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A key for the identification of the known larval stages of the Mediterranean
Brachyura
Article in Invertebrate Reproduction and Development · January 1998
DOI: 10.1080/07924259.1998.9652631
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A key for the identification of the known larval
stages of the Mediterranean Brachyura
DANIELA PESSANI
a
, RENATA BURRI
a
& LAURA SALTON
a
a
Dipartimento di Biologia Animale e dell'Uomo, Via Accademia Albertina 17,
1-10123, Torino, I Phone: +39 (11) 812-2374 Fax: +39 (11) 812-2374
Available online: 01 Dec 2010
To cite this article: DANIELA PESSANI, RENATA BURRI & LAURA SALTON (1998): A key for the identification of the
known larval stages of the Mediterranean Brachyura, Invertebrate Reproduction & Development, 33:2-3, 191-199
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3. Invertebrate Reproduction and Development, 33:2-3 (1998) 191-1 99
Balaban, PhiladelphiaIRehovot
0168-8170/98/$05.00O 1998Balaban
A key for the identificationof the known larval stages of the
MediterraneanBrachyura
DANIELA PESSANI*,RENATA BURRI and LAURA SALTON
Dipartimento di BiologiaAnimale e dell'Uomo, ViaAccademia Albertina 17,I-10123 Torino,I
Tel. +39 (11) 812-2374; Fax +39 (11) 812-4561
Received 14October 1996;Accepted 16December 1996
Summary
Based on the accounts by various authors, an identification key has been constructed for the
planktonic zoeae of the 70 species of Mediterranean Brachyura whose larvae are known. The
key is based on primary (spinesofthe carapace,exopodite ofthe antenna,spinesoftelson forks)
as well as secondary (setation of mouthparts, processes and spines on abdominal somites)
morphological characteristics.It allows the identification of the zoeae of 64 species plus those
of Liocarcinus genus (six species).
Key worris:Decapoda Brachyura, zoeae, identificationkey, Mediterranean Sea
Introduction decapod species whose larvae were known until 1988;
the list was accompanied by information relevant to the
The identification of planktonic crab zoeae is very
origin of the material (either planktonic or laboratory
due both the dimensions and the reared), the geographical area researched by authors
likeness among specimens belonging to different
and the number of known stages described. A list of
families/species/stages. Moreover, in some cases the
relevant species is given in Table 1, with updates until
larvae of congeneric species, described on the basis of
1996 and changes where necessary. Where no refer-
specimens hatched in the laboratory from ovigerous
ence is indicated, see Pessani and Robotti (1992).
females, are identifiable only at generic level when
collected in nature [e.g., the larvae of Inachus
dorsettensis and I. phalangium; those of six species of
Liocarcinus genus (Clark, 1983, 1984)l.
The aim of this work is to provide a key for the
identification of the Mediterranean planktonic crab
zoeae, based on the descriptions by various authors.
In the Mediterranean Seaabout 130speciesof adult
Brachyura are present (Anon., 1991),but one or more
larval stages of only 70 species are known. Pessani and
Robotti (1992) compiled a list of Mediterranean
Materials and Methods
To create the key, the descriptions by various
authors (Table 1) have been used, selecting mainly
those referring to specimens obtained from ovigerous
females and reared in the laboratory.
The key is based mainly on external morphological
characteristics which are visible, by using an optical
microscope, without the necessity of dissecting
*corresponding
author.
