Larval Development of the Wrinkled Swimming Crab Liocarcinus corrugatus (Decapoda: Brachyura: Portunidae) Reared in the Laboratory

Larval Development of the Wrinkled Swimming Crab Liocarcinus corrugatus (Decapoda:Brachyura: Portunidae) Reared in the LaboratoryAuthor(s): Kwang Bong Kim and Sung Yun HongSource: Journal of Crustacean Biology, Vol. 19, No. 4 (Nov., 1999), pp. 792-808Published by: The Crustacean SocietyStable URL: http://www.jstor.org/stable/1549302 .

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JOURNAL OF CRUSTACEAN BIOLOGY, 19(4): 792-808, 1999

LARVAL DEVELOPMENT OF THE WRINKLED SWIMMING CRAB LIOCARCINUS CORRUGATUS (DECAPODA: BRACHYURA: PORTUNIDAE)

REARED IN THE LABORATORY

Kwang Bong Kim and Sung Yun Hong

ABSTRACT

The complete larval development of Liocarcinus corrugatus is described, based on laboratory- reared material. The species has 5, or occasionally 6, zoeal stages and a megalopal stage. Mor- phological characters of larvae of L. corrugatus were compared with this and other species of Lio- carcinus reported from other regions.

The wrinkled swimming crab Liocarcinus corrugatus (Pennant, 1777) inhabits coarse sand and gravel beaches, extending to water depths of 100 m on marl and gravel (Bour- don, 1965; Kim, 1973; Ingle, 1980). The species has a wide range of distribution. It has been reported from the Korean coast, the East China Sea, and the Pacific coast of Japan (Kim, 1973; Sakai, 1976; Dai and Yang, 1991). It also occurs in South Australia, New Zealand, the Mediterranean, and the British Isles (Bennett, 1964; Zariquiey-Alvarez, 1968; Ingle, 1980). In Korean waters L. corrugatus is the only species representing the genus.

Complete larval development of the fol- lowing five species of Liocarcinus is already known: L. marmoreus (Leach), L. holsatus (Fabricius), L. pusillus (Leach), L. depurator (Linnaeus), and L. arcuatus (Leach). These species each have five zoeal stages be- fore the megalopal stage.

Lebour (1928) described the prezoea and the first zoeal stages of Liocarcinus corrugatus (as Portunus corrugatus) from a crab obtained in the Plymouth region. Clark (1984) briefly described the five zoeal stages of the species based on laboratory-reared material from the west coast of Scotland. However, he did not provide illustrations. Wear and Fielder (1985) described the first zoea of the species from laboratory-reared material and reported the second and fourth zoeas from plankton samples collected in New Zealand waters. Hong (1988a, b) described the prezoeal stage and larval gill development based upon hatched material collected off the Isle of Man, U.K. Most of these works have been based on the species from British and New Zealand waters, and no studies have been made on the

species in the western Pacific region. The megalopal stage of L. corrugatus is still un- described.

The present study describes the zoeal and megalopal stages of Liocarcinus corrugatus collected in Korean waters, and the larvae are compared with this and other species of Lio- carcinus species reported from other regions.

MATERIALS AND METHODS

On 25 May 1995 an ovigerous crab of Liocarcinus corrugatus was collected in a crab trap in the vicinity of Hae- undae (35?10'N, 129?10'E), Korea. The crab was placed in an aquarium with running sea water and maintained in the laboratory until the larvae hatched. The eggs hatched on 28 May 1995 and the first megalopa appeared on 20 June 1995.

Seventy larvae were reared individually in glass bottles (0.13-1 capacity), and mass cultures were made in glass beakers (1 1) with sea water (31.80-32.65%o salin- ity). The glass bottles and beakers were kept at a constant temperature of 20 ? 1 ?C. The larvae were fed with newly hatched nauplii of Artemia. Molting and mortality were checked daily, at which time the larvae were transferred to freshly prepared bottles and beakers.

Larvae and molts were fixed and preserved in 4% neu- tral Formalin. A minimum of 5 specimens of each stage were dissected in ethylene glycol. The larvae were dissected using an M5 Wild binocular microscope. Setal ob- servations and drawings were made using an M 11 Wild microscope equipped with a drawing tube. The female crab and the reared larvae have been deposited in the In- vertebrate Laboratory, Pukyong National University, as PKNU-INVERT number 95-01.

Measurements taken were: (a) distance between tip of dorsal and rostral spines (TT); (b) carapace length (CL) from between eyes to posterolateral carapace margin for zoea; from rostral tip to posteromedian carapace margin for megalopa; (c) rostral spine length (RS) for zoea; (d) dorsal spine length (DS) for zoea. Measurements of at least 10 specimens were made with a calibrated ocular micrometer.

