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Larval Development of a Warm-Water Immigrant Barnacle, Solidobalanus Fallax (Cirripedia: Archaeobalanidae) Reared in the Laboratory

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Larval Development of a Warm-Water Immigrant Barnacle, Solidobalanus Fallax (Cirripedia: Archaeobalanidae) Reared in the Laboratory J. Mar. Biol. Ass. U.K. (1999),79,1039^1044 Printed in the United Kingdom Larval development of a warm-water immigrant barnacle, Solidobalanus fallax