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4. 192 D. Pessani et al. /IRD 33 (1998) 191-199
Table 1. List of the Mediterranean species of Brachyura, whose larval stages (one or more) are known [the family sequence
follows (Bowman and Abele, 1982)l
DROMIIDAE Drorniapersonata (Linneus)
HOMOLIDAE Hornola barbata (Fabricius)
Parornola cuvieri (Risso)
LATREILLIDAE Latreillia elegans (Roux)
DORIPPIDAE Dorippe lanata (Linneus)
Ethusa mascarone (Herbst)
LEUCOSIIDAE Ebalia cranchii Leach
Ebalia nux A. Milne-Edwards
Ebalia tuberosa (Pennant)
Ilia nucleus (Linneus
Philyra globosa (Fabricius)
MAJIDAE Acanthonyx lunulatus (Risso)
Achaeus cranchii Leach
Dorhynchus thornsoni Thomson
Eurynorne aspera (Pennant)
Herbstia condyliata (Fabricius)
Inachus dorsettensis (Pennant)
Inachus leptochirus Leach
Inachusphalangium (Fabricius)
Inachus thoracicus (Roux)
Lissa chiragra (Fabricius)
Macropodia longirostris (Fabricius)
Macropodia rostrata (Linneus)
Maja squinado (Herbst)
Maja verucosa (H. Milne-Edwards)
Pisa armata (Latreille)
Pisa nodipes (Leach
Pisa tetraodon (Pennant)
Rochinia carpenteri (Thomas)
Rice and Williamson, 1977
Chhapgar, 1958
Lebour, 1928
Lebour, 1928
Ingle, 1979
PARTHENOPIDAE Heterocrypta rnaltzani(Miers)
Parthenope angulifrons (Latreille)
Parthenope rnassena (Roux)
ATELECYCLIDAE Atelecyclus rotundatus (Olivi)
CANCRIDAE Cancerpagurus Linneus
CORYSTIDAE Corystes cassivelaunus (Pennant)
PIRIMELIDAE Pirimela denticulata (Montagu)
Sirpus zariquieyi (Gordon)
THIIDAE Thiascutellata (Fabricius)
GERYONIDAE Geryon longipes (A. Milne-Edwards)
Ingle, 1985
Costlow and Bookhout, 1959
PORTUNIDAE Bathynectes longipes (Risso)
Bathynectes superba (Costa)
Callinectes sapidus (Rathbun)
Carcinus mediterraneus (Czerniavsky)
Liocarcinus arcuatus (Leach
Liocarcinus corrugatus (Pennant)
Liocarcinus depurator (Linneus)
Liocarcinus marmoreus (Leach)
Liocarcinuspuber (Linnaeus)
Liocarcinuspusillus (Leach)
Polybius henslowii Leach Paula 1985
Porturnnus iatipes (Pennant)
Portunus hastatus (Linneus)
Portunuspelagicus (Linneus) Yatsuzuka and Sakai, 1980
Portunus sayi (Gibbes) Roff et al., 1984
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5. D. Pessani et al. / IRD 33 (1998) 191-1 99 193
Table 1, continued
GONEPLACIDAE Goneplax rhomboides (Linneus)
XANTHIDAE
GRAPSIDAE
Eriphia verrucosa (Forskal)
Medaeus couchi (Couch)
Pilumnus hirtellus (Linneus)
Xantho incisusgranulicarpus (Forest)
Xantho poressa (Olivi)
Brachynotus atlanticus (Forest)
Brachynotusgemmellari (Rim)
Brachynotus sexdentatus (Risso)
Pachygrapsus marmoratus (Fabricius)
Pachygrapsus maurus (Lucas)
Pachygrapsus transversus (Gibbes)
Planes minutus (Linneus)
PINNOTHERIDAE Pinnotherespinnotheres (Linneus)
Pinnotherespisum (Linneus)
PALICIDAE Palicus caronii (Roux)
Rodriguez et al., 1992
Guerao et al., 1995
Paula, 1987
Cuesta and Rodriguez, 1994
Cuesta and Rodriguez, 1994
Cuesta and Rodriguez, 1994
Roff et al., 1984
specimens. When these features are insufficient, the
setation present on oral appendages (endopod of
maxillula; endopod and exopod of maxilla; basis,
endopod, and exopod of maxillipeds) has been
included (Fig. 1).
Identification Key to the Zoeae of the
Mediterranean Brachyura
The key has been constructed by subdividing the
species into groups on the basis of some zoeal
characteristics, i.e.:
presencelabsence of the rostral, dorsal, and lateral
spines, and processes on abdominal somites;
development and setation of antenna1exopod;
number of spines on telson forks.