RESULTS

In the complete larval development of Lio- carcinus corrugatus there were five, or oc-

792



KIM AND HONG: LARVAL DEVELOPMENT OF LIOCARCINUS CORRUGATUS

Table 1. Duration and survival of each larval stage of Liocarcinus corrugatus at 20 ? 1?C and 31.80-32.65%o. No megalopas molted.

Duration (days)*

Larval stages Mean ? SD Range Survivors at end of stage

Z-1 5.44 ? 0.08 4-8 62 Z-2 3.93 ? 1.18 3-11 58 Z-3 7.41 ? 7.55 3-13 54 Z-4 5.09 ? 0.94 3-9 44 Z-5 -*Megalopa 3.51 ? 0.9 4-7 13 Z-5 --Z-6 4.64 ? 0.63 4-6 14 Z-6 5.14 ? 0.38 5 or 6 5 Megalopa (from Z-5) 1-3 (survival) 0 Megalopa (from Z-6) 1-3 (survival) 0

* Duration refers to survival time.

casionally six, zoeal, and one megalopal stages. Duration of each larval stage and survival of L. corrugatus are shown in Table 1. From a total of 70 zoeas, 27 reached zoea 5, of which 14 molted to zoea 6. Five individ- uals successfully molted to megalopa.

Larval Description First Zoea

Fig. 1

Size.-CL = 0.56 mm (0.49-0.62 mm); DS = 0.54 mm (0.34-0.63 mm); TT = 1.28 mm (1.11-1.40 mm); RS = 0.34 mm (0.25-0.41 mm).

Color.-Pinkish gray chromatophores on: posterior base of dorsal spine of carapace; an- teroventral, posteroventral, and posterolateral carapace; mandible; basis of first and second maxillipeds; posterolateral parts of first-fifth abdominal somites; and telson.

Carapace (Fig. 1A, B).-Dorsal spine stout, curved backward; rostral spine acute distally and longer than antennal protopod; lateral spines prominent; anteromedial process and pair of posterodorsal setae present; posterolateral marginal setae absent; eye stalks sessile.

Antennule (Fig. 1C).-Uniramous, endopod absent; exopod unsegmented with 1 short terminal seta and 3 aesthetascs of varying size.

Antenna (Fig. 1D).-Protopodal process about 2.3 times length of exopod, with 2 rows of spinules on distal half; endopod absent; exopod with 2 unequal terminal setae.

Mandible (Fig. 1E).-Incisor and molar processes developed; mandibular palp absent.

Maxillule (Fig. 1F).-Coxal endite with 7 setae; basal endite with 5 setal processes; endopod 2-segmented, proximal segment with 1 seta, distal segment with 6 (2 subterminal + 4 terminal) setae; exopod seta absent.

Maxilla (Fig. 1G).-Coxal endite bilobed with 3 + 4 setae; basal endite bilobed with 4 + 4 setae; endopod bilobed with 3 + 5 setae; scaphognathite (exopod) with 4 dense plumose setae and 1 long thick plumose posterior projection.

First Maxilliped (Fig. 1H).-Coxa without setae; basis with 10 medial denticulate setae arranged as 2, 2, 3, 3; endopod 5-segmented with 2, 2, 1, 2, 5 (1 subterminal + 4 terminal) denticulate setae; exopod 2-segmented, distal segment with 4 plumose natatory setae.

Second Maxilliped (Fig. 1I).-Coxa without setae; basis with 4 medial setae; endopod 3- segmented with 1, 1, 5 (1 subterminal + 4 terminal) denticulate setae; exopod 2-segmented, distal segment with 4 terminal plumose natatory setae.

Third Maxilliped.-Not developed.

Pereiopods.-Not developed.

Abdomen (Fig. 1A, J).-Five somites; second somite with pair of forwardly directed dorsolateral processes, third somite with small pair of backwardly directed dorsolateral processes; posterolateral margin of second somite with obtuse angle bearing 2 minute teeth; posterolateral margin of third-fifth

793



JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 19, NO. 4, 1999

A,B

E ,H-J I I~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

C,D \ N I f \w F,G

Fig. 1. Liocarcinus corrugatus (Pennant). First zoea. A, lateral view; B, frontal view; C, antennule; D, antenna; E, mandible; F, maxillule; G, maxilla; H, first maxilliped; I, second maxilliped; J, abdomen and telson. Scale bars = 0.5 mm.

794




somites with acute spines; 1 pair of minute setae on posterodorsal margin of second-fifth somites; pleopod buds absent.

Telson (Fig. 1A, J).-Each fork long; 1 fused and 1 minute secondary lateral spine; 1 dorsal articulated spine; posterior margin con- cave, armed with 3 pairs of setae; innermost pair with 4 long interior spinules located on proximal quarter of spine.

