosomas from the Macaronesia 2000 samples in the Cape Verde Islands

 Ia II III IV VIII or IX? 
Number 
Total length (mm) 1.44–1.68 2.1 2.8–2.9 3.6–3.8 17.5 
Eye and stalk 0.4–0.6 0.88 0.9–1.2  6.8 
Eyestalk length (mm) Nil 0.28 0.4 0.4–0.5 5.1 
Eyestalk/total  0.13 0.14 0.12–0.13 0.29 
Cephalic shield width (mm) 0.66–0.76 1.0 1.3–1.4 1.14–1.8 10.3 
Thorax width (mm) 0.46–0.48 0.72 1.1–1.2 1.5–1.6 10.3 
CS/T 1.43–1.58 1.39 1.18–1.22 0.71–1.2 1.0 
Coxal spines Mp3?, P1-3 Mp3, P1-3 Mp3?, P1-3 Mp3, P1-3 P3, P4 
Sternal spines Nil Nil Nil Nil At base of Mpd3, P1-4 
Setae on maxilliped III exopod  7–8 7–11 Ca 48 
Setae on P1 exopod 9–10 12 14–15 13–20 Ca 50 
Setae on PII exopod 10 12 14–15 14–20  
PIII exopod Bud Elongate, non-setose 9–12  
PIV Nil Nil Bud Large bud Exopod elongate bud 
PV Nil Nil Nil Small bud Bud, basal segment distinct 
Pleopods Nil Nil Nil Nil Buds, slightly bifid 
Uropods Nil Nil Nil Nil or small buds Exopod and endopod developed, non-setose 
 Ia II III IV VIII or IX? 
Number 
Total length (mm) 1.44–1.68 2.1 2.8–2.9 3.6–3.8 17.5 
Eye and stalk 0.4–0.6 0.88 0.9–1.2  6.8 
Eyestalk length (mm) Nil 0.28 0.4 0.4–0.5 5.1 
Eyestalk/total  0.13 0.14 0.12–0.13 0.29 
Cephalic shield width (mm) 0.66–0.76 1.0 1.3–1.4 1.14–1.8 10.3 
Thorax width (mm) 0.46–0.48 0.72 1.1–1.2 1.5–1.6 10.3 
CS/T 1.43–1.58 1.39 1.18–1.22 0.71–1.2 1.0 
Coxal spines Mp3?, P1-3 Mp3, P1-3 Mp3?, P1-3 Mp3, P1-3 P3, P4 
Sternal spines Nil Nil Nil Nil At base of Mpd3, P1-4 
Setae on maxilliped III exopod  7–8 7–11 Ca 48 
Setae on P1 exopod 9–10 12 14–15 13–20 Ca 50 
Setae on PII exopod 10 12 14–15 14–20  
PIII exopod Bud Elongate, non-setose 9–12  
PIV Nil Nil Bud Large bud Exopod elongate bud 
PV Nil Nil Nil Small bud Bud, basal segment distinct 
Pleopods Nil Nil Nil Nil Buds, slightly bifid 
Uropods Nil Nil Nil Nil or small buds Exopod and endopod developed, non-setose 
a

Stage.

Stages I–III are illustrated in Figure 1 and stage IV in Figure 2.

These stages I–IV are attributed to Panulirus as the early stages of Palinurus spp. show more advanced development of the appendages (Cunningham, 1891; Bouvier, 1914; Berry, 1974). In particular, pereiopods 4 and 5 are both clearly visible as buds from the earliest stages of Palinurus. Also, the setation of the second maxillae (Figure 1d; Table II) conforms to the descriptions of Panulirus (Baisre, 1994). The stage VIII was similar to the Panulirus A of Gurney (Gurney, 1936). No phyllosomas referred to Palinurus sp. have been taken in the Cape Verde Islands.