In this way, the main couplets (bold) have been
obtained; these are subdivided into secondary branches
(normal types), whose minor subdivisions (figures plus
letters or asterisk) lead to a single species (Fig. 2).
S.P. Fig. 1. Schematic representation of the zoea 1 of Brachyura
and characteristics used for the key. A=zoea in toto; B =
antenna; C=maxillule; D =maxilla B-D= Portunus
pelagicus; (Yatsuzuka and Sakai, 1980). Abbreviations: A2,
eKp. antenna;' d.l.p., dorso-lateral processes; d.s., dorsal spine;
end., endopod; e.s., external spines; exp., exopod; I.s., lateral
) spine; mxl., maxillule; mx., maxilla; mxp. 1 (2), maxilliped
1 (2); p.l.s., posterio-lateral spine; p.p., posterior processes;
B C D s.p., spinous process; r.s., rostra1 spine; t.f., telson fork.
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6. D.Pessani et al./IRD 33 (1998) 191-199
Fig. 2. Schematicrepresentationof the main (bold type) and secondary (normal type) branches of the key.
Abbreviations: A2, antenna; crp., carapace;end., endopod;exp., exopod; mxl., maxillule;mx., maxilla; mxp. 1 (or 2), first
(or second)maxilliped;tls.,telson.The antenna1spinousprocess andthe abdominal somite(or somites)are indicated simply
as, respectively, spinous process and somite(or somites).
No lateral spine on crp. 2
Lateral spines on crp. (sharp, rounded, or bulge-like) 7
No dorsal spine on crp. 2A
Dorsal spine on crp. 3-
Crp. almost cylindrical; 2nd posterior process of tls., hairlike Dromiapersonata
Crp. almost spherical; a seta as 2nd posterior process of tls., not hairlike Latreillia elegans
No rostra1spine on crp. 4
Rostra1spine on crp. 5
1 external, lateral, small spine on tls. forks; 0,3 setae on mxl. end. 4A
1 external, lateral, long spine on tls. forks; 0,4 setae on mxl. end. 4B
Dorsal spine straight; spinous process without distal spines Macropodia rostrata
Dorsal spine curved; spinous process with distal spines Macropodia longirostris
Dorsal spine spinous; posterio-lateral spines on somite 3 shorter
than those on somites 4-5 Achaeus cranchii
Dorsal spine smooth; posterio-lateral spines on somites 3-5
of the same length 4B1
Without posterio-dorsal setae on somite 1 Inachus dorsettensis/I.phalangium
With a pair of posterio-dorsal setae on somite 1 4B2
Posterio-lateral spines on somite 3 shorter than those on somite 4 lnachus leptochirus
Posterio-lateral spines of the same length on somites 3-4 Inachus thoracicus
No external spine on tls. forks 5A
External spines on tls. forks 6
Dorsal and rostral spines longer than crp.; somites longer than width;
1 pair of posterior processes on tls. Dorippe lanata
Dorsal and rostral spines shorter than crp.; somites as long as wide;
3 pairs of posterior processes on tls. 5A1
A2 bearing exp. tubercle-shaped and spinous process Planes minutus
A2 rudimentary Phylira globosa (zoea 1 )
On each tls. fork 1 external spine or 1 external spine plus 1 seta 6A
On each tls. fork 2-3 external spines 6B
Spinous process shorter than rostral spine; 2 setae on proximal segment
of mxp. 1 end. Sirpus zariquieyi
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7. D. Pessani et a/./ IRD 33 (1998) 191-1 99 195
Spinous process longer than rostral spine; 3 setae on proximal segment
of mxp. 1 end. 6A1
Dorsal spine longer than crp.; rostral spine as long as 314 spinous process;
dorso-lateral processes on somites 2-3 Herbstia condyliata
Dorsal spine shorter than crp.; rostral spine as long as 115spinous
process; dorso-lateral processes on somite 2 6A2
1dorsal, small, external spine on tls. forks; somites without
posterio-lateral spines Acanthonyx lunulatus
1 lateral, well developed, external spine on tls. forks; somites
with posterio-lateral spines 6A3
A2 end. as long as 114 of A2 exp.; proximal segment of mxl. end.