Second Zoea Fig. 2


Carapace (Fig. 2A, B).-Two pairs of anterodorsal setae between eyes; posterolateral margin with 1 anterior plumose seta + 3 posterior setae inserted near margin; eyes stalked and movable; otherwise unchanged.

Antennule (Fig. 2C).-Uniramous, endopod absent; exopod unsegmented with 1 or 2 terminal setae and 6 aesthetascs.

Antenna (Fig. 2D).-Unchanged.

Mandible (Fig. 2E).-Unchanged.

Maxillule (Fig. 2F).-Basal endite with 8 setae; exopod seta present; otherwise unchanged.

Maxilla (Fig. 2G).-Basal endite 4 + 5 setae; scaphognathite (exopod) fringed with 9 plumose setae; otherwise unchanged.

First Maxilliped (Fig. 2H).-Coxa with minute epipod bud; exopod distal segment now with 6 terminal plumose natatory setae; otherwise unchanged.

Second Maxilliped (Fig. 21).-Coxa without setae; exopod distal segment now with 6 terminal plumose natatory setae; otherwise unchanged.

Third Maxilliped (Fig. 2J).-Minute bud.

Pereiopods (Fig. 2J).-Five minute buds.

Abdomen (Fig. 2A, K).-Posterolateral margin of third-fifth somites with acute processes; otherwise unchanged.

Telson (Fig. 2A, K).-Posterior margin with additional small pair of setae; and increase in setal number as for later stages.

Third Zoea Fig. 3

Size. -CL = 0.83 mm (0.73-0.97 mm); DS = 0.82 mm (0.63-0.97 mm); TT = 2.00 mm (1.68-2.25 mm.); RS = 0.63 mm (0.51-0.71 mm).

Carapace (Fig. 3A).-Dorsal spine with 3 setae; rostral spine with 1 seta at base; 5 pairs of anterodorsal setae; posterolateral margin with 1 anterior plumose seta + 4 posterior setae; otherwise unchanged.

Antennule (Fig. 3B).-Uniramous, endopod absent; exopod unsegmented with 1 terminal seta and 6 aesthetascs.

Antenna (Fig. 3C).-Spinous process about 2 times length of exopod; endopod bud developed; exopod with 2 unequal terminal setae and minute setules.

Mandible (Fig. 3D).-Unchanged.

Maxillule (Fig. 3E).-Coxal endite with 7 or 8 setae; basal endite with 9 setal processes; otherwise unchanged.

Maxilla (Fig. 3F).-Basal endite with 5 + 5 setae; scaphognathite (exopod) with 14-17 plumose setae; otherwise unchanged.

First Maxilliped (Fig. 3G).-Coxa with 1 epipod bud; exopod distal segment with 8 terminal plumose natatory setae; otherwise unchanged.

Second Maxilliped (Fig. 3H).-Coxa with 1 epipod bud; exopod distal segment with 8 terminal plumose natatory setae; otherwise unchanged.

Third Maxilliped (Fig. 31).-Biramous with distinct endopod and exopod bud; 3 gill buds on basocoxal part (future epipod and anterior and posterior arthrobranchs).

795




A

C

F

't6,

^^^\ An\ i{//i//////

Fig. 2. Liocarcinus corrugatus (Pennant). Second zoea. A, lateral view; B, frontal view; C, antennule; D, antenna; E, mandible; F, maxillule; G, maxilla; H, first maxilliped; I, second maxilliped; J, third maxilliped and pereiopod buds; K, abdomen and telson. Scale bars = 0.5 mm.

796




B CI

A . I

G-K _

B-D

E I F

Fig. 3. Liocarcinus corrugatus (Pennant). Third zoea. A, lateral view; B, antennule; C, antenna; D, mandible; E, maxillule; F, maxilla; G, first maxilliped; H, second maxilliped; I, third maxilliped; J, first-fifth pereiopod buds; K, abdomen and telson. Scale bars = 0.5 mm.

797

I

.

I

/n1




Pereiopods (Fig. 3J).-Five small buds; first leg with one gill bud (future arthrobranch) on basocoxal part; second-fourth legs each with one gill bud (future pleurobranch).

Abdomen (Fig. 3A, K).-Six somites; dorsomedial seta on first somite; posterolateral margin of third-fifth somites with well-developed acute processes; otherwise unchanged.

Telson (Fig. 3A, K).-Apparent length re- duced by formation of sixth abdominal somite; posterior margin with 2 pairs of medial setae; otherwise unchanged.

Fourth Zoea Fig. 4


Carapace (Fig. 4A).-Dorsal spine with 6 pairs of setae; lateral spine shorter than in third stage; posterolateral margin with 1 anterior + 12-14 posterior setae; otherwise unchanged.