Panulirus D from 14°39′N, 25°52′W shown in figure 25 of Gurney (Gurney, 1936) is a later stage than the specimens described here as stage IV. It was 5 mm long with 12 setae on the exopod of the third maxilliped and, apparently, 26, 24 and 16 (8 pairs?) of setae on the exopods of the first to third pereiopods, respectively. The fourth and fifth pereiopods were still non-setose, but longer than in the stage IV described here (Table II; Figure 2).

The intraspecific and interspecific variations in Panulirus in the numbers of coxal spines present at given development stages (Lewis, 1951; Berry, 1974) suggest that the difference between the present specimen described as stage VIII and Gurney’s (Gurney, 1936) Panulirus A need not indicate specific difference.

All the present specimens have setae on the scaphognathite of the second maxilla. Both Panulirus A and Panulirus D of Gurney (Gurney, 1936) had such setae, but in Panulirus B setae were missing until the last phyllosoma stage and the second maxilla was vestigial in the earliest stages, described as V by Gurney. We conclude that the Panulirus specimens identified here are a single species, the same as Gurney’s Panulirus A and Panulirus D from the seas near to the Cape Verde Islands. Panulirus B from this area must be the second species of this genus. Gurney states that his Panulirus D probably includes more than one species and the distributions of all three are more extensive than the known distributions (Holthuis, 1991) of Panulirus echinatus and Panulirus regius. Gurney believed that Panulirus A was most likely to be P. regius, as some of his smallest phyllosomas of this type were from the Cape Verde Islands and at that time this was the only species of the genus described from the archipelago (Gurney, 1936). In view of present knowledge of distributions and Holthuis’ (Holthuis, 1991) comments about the abundance of P. echinatus, the specific identity of these phyllosomas remains uncertain. Rearing of early phyllosomas from the eggs carried by identified females or matching genetic data from adults and larvae would resolve this uncertainty.

Kittaka and Kimura, in laboratory culture of Panulirus japonicus (Von Siebold, 1824), found that a morphological stage may include several instars (Kittaka and Kimura, 1989). The number of instars per stage increased until the eighth stage, after which the number fell again. The presence of more than one instar in a stage may explain the greater variability between the stage IV specimens observed here than in the earlier stages and variability noted by Gurney in later developmental stages (Gurney, 1936).

Two specimens of each of two successive stages of development of Scyllaridae were taken in the samples from the Cape Verde Islands. One of the earlier development stages was taken in sample 24C 98N and all the others in sample 28C 98T. All had telson spines approximately as long as the main part of the abdomen, similar to the phyllosomas attributed to Scyllarides by Stephensen (Stephensen, 1923) and figure 36 of Gurney (Gurney, 1936). The two successive stages were comparable in size and development of uropod buds to stages IV and V of Scyllarus arctus as described by Stephensen (Stephensen, 1923). However, Stephensen’s S. arctus stage IV had non-setose exopods on P4, in contrast to the present specimens (Table III). The second antennae of the earlier stage specimens from the Cape Verde Islands were slightly longer than the first antennae, more closely resembling Stephensen’s ‘Scyllarides’ than his Scyllarus arctus. Stephensen’s S. arctus had a small lateral process on the second antenna not reaching the junction of the eyestalk and eye, in contrast to the comparable specimens from the Cape Verde Islands in which the process reached beyond that point.

Table III:

Some characteristics of Scyllaridae from the Macaronesia 2000 samples from the Cape Verde (CV) and samples from the Canary Islands