without setae Lissa chiragra
A2 end. as long as 113 A2 exp.; proximal segment of mxl. end. with setae 6A4
Long posterio-lateral spines on somites 3-5 Pisa armata
Short posterio-lateral spines on somites 3 4 6A5
A2 exp. bearing a spinous terminal spine; tls. fork spinous Pisa tetraodon
A2 exp. bearing a smooth terminal spine; tls. fork smooth Pisa nodipes
Rostral spine shorter than spinous process; 3 setae on proximal
segment of mxp. 1 end. Eurynome aspera
Rostral spine not shorter than spinous process; 1-2 setae on proximal
segment of mxp. 1 end. 6B1
Rostral spine longer than spinous process; 2 setae on proximal
segment of mxp. 1end. 6B2
Rostral spine as long as spinous process; 1 seta on proximal segment
of mxp. 1 end. 6B3
2 terminal setae of different length on A2 exp.; dorsal spine long
and curved Carcinus mediterraneus
3 terminal setae of different length on A2 exp.; dorsal spine long
and straight Portumnus latipes
Lateral margins of somite 4 straight Pachygrapsus marmoratus (zoea 1)
Lateral margins of somite 4 rounded and protruding 6B4
End. of mxp. 2 3-segmented Pachygrapsus maurus (zoea 1 )
End. of mxp. 2 2-segmented Pachygrapsus transversus (zoea 1)
No dorsal spine on crp. 7A
Dorsal spine on crp. 8
Posterior margin of tls. trilobed Pinnotherespisum
Posterior margin of tls. almost straight 7A1
1 tooth on posterio-lateral corners of tls.; mxl. end. 2-segmented
with 4 apical setae Ebalia tuberosa
3 teeth on posterio-lateral comers of tls.; mxl. end. unsegmented with
3 apical setae 7A2
Setal formula of mxp. 1 end.: 2, 1, I, 2, 5; 3-4 setae on mxp. 2 end. Ebalia n u
Setal formula of mxp. 1end.: 1, 1, 0,2, 5; 2 apical setae on mxp. 2 end. Ebalia cranchii
A2 exp. flattened 8A
A2 exp. not flattened (cylindrical or rudimentary) 9
A middle-dorsal spine and 2 dorso-lateral spines on somites 2-5;
dorsal spines on tls. Homola barbata
Very small lateral spines on somites 4-5; no dorsal spine on tls. Paromola cuvieri
No external spine on tls. forks 10
External spines on tls. forks 12
Somites longer than width; 2 pairs of posterior processes on tls. 10A
Somites as long as wide;. 3 pairs of posterior processes on tls. 11
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8. D. Pessani et al. / IRD 33 (1998)191-199
Rostra1and dorsal spines forming an obtuse angle Palicus caronii
Rostra1 and dorsal spines forming a straight angle Ethusa mascarone
A2 with exp. and end.; tls. fork-shaped 11A
A2 rudimentary; tls. flattened 11B
Processes on somite 2 Brachynotus atlanticus
Processes on somites 2-3 1lAl
Crp. lateral margins denticulate; 2 subterminal spines on A2 exp.;
small processes on somite 3 inserted laterally Brachynotussexdentatus
Crp. lateral margins smooth; 2 medial spines on A2 exp.;
processes on somite 3 as minute dorsal knobs Brachynotusgemmellari
Posterior margin of tls. trilobed; mxp. 2 end. 3-segmented Pinnotherespinnotheres
Posterior margin of tls. almost straight; mxp. 2 end. unsegmented 1lBl
Lateral spines on crp. long and sharp; 1terminal seta on mxp. 2 end. Ilia nucleus
Lateral spines on crp. small and rounded; 2 terminal setae on mxp.