Antennule (Fig. 4B).-Biramous, endopod present as small bud; exopod with 2 subterminal aesthetascs and 1 or 2 terminal setae and 5 aesthetascs.

Antenna (Fig. 4C).-Endopod elongate bud about 0.5 times length of exopod; otherwise unchanged.

Mandible (Fig. 4D).-Teeth on incisor and molar processes differentiated.

Maxillule (Fig. 4E).-Coxal endite with 8 setae; basal endite with 13 or 14 setae; epipod seta now present; otherwise unchanged.

Maxilla (Fig. 4F).-Coxal endite with 3 or 4 + 4 setae; basal endite with 6 or 7 + 6-8 setae; scaphognathite (exopod) fringed with 24-26 plumose setae; otherwise unchanged.

First Maxilliped (Fig. 4G).-Coxa with 1 epipod bud and 1 seta; endopod distal segment with 6 setae; exopod distal segment with 10 terminal plumose natatory setae; otherwise unchanged.

Second Maxilliped (Fig. 4H).-Exopod distal segment with 10 terminal plumose natatory setae; otherwise unchanged.

Third Maxilliped (Fig. 41).-Endopod and exopod buds elongate, with 1 epipod bud and 2 gill buds (future arthrobranchs) on basal part.

Pereiopods (Fig. 4J).-Incipiently segmented buds; chelipeds bilobed; gill formula unchanged.

Abdomen (Fig. 4A, K).-First somite with 2 or 3 dorsomedial setae; posterolateral processes on third-fifth somites longer than in third stage; pleopod buds on second-sixth abdominal somites.

Telson (Fig. 4A, K).-Medial length in- creased compared with third zoea; posterior margin with 3 or 4 medial setae; otherwise unchanged.

Fifth Zoea Fig. 5


Carapace (Fig. 5A).-Dorsal spine shorter (relative to carapace length) and with 8 pairs of setae; 6 pairs of anterodorsal setae; posterolateral margin with 1 anterior plumose seta + 14-17 posterior setae; otherwise unchanged.

Antennule (Fig. 5B).-With 2 small setae on proximal part; endopod bud elongate; exopod with 7 or 8 subterminal and 5 terminal aesthetascs.

Antenna (Fig. 5C).-Spinous process about 2 times length of exopod; endopod elongate, more than 1.5 times length of exopod, distal margin of endopod with 1-3 minute spinules; otherwise unchanged.

Mandible (Fig. 5D).-Mandibular palp present as small bud.

Maxillule (Fig. 5E).-Coxal endite with 11-13 setae; basal endite with 17 or 18 setae; otherwise unchanged.

798




G-J a ( \

E I F~~~~~~~~~~~~~~~~~~~~~~~~~~~~~I

Fig. 4. Liocarcinus corrugatus (Pennant). Fourth zoea. A, lateral view; B, antennule; C, antenna; D, mandible; E, maxillule; F, maxilla; G, first maxilliped; H, second maxilliped; I, third maxilliped; J, pereiopod buds; K, abdomen and telson. Scale bars = 0.5 mm.

799




G"-' I ?I

B-D

E , F

Fig. 5. Liocarcinus corrugatus (Pennant). Fifth zoea. A, lateral view; B, antennule; C, antenna; D, mandible; E, maxillule; F, maxilla; G, first maxilliped; H, second maxilliped; I, third maxilliped; J, pereiopod buds; K, abdomen and telson. Scale bars = 0.5 mm.

800

4,/

I




Maxilla (Fig. 5F).-Coxal endite with 4-6 + 5-7 setae; basal endite with 8 or 9 + 7 or 8 setae; scaphognathite (exopod) with 35-40 plumose setae; otherwise unchanged.

First Maxilliped (Fig. 5G).-Elongate epipod and 2 setae on coxa; exopod distal segment now with 12 or 12+2 plumose setae; otherwise unchanged.

Second Maxilliped (Fig. 5H).-With biramous epipod bud (future epipod and podobranchs) on coxa; endopod distal segment with 6 (3 subterminal + 3 terminal) setae; exopod distal segment now with 12 or 12 + 2 plumose natatory setae; otherwise unchanged.

Third Maxilliped (Fig. 51).-With epipod and 2 gill buds (future arthrobranchs) elongate; endopod bud longer than exopod bud and segmented.

Pereiopods (Fig. 5J).-Developing; otherwise unchanged.

Abdomen (Fig. 5A, K).-First somite with 3 or 4 dorsomedial setae; second somite with 1 pair of dorsomedial setae; pleopods elongate, biramous with endopod buds on pleopods 2-6.

Telson (Fig. 5A, K).-Medial length in- creased compared with fourth stage; 1 pair of dorsal setae present in some specimens; posterior margin with 4 or 5 medial setae.