 IVa (?) CV V (?) CV IV Canary V Canary 
Number 
Total length (mm) 3.6–3.8 4.5–5.4 3.5–3.7 4.4–5.3 
Eye and stalk 1.3–1.4 1.4–1.6 1.2–1.3 1.6–1.8 
Eyestalk length (mm) 0.6 0.7–0.8 0.6 0.8–1.1 
Eyestalk/total 0.16–0.17 0.15–0.16 0.16–0.17 0.18–0.2 
Antenna 1 (mm) 0.7 not segmented 1.0 segmented 0.9–1.0 not segmented 1.1–1.2 segmented 
Antenna 2 (mm) 0.8 0.9–1.2 0.6 0.9–1.1 
Cephalic shield width (mm) 2.7–3.1 3.3–3.8 2.4–2.6 3.3–3.5 
Thorax width (mm) 1.5 1.7–1.9 1.25–1.4 1.6–1.8 
CS/T 1.8–2 1.9 
Coxal spines Mpd3, P1-4 Mpd3, P1-4 Mpd3, P1-4 Mpd3, P1-4 
Setae on P1 exopod 20 26 24 24+ 
Setae on PII exopod 22–24 26–27 24+ 
Setae on PIII exopod 14–16 20 8–16 14 
Setae on PIV exopod 7–8 12–14 Nil 10 
PV Bud Bud Bud Bud 
Pleopods Nil Nil Nil Nil 
Uropods Folds/small bud Bud Bud Bud 
 IVa (?) CV V (?) CV IV Canary V Canary 
Number 
Total length (mm) 3.6–3.8 4.5–5.4 3.5–3.7 4.4–5.3 
Eye and stalk 1.3–1.4 1.4–1.6 1.2–1.3 1.6–1.8 
Eyestalk length (mm) 0.6 0.7–0.8 0.6 0.8–1.1 
Eyestalk/total 0.16–0.17 0.15–0.16 0.16–0.17 0.18–0.2 
Antenna 1 (mm) 0.7 not segmented 1.0 segmented 0.9–1.0 not segmented 1.1–1.2 segmented 
Antenna 2 (mm) 0.8 0.9–1.2 0.6 0.9–1.1 
Cephalic shield width (mm) 2.7–3.1 3.3–3.8 2.4–2.6 3.3–3.5 
Thorax width (mm) 1.5 1.7–1.9 1.25–1.4 1.6–1.8 
CS/T 1.8–2 1.9 
Coxal spines Mpd3, P1-4 Mpd3, P1-4 Mpd3, P1-4 Mpd3, P1-4 
Setae on P1 exopod 20 26 24 24+ 
Setae on PII exopod 22–24 26–27 24+ 
Setae on PIII exopod 14–16 20 8–16 14 
Setae on PIV exopod 7–8 12–14 Nil 10 
PV Bud Bud Bud Bud 
Pleopods Nil Nil Nil Nil 
Uropods Folds/small bud Bud Bud Bud 
a

Stage.

Four phyllosomas of Scyllaridae were found in the samples from the Canary Islands. These specimens had the same degree of development, and characteristics of the second antenna, as the stages IV and V of Scyllarus arctus described by Stephensen (Stephensen, 1923). These descriptions differ from those of Kurian (Kurian, 1956), who found that the larvae were smaller and the ZIV and ZV antennae were similar to those of Stephensen’s ZIII and ZIV, respectively (Kurian, 1956).

The similarity of the present Scyllarus phyllosomas taken in the Cape Verde Islands to the ‘Scyllarides’ of Stephensen (Stephensen, 1923) from the Mediterranean makes it likely that these are the same species. The distributions of the species of Scyllarus spp. in the Mediterranean and eastern Atlantic are summarized by Holthuis (Holthuis, 1991) and Udekem d’Acoz (Udekem d’Acoz, 1999). Within the Mediterranean, S. paradoxus does not occur, the only record of S. caparti is from the Adriatic, probably an escape from an aquarium (Froglia, 1979), and S. posteli is known only from the Alboran Sea. Scyllarus pygmaeus is known throughout the Mediterranean, so Stephensen’s records, which were from the eastern Basin, were probably of this species. The ‘Scyllarides’ illustrated in figure 36 of Gurney (Gurney, 1936) was taken at 14°39′N 25°52′W. This specimen, which is shown with coxal spines only on P3 and P4, was larger (6 mm) and had more setae on the exopodite of P4 than the later stage specimens in the present samples from the Cape Verde Islands and probably is the next stage.