2 end Philyra globosa (zoea 2-3)
5 lateral spines on crp. Dorhynchus thompsoni
1 lateral spine on crp. 13
A2 exp. almost rudimentary 13A
A2 exp. developed 14
Spinous process as long as 112-314 rostra1spine; lateral spines Callinectessapidus
on crp. very short
Spinous process almost as long as (or longer than) rostral spine;
lateral spines on crp. well developed 13A1
Spinous process as long as rostra1spine Xantho incisusgranulicarpus
Spinous process longer than rostra1spine 13A2
Rostra1 and dorsal spines denticulate Xantho poressa
Rostra1and dorsal spine smooth 13A3
Lateral margins of somite 4 straight Pachygrapsus marmoratus (zoea 2)
Lateral margins of somite 4 rounded and protruding 13A4
End. of mxp. 2-3-segmented Pachygrapsus maurus (zoea 2)
End. of mxp. 2-2-segmented Pachygrapsus transversus (zoea 2)
A2 exp. as long as spinous process and bearing 2 setae 14A
A2 exp. shorter than spinous process and bearing 2-3 setae 15
3 external spines on each tls. fork; dorso-lateral processes on
somites 2-3 Pilumnus hirtellus
2 external spines on each tls. fork; dorso-lateral processes on
somites 2 4 Goneplax rhomboides
On each tls. fork 1 external spine or 1 external spine plus 1 seta 16
On each tls. fork 2-3 external spines 17
Dorso-lateral processes on somite 2 16A
Dorso-lateral processes on somites 2-3 168
3 terminal setae on A2 exp. Rochinia carpenteri
2 terminal setae on A2 exp 16A1
Dorsal and rostral spines longer than twice crp.; 2 setae on A2 exp Coiystes cassiveiaunus
Dorsal and rostral spines as long as crp.; 2 setae +2 spines on
A2 exp. Pirimela denticulata (zoea 2 4 )
Lateral spines longer than 1/2dorsal spine Heterociypta maltzani
Lateral spines shorter than 112dorsal spine 16B1
Dorsal spine stout and straight; spinous process as long as
1I2 rostra1 spine Parthenope angul~fions
Dorsal spine thin and curved; spinous process shorter than
rostra1 spine Parthenope massena
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9. D.Pessani et al. / IRD 33 (1998) 191-199 197
2 external spines on each tls. fork 17A
3 external spines on each tls. fork 19
3 terminal setae on A2 exp.; 3 setae on proximal segmentof mxp. 1end. Cancerpagurus
2 terminal setae on A2 exp.; 2 setae on proximal segmentof mxp. I end. 18
Dorso-lateralprocesses on somites2-3 18A
Dorso-lateral processes on somite 2 18B
Crp. lateral spines short; dorsal spine straight Portunuspelagicus
Crp. lateral spines long; dorsal spine curved Portunus hastatus
Spinousprocess shorterthan 112rostral spine; somites 3-5
without posterio-lateral spines Thia scutellata
Spinous process as long as 314 rostral spine; somites3-5 with
posterio-lateral spines Atelecyclusrotundatus
3 apical setae on A2 exp. 19A
2 apical setae on A2 exp. 20
Spinousprocess as long as rostral spine; 3 terminal setae of same
length on A2 exp. Maja squinado
Spinous process as long as 112rostral spine; 3 terminal setae of
different length on A2 exp. 19A1
Dorso-lateral processes on somites2-5; A2 exp. as long as 112-314
spinous process Eriphia verrucosa
Dorso-lateral processeson somites2-3; A2 exp. as long as
1I3 spinous process Medaeus couchi
Dorso-lateral processes on somite 2 20A
Dorso-lateral processes on somites 20B
On each tls. fork a group of 3 external spines inserted laterally Maja verrucosa
On each tls. fork 3 external spines, 1(or 2) inserted dorsally and
2 (or 1) inserted laterally Pirimela denticulata (zoea 1)
Dorso-lateral processes on somites2-4 Geryon longipes
Dorso-lateral processes on somites2-3 21
Crp. lateral spines much shorter than 112rostral spine; external
spines on tls. forks almost equal 21A
Crp. lateral spines as long as 112rostral spine; 1external spine on
tls. forks much longer than the others 21B
A2 exp. as long as 112spinousprocess Liocarcinus spp.