Sixth Zoea Fig. 6


Carapace (Fig. 6A).-Dorsal spine with 10 pairs of setae; 8 pairs of anterodorsal setae; posterolateral margin with 1 anterior plumose seta + 16-20 posterior setae; otherwise unchanged.

Antennule (Fig. 6B).-With 2 small setae on proximal part; endopod developing; exopod with 1 terminal seta and 5 aesthetascs and 15 subterminal aesthetascs.

Antenna (Fig. 6C).-Endopod elongate, more than 1.8 times length of exopod, distally with 5 setae; otherwise unchanged.

Mandible (Fig. 6D).-Unchanged.

Maxillule (Fig. 6E).-Coxal endite with 13 setae; basal endite with 20 or 21 setae; otherwise unchanged.

Maxilla (Fig. 6F).-Coxal endite with 5 or 6 + 5-8 setae; basal endite with 8 or 9 + 7-10 setae; scaphognathite (exopod) with 37-43 plumose setae; otherwise unchanged.

First Maxilliped (Fig. 6G).-Coxa with 1 epipod and 6 setae; exopod distal segment with 14+2 plumose natatory setae; otherwise unchanged.

Second Maxilliped (Fig. 6H).-Coxa with 1 epipod and 1 seta; endopod distal segment with 7 setae; exopod distal segment with 14+2 plumose natatory setae; otherwise unchanged.

Third Maxilliped (Fig. 61).-Epipod with 3 terminal setae; exopod 2-segmented, distal segment with 6 terminal plumose natatory setae; otherwise unchanged.

Pereiopods (Fig. 6J).-First pereiopods with several spines on propodus and dactylus; gill formula unchanged.

Abdomen (Fig. 6A, K, L).-First somite with 3-5 dorsomedial setae; second and third somites each with pair of dorsomedial setae; pleopods elongate, biramous with endopod buds on pleopods 2-6; in some specimens, endopod with minute curved processes; exopods fringed with 15 plumose natatory setae on second somite.

Telson (Fig. 6A, K, L).-With 1 pair of dorsal setae; posterior margin with 4 or 5 medial setae.

Megalopa Figs. 7, 8

Size.-CL = 2.40 mm (2.44-2.46 mm).

Carapace (Fig. 7A, B).-Longer than broad, narrowing anteriorly; rostrum directed ven-

801




D D /T r\ [*~~~~~~~~~~~~~~~

/*W~~~~~~~a Ni /

C

K

F

A,J, L

G-I,K

B-D -

E , F I -I

Fig. 6. Liocarcinus corrugatus (Pennant). Sixth zoea. A, lateral view; B, antennule; C, antenna; D, mandible; E, maxillule; F, maxilla; G, first maxilliped; H, second maxilliped; I, third maxilliped; J, pereiopod buds; K, abdomen and telson; L, abdomen showing pleopod setae. Scale bars = 0.5 mm.

802




A

K

A, B

C-G

Fig. 7. Liocarcinus corrugatus (Pennant). Megalopa. A, lateral view; B, dorsal view; C, antennule; D, antenna; E, mandible; F, maxillule; G, maxilla; H, first maxilliped; I, second maxilliped; J, third maxilliped; K, abdomen and telson; L, lateral view of abdomen and telson. Scale bars = 0.5 mm.

803



JOURNAL OF CRUSTACEAN BIOLOGY, VOL. 19, NO. 4. 1999

A ,,-- A I' L~

D -

I

I /;

-/ Lr ---

< ^

,C

1/

Fig. 8. Liocarcinus corrugatus (Pennant). Megalopa. A-E, first-fifth pereiopods; F, pleopod on second somite; G, pleopod on fifth somite; H, uropod and telson. Scale bars = 0.5 mm.

804




Table 2. Comparisons of selected zoeal characters of Liocarcinus corrugatus in the present study and in Clark (1984).

Characters Zoeal stage Present study Clark (1984)

Locality Number of zoeal stages Carapace

Setae on posterolateral margin

Antennule Aesthetascs+(setae) (terminal/subterminal)

Maxillule Setae on coxal endite

Setae on basal endite

Setae on endopod (proximal/distal segment) Maxilla

Exopod (scaphognathite) marginal setae

First maxilliped Setae on exopod distal segment

Second maxilliped Setae on endopod distal segment Setae on exopod distal segment

Abdomen Somites with dorsolateral process Dorsomedial setae on first somite

Telson Lateral spine Dorsal spine Posterior medial setae

trally and not prominent in dorsal view; sub- medial lobes rounded and raised; dorsal surface with 2 pairs of lateral tubercles; cardiac region with well-defined carina; posterome- dial margin with carina; posteroventral margin with 24-26 setae inserted near margin; eyes large and suboval.

Antennule (Fig. 7C).-Peduncle 3-segmented with 15, 5, 2 setae, respectively; endopod 2-segmented, distal segment with 2+4 setae; exopod 4-segmented with 0, 6, 4, 3 aesthetascs, respectively, third segment with 2 setae, fourth segment with 2 setae.

Antenna (Fig. 7D).-Peduncle 3-segmented with 4, 1, 3 setae, respectively; 7-segmented flagellum with (from proximal to distal) 0, 0, 4, 5, 1, 4, 4 setae, respectively.

Mandible (Fig. 7E).-Molar and incisor por- tions not differentiated; palp 2-segmented, distal segment with 15 marginal setae.

Korea 5 or 6

III IV V

I IV v V

III V II IV V

I-V

II III IV V

V

V V

I-V IV

I-V I-V III

Britain 5

5 13-15 15-18

9 11 14

3+(1) 5+(1 or 2)/2+(0)

5/6+(1 or 2)

3+(1) 4+(1)/2+(0)

5/4+(2)

7 or 8 11-13

8 13 or 14 17 or 18

1/6

7 10 9 13 17 1/6

9 14-17 24-26 35-40

10 17 23 28

12 or 14 12

6 12 or 14

5 12

2 or 3 2 or 3

2 or 3 3 or 4

2 pairs 1 pair

4

2 pairs 1 pair

3

Maxillule (Fig. 7F).-Coxal endite with 16 setae; basal endite with 26 setae; endopod 2-segmented, proximal segment with 2 setae, distal segment with 4 (2 subterminal + 2 terminal) setae.

Maxilla (Fig. 7G).-Coxal endite bilobed with 7 or 8 + 8 or 9 setae; basal endite bilobed with 10-12 + 11-13 setae; endopod subacute lobe, with 4 setae on outer margin; scaphognathite (exopod) margin with 49-56 plumose setae and 3 lateral setae.

First Maxilliped (Fig. 7H).-Epipod well-developed process, with 2+21 long setae; coxal endite with 23 setae; basal endite with 43 setae; endopod not distinctly segmented, distal part flattened, with convex outer margin with 15 setae; exopod 2-segmented, proximal segment with 6 setae, distal segment with 7 terminal plumose setae.

Second Maxilliped (Fig. 71).-Epipod with 5 setae and 1 gill bud (future podobranch); en-

805



Table 3. Comparison of larval characters in six species of Liocarcinus. (1984); 5, from Ingle (1985); T, present study.

1, from Rice and Ingle (1975); 2, from Goldstein (1971); 3, from Rice and Ingle (1978); 4, from Clark

Characters L. holsatus L. marmoreus2 L.corrugatust L. pusillus' L. depurator L.arcuatus4

Z-I Tip of dorsal to tip of rostral spine (mm) Carapace: anterodorsal setae Carapace: posterodorsal setae Maxilla: endopod setation for all stages First maxilliped: basal setae

Z-II Tip of dorsal to tip of rostral spine (mm) Maxilla: scaphognathite marginal setae Abdominal somite 3: lateral processes

Z-III Tip of dorsal to tip of rostral spine (mm) Carapace: anterodorsal setae Maxilla: scaphognathite marginal setae Abdominal somite 3: lateral processes Telson: posteromarginal setae

Z-IV Tip of dorsal to tip of rostral spine (mm) Carapace: anterodorsal setae Maxilla: scaphognathite marginal setae Abdominal somite 3: lateral processes Telson: dorsal setae

Z-V Tip of dorsal to tip of rostral spine (mm) Carapace: anterodorsal setae Maxilla: scaphognathite marginal setae Rostral spine: setae on base Abdominal somite 3: lateral processes

MEGALOPA Carapace length (mm) Antennal flagellum: number of segments Antenna: setation of segments

Pleopods 2, 5: exopod setae Uropod: exopod setae Telson: dorsal setae

0.955 absent present

5+3 10

1.1-1.3 absent present

5+3 10

1.40-1.60 9

present

1.90-2.20 4 pairs

17 absent

4 or 5 pairs

2.60-2.90 5 pairs

26 absent present

3.00-3.35 6 pairs 30-36 1 pair absent

1.86-2.16 8

0, 0, 2, 0, 5, 1, 2, 3

17-20, 14-16 9 or 10 2 pairs

1.18-1.24 absent4 present4

5+3 6( 104)

1.40 11

absent

1.70 4 pairs

13 absent 5 pairs

2.0 5 pairs

26 absent absent

3.70 6 pairs

40 2 pairs absent

2.0 6

0, 4, 5, 4, 4, 4

? 8

4 pairs


5+3 10

1.31-1.68 9

present

1.68-2.25 5 pairs 14-17

present 5 pairs

2.04-2.78 5 pairs 24-26 present absent

2.81-3.32 6 pairs 35-40 1 pair

present

2.44-2.46 7

0, 0, 4, 5, 1, 4,4 25, 18

11 2 pairs


5+3 10

1.43-1.58 9

present

1.66-1.76 4 pairs

15 present 4 pairs

2.11-2.25 5 pairs

22 absent absent

2.54-2.78 6 pairs

32 1 pair absent

1.67-1.93 7

0, 3, 0, 4, 0, 4, 3

14-17, 12-14 7 or 8 2 pairs

1.525 absent present

5+3 10

1.655 9

present

2.025 4 pairs

18 absent

4 or 5 pairs

2.505 5 pairs

27 present present

2.805 6 pairs

32 1 pair

present

2.015 85

0,0, 4, 0, 4, 1, 3, 45

9

8-105 9

1.125 9

present

1.405 4 pairs

15 present

4 or 5 pairs

1.955 5 pairs

28 absent absent

2.455 6 pairs

31 1 pair absent

0

z

0 (IT

C.'-

z

0 0

0

z p

sO

1.255 85

0, 0, 2, 0, 4, 0, 3, 55 16, ?5

7 or 85 9




dopod 5-segmented with 2, 4, 1, 9, 10 setae; exopod 2-segmented, distal segment with 8 terminal plumose natatory setae.

Third Maxilliped (Fig. 7J).-Epipod with 25 long setae and 2 gill buds (future arthrobranchs); endopod 5-segmented, ischium with 28 setae, merus with 16 setae, carpus with 9 setae, propodus with 17 setae, dactylus with 10 setae; exopod 2-segmented, proximal segment with 5 setae, distal segment with 5 terminal plumose natatory setae.

Pereiopods (Fig. 8A-E).-All segmented and sparsely spinous; first pereiopods with 2 gill buds (future arthrobranchs) at base of coxa, second pereiopods with 1 gill bud (future pleurobranch) on coxa, third pereiopods with 1 gill bud (future pleurobranch) on coxa, fourth and fifth pereiopods devoid of gill buds; first pereiopods without coxal or ischial spine, second-fourth pereiopods with coxal spine, dactylus of fifth pereiopods with 1 long straight seta and 2 long curved setae, one of them highly pectinate on distal part.

Abdomen (Fig. 7A, B, K, L).-Six somites, with 5, 5, 6, 6, 2 pairs of surface setae on somites 2-6; posterolateral margin of second-fifth somites broadly truncate.

Pleopods (Figs. 7A, L, 8F, G).-Somites 2-5 each with pair of biramous pleopods; endopod unsegmented with subterminal hooks on internal margin; exopod fringed with 25 plumose natatory setae on second somite; fifth somite with 18 plumose natatory setae.

Uropod (= pleopod on sixth somite) (Figs. 7A, L, 8H).-Endopod absent, exopod 2-segmented, proximal segment with 2 setae, distal segment with 11 plumose natatory setae.

Telson (Fig. 8H).-Broader than long, sub- quadrate with 2 pairs of medial dorsal setae.

DISCUSSION

Zoeas of the wrinkled swimming crab Li- ocarcinus corrugatus were described by Lebour (1928), Clark (1984) and Wear and Fielder (1985). Lebour (1928) described partially the first zoeal stage of the species (as Portunus corrugatus) from the Plymouth region. In chromatophores and TT length (distance between tip of dorsal and rostral spines), her description agrees closely with

that of the present study. Wear and Fielder (1985) partially described the first zoea of the species from laboratory-reared material, and also described the second and fourth zoeas from plankton samples collected in New Zealand waters, and assigned them to this species. The description of the zoeas of the species given by them is too brief for comparison with the present study. Clark (1984) described five zoeal stages of the species based on laboratory-reared material from the west coast of Scotland. The present study shows minor differences from Clark's account in some features. These are chiefly in the number of zoeal stages and the setation of the posterolateral margin of the carapace, anten- nular exopod, and scaphognathite (Table 2). It is unknown whether these are genotypic or the result of dissimilar rearing conditions, but they do not seem to amount to specific differences.

The zoeal stages of Liocarcinus corrugatus were compared with those of five other species of Liocarcinus. Clark (1984) and In- gle (1985) were unable to discover satisfac- tory characters to distinguish zoeas of species of Liocarcinus in all stages. The present study has shown that the zoeas of L. corrugatus are not easily distinguished from those of the other five species of Liocarcinus. Particularly in the early zoeal stages, there may be con- siderable variation in the length of carapace spines, even in larvae from the same female. Distinctions based on small differences in spine length are, therefore, unreliable. Table 3 gives comparison of the six species.

Specific larval characters of Liocarcinus are recognizable only in the megalopal stage. The megalopal stage of L. corrugatus has been compared with those of the other five species of Liocarcinus. These comparisons are listed in Table 3. The features that distinguish the megalopal stage of L. corrugatus are: (1) antennal flagellum 7-segmented, with setal formula 0, 0, 4, 5, 1, 4, 4; (2) exopods on second and fifth pleopods with 25 and 18 setae, respectively; and (3) exopods on uropods with 11 setae.

The complete larval development of Lio- carcinus is known for five species (Table 3). Based on these works, it is evident that these species have five zoeal stages and one megalopal stage. However, in the larval development of L. corrugatus, there are occasionally six zoeal stages.

807




ACKNOWLEDGEMENTS

The authors thank Dr. D. I. Williamson (Port Erin Ma- rine Laboratory, Isle of Man, England) and Dr. C. Sankarankutty (Universidade Federal do Rio Grande do Norte, Natal, Brazil) for their comments on an earlier draft of the manuscript. We also thank Paul F. Clark (Natural History Museum, London, England) for his valuable crit- icism and suggestions for improving the manuscript.

LITERATURE CITED

Bennett, E. W. 1964. The marine fauna of New Zealand. Crustacea Brachyura.-New Zealand Department of Scientific and Industrial Research Bulletin 153: 1-119.

Bourdon, R. 1965. Inventaire de la fauna marine de Roscoff: D6capodes-Stomatopodes.-Travaux de la Station Biologique de Roscoff 16: 1-45.

Clark, P. F. 1984. A comparative study of zoeal morphology in the genus Liocarcinus (Crustacea: Brachyura: Portunidae).-Zoological Journal of the Linnean Society of London 82: 273-290.

Dai, A., and S. Yang. 1991. Crabs of the China Seas.- China Ocean Press, Beijing, China. Pp. i-xxi + 1-682.

Goldstein, B. 1971. Developpement larvaire de Macropipus marmoreus (Leach) en laboratoire (Crus- tacea, Decapoda, Portunidae).-Bulletin du Museum national d'Histoire naturelle, Paris 42: 919-943.

Hong, S. Y. 1988a. The prezoeal stage in various deca- pod crustaceans.-Journal of Natural History 22: 1041-1075.

. 1988b. Development of epipods and gills in some pagurids and brachyurans.-Journal of Natural History 22: 1005-1040.

Ingle, R. W. 1980. British crabs.-British Museum (Nat- ural History), Oxford University Press, Oxford, En- gland. Pp. 1-222.

. 1985. Larval development of the red swimming crab, Bathynectes longipes (Risso, 1816) (Crustacea: Decapoda: Portunidae).-Bulletin of the British Mu- seum of Natural History (Zoology) 49: 239-255.

Kim, H. S. 1973. Anomura, Brachyura. Illustrated en- cyclopedia of fauna and flora of Korea.-Samwha Pub- lishing Company, Seoul, Korea. 14: 1-694. [In Korean.]

Lebour, M. V. 1928. The larval stages of the Plymouth Brachyura.-Proceedings of the Zoological Society of London 2: 473-560.

Pennant, T. 1777. British Zoology. IV. Crustacea, Mol- lusca, Testacea.-London, England. Pp. i-viii + 1-136.

Rice, A. L., and R. W. Ingle. 1975. A comparative study of the larval morphology of the British portunid crabs Macropipus puber (L.) and M. holsatus (Fabricius), with a discussion of generic and sub-familial larval characters within the Portunidae.-Bulletin of the British Museum of Natural History (Zoology) 28: 123-151.

, and . 1978. The larval development of the portunid crab Macropipus pusillus (Leach) reared in the laboratory.-Bulletin of the British Museum of Natural History (Zoology) 33: 287-296.

Sakai, T. 1976. Crabs of Japan and the adjacent seas.- Kodansha Ltd., Tokyo, Japan. 3 volumes; English text, pp. i-xxix + 1-773; Japanese text, pp. 1-461.

Wear, R. G., and D. R. Fielder. 1985. The marine fauna of New Zealand: larvae of the Brachyura (Crustacea, Decapoda).-New Zealand Oceanographic Institute Memoir 92: 1-89.

Zariquiey-Alvarez, R. 1968. Crustfceos Decapodos Ib6ricos.-Investigaci6n Pesquera 32: i-xv + 1-510.

RECEIVED: 4 November 1998. ACCEPTED: 17 February 1999.

Address: Department of Marine Biology, Pukyong Na- tional University, Pusan 608-737, Korea.

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Documents

Larval Development of the Wrinkled Swimming Crab Liocarcinus corrugatus (Decapoda: Brachyura: Portunidae) Reared in the Laboratory