The specific identities of Gurney’s ‘Scyllarus’ spp. remain uncertain. He stated that the specimens from the eastern Atlantic agree in all respects with Stephensen’s S. arctus. He recorded specimens from 29°27′N, 15°7′W, just north of the Canary Islands where the species is reported as frequent, whereas S. pygmaeus, the only other species of the genus recorded in the archipelago, is rare or occasional (González Pérez, 1995). Those specimens and the present specimens from the Canaries are referred to S. arctus. Gurney’s other records off the African coast from 13°25′N, 18°22′W to 15°55′S, 10°35′E are well outside the limits of the distribution of that species. Records of S. paradoxus are limited to the region from Dakar to Sierra Leone and São Tomé, S. caparti is known from as far south as 9°47′S and S. posteli is known from south-west Spain to the Congo. González-Gordillo and Rodríguez found the stage I phyllosoma of S. posteli to be indistinguishable from S. arctus, so it is suggested that Gurney’s ‘Scyllarus’ from the west coast of Africa are this species (González-Gordillo and Rodríguez, 2000).

The phyllosomas referred to Scyllarus pygmaeus by Ribeiro (Ribeiro, 1973) were later stages than the examples described here, comparable with the S. arctus stages VI–IX of Stephensen (Stephensen, 1923). The lengths of the specimens were 17.5–30.5 mm and the uropods were rounded, like those of the specimens that Gurney (Gurney, 1936) referred to Scyllarus, in contrast to the pointed uropods of the specimens that he referred to Scyllarides.

The phyllosomas of other species of Scyllarus have distinctly different morphologies. For example, S. americanus S. I. Smith 1869, S. demani Holthuis 1946 and S. martensii Pfeffer 1881 never develop elongate telson spines and the second antenna is much shorter than the first as late as stage VI (Robertson, 1968; Phillips and MacWilliam, 1986; Ito and Lucas, 1990). It would seem likely that Scyllarus phyllosomas distinctly different from those described here and by Gurney will be found in the Cape Verde Islands or nearby African shelf waters.

No true Scyllarides phyllosomas have been described from the Cape Verde Island area. However, the stage I Scyllarides latus described by Santucci (Santucci, 1925) and the ‘Thenus’ of Stephensen (Stephensen, 1923) must be the phyllosomas of S. latus, the only species of the genus known to occur in the Cape Verde Islands. Gurney’s ‘Thenus’ from 4°18′N, 16°51′W would probably be a phyllosoma of S. herklotsii.

We thank Dr Juan José Bacallado Aránega, Director of the Museo de Ciencias Naturales de Tenerife, for facilitating the cruise TFMCBM/98 as part of the Macaronesia 2000 project, and Dr Sebastián Jiménez and Mr Pedro Ortega for collaboration in sampling. The study of the phyllosoma larvae comprised part of a project funded by the UK Darwin Initiative.