A2 exp. as long as 113spinousprocess 21A1
2 terminal setae of different length on A2 exp. Polybius henslowii
2 terminal setae of equal length on A2 exp. Portunus sayi
Long posterio-lateral spines on somites Bathynectessuperba
Short posterio-lateral spines on somites Bathynectes longipes
Discussion
The key has been found satisfactoryfor identifying
the planktonic zoeae of 62 species of crabs; the
remaining eight species (Inachus dorsettensisll.
phalangium,plusLiocarcinusarcuatus,L. corrugatus,
L,depurator, L. marmoreus, L.puber, and L.pusillus)
can be identified only to generic level (Clark, 1983,
1984). In many cases, the distinction between two
congenericspecies is based on slight differences since
both the species (at least as regards zoea 1) are
morphologically similar [e.g., Pisa tetraodonlP.
nodipes:Heegaard, 1963;Pachygrapsus rnarmoratusl
P. transversus (Cuesta and Rodriguez, 1994; Ingle,
1987)],and the accountsby two or more authors of the
same characteristics are often not described with the
same accuracy [Macropodia rostratalM longirostris
(Ingle, 1982; Lebour,1927); Inachus leptochirusll.
thoracicus (Lebour, 1928;Heegaard, 1963)l. In some
cases specimens used for the description were plank-
tonic for one species and reared for the other [e.g.,
Brachynotus sexdentatuslB. gemmellari (Bourdillon-
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10. 198 D. Pessani et al. / IRD 33 (1998) 191-199
Casanova, 1960; Guerao et al., 1995)l. According to
Roff et al. (1984), there may be differences in setation
between planktonic and laboratory reared specimens.
Characters have been used in this key that do not
change or only change slightly throughout zoeal
development. Any changes are usually most evident at
the third zoeal stage, for example, in relative propor-
tions of carapace spines and appendages setation. All
are normal expressions of progressive metamorphosis
but must be recognised when identifying zoeae from
secondtthird stages onwards. In this respect, for a few
species (Phylira globosa, Pirimela denticulata and
Pachygrapsus spp.), it was found necessary to assign
their first zoeal stage to one part of the key and their
later stages (second onwards) to another part.
More species of North Eastern Atlantic/ Mediter-
ranean brachyurans are represented in this present key
than were included in keys by Lebour (1928) to the
species occurring in the Plymouth region of the British
Isles and by Bourdillon-Casanova (1960) for the
decapods of the Gulf of Marseille.
Recently, Ingle (1992) and Paula (1996) published
keys for the identification of North Eastern Atlantic
brachyuran larvae;they took into account all the stages
described and therefore used the dimensions of the
zoeae as a discriminating feature as well as many
characters adopted in the present key.
Despite the great number of species common to the
two geographical areas (North Eastern Atlantic Ocean
and Mediterranean Sea), we preferred to construct a
new key where it was easier to include both the species
whose larvae have been more recently described and
the strictly Mediterranean speciesnot considered in the
keys previously published.
The features used in this key were chosen entirely
to enable identification of zoeae to be made to species
levels and do not reflect any systematic arrangement
(e.g., Bowman and Abele, 1982) of decapod families.
Acknowledgements
The authors are sincerely grateful to the referee for
giving his time with patience and extreme care to
improve the form of this work.
References
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Sta. Mar. Endoume, 30 (1960) 1-286.
Bowman, T.E. and Abele, L.G., Classification of the recent
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Bliss (ed.), Academic Press, New York, 1982,pp. 1-27.
Chhapgar, B.F., On the life history of Philyra globosa
(Fabricius) (Decapoda: Brachyura). Rec. Ind. Mus., 53
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Clark, P.F., The larval and first crab stages of three Inachus
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ogical and statistical analysis. Bull. Br. Mus. nat. Hist.
H ZOO^.), 44 (1983) 179-190.
Clark,P.F., A comparative study of zoeal morphology in the
genus Liocarcinus (Crustacea: Brachyura: Portunidae).
Zool. J. Linn. Soc., 82 (1984) 273-290.
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