REFERENCES

Baisre
,
J. A.
(
1994
) Phyllosoma larvae and the phylogeny of Pali-nuroidea (Crustacea: Decapoda): a review.
Aust. J. Mar. Freshwater Res.
 ,
45
,
925
–944.
Berry
,
P. F.
(
1974
) Palinurid and Scyllarid lobster larvae of the Natal coast, South Africa.
Invest. Rep. Oceanogr. Res. Inst.
 ,
34
,
1
–44.
Bouvier
,
E. L.
(
1914
) Reserches sur le développment post-embryonnaire de la langouste commun, Palinurus elephas.
J. Mar. Biol. Assoc. UK
 ,
10
,
179
–193.
Cunningham
,
J. T.
(
1891
) On the development of Palinurus vulgaris, the Rock Lobster or sea crayfish.
J. Mar. Biol. Assoc. UK
 ,
2
,
141
–150.
Froglia
,
C.
(
1979
) Segnalazione di alcuni crostacei nuovi o rari per l’Adriatico.
Quad Lab. Tecnol. Pesca Ancona
 ,
1
,
43
–52.
González Pérez
,
J. A.
(
1995
)
Crustáceos Decápodos de las Islas Canarias
 . Publicaciones Turquesa, Santa Cruz de Tenerife.
Gonzaílez-Gordillo
,
J. I.
and Rodríguez, A. (
2000
) First stage of Scyllarus posteli Forest, 1963 and Processa macrodactyla Holthuis, 1952 hatched in the laboratory (Crustacea, Decapoda).
Ophelia
 ,
53
,
91
–99.
Gurney
,
R.
(
1936
) Larvae of decapod Crustacea, Pt III Phyllosoma.
Discovery Rep.
 ,
12
,
400
–440.
Holthuis
,
L. B.
(
1991
) Marine Lobsters of the World.
FAO Fish. Synop.
 ,
125
, No. 13,
1
–292.
Ito
,
M.
and Lucas, J. S. (
1990
) The complete larval development of the scyllarid lobster, Scyllarus demani Holthuis, 1946 (Decapoda: Scyllaridae), in the laboratory.
Crustaceana
 ,
58
,
144
–167.
Kittaka
,
J.
and Kimura, K.-i. (
1989
) Culture of the spiny lobster Panulirus japonicus from egg to juvenile stage.
Nippon Suisan Gakkaishi
 ,
55
,
963
–970.
Kurian
,
C. V.
(
1956
) Larvae of decapod Crustacea from the Adriatic Sea.
Acta Adriat.
 ,
6
,
3
–106.
Lebour
,
M. V.
(
1959
) The larval decapod Crustacea of tropical West Africa.
Atlantidae Rep.
 ,
5
,
119
–143
Lewis
,
J. B.
(
1951
) The phyllosoma larvae of the spiny lobster Panulirus argus.
Bull. Mar. Sci.
 ,
1
,
89
–104.
Lindley
,
J. A.
, Hernández, F., Tejera, E. and Jiménez, S. (
2002
) Decápodos pelágicos (larvas y adultos) de las Islas de Cabo Verde (campaña TFMCBM/98).
Rev. Acad. Canar. Cienc.
 ,
XIII
,
87
–99.
Phillips
,
B. F.
and MacWilliam, P. S. (
1986
) Phyllosoma and nisto stages of Scyllarus martensii Pfeffer (Decapoda, Scyllaridae) from the Gulf of Carpentaria, Australia.
Crustaceana
 ,
51
,
133
–154.
Ribeiro
,
A.
(
1973
) Crustáceos decápodos capturados em águas do Arquipélago de Cabo Verde pelo navio de pesquisas ‘Walter Herwig’ em 1970.
Notas Centro Biol. Aquát. Trop.
 ,
36
,
1
–21.
Robertson
,
P. B.
(
1968
) The complete larval development of Scyllarus americanus (Smith), (Decapoda, Scyllaridae) in the laboratory with notes on larvae from the plankton
. Bull. Mar. Sci.
 ,
18
,
294
–342.
Robertson
,
P. B.
(
1969
) The early larval development of the Scyllarid lobster Scyllarides aequinoctialis (Lund) in the laboratory, with a revision of the larval characters of the genus.
Deep-Sea Res.
 ,
16
,
557
–586.
Santucci
,
R.
(
1925
) Contributo allo studio dello sviluppo post-embrionale degli Scyllaridae del Mediterraneo. II: Scyllarus arctus III: Scyllarides latus.
Mem. R. Comitato Talassogr. Ital.
 ,
121
,
1
–16.
Stephensen
,
K.
(
1923
) Decapoda-Macrura excl. Sergestidae. (Penaeidae, Pasiphaeidae, Hoplophoridae, Nematocarcinidae, Scyllaridae, Eryonidae, Nephropsidae, Appendix).
Rep. Danish Oceanogr. Exp. 1908–10
 ,
II
, D3,
1
–85.
Udekem d’Acoz
,
C.
(
1999
) Inventaire et distribution des Crustacés Décapodes de l’Atlantique nord-orientale, de la Mediterranée et des eaux continentales adjacent au nord de 25°N.
Patrimoines naturels (MNHN/SPN)
 ,
40
,
1
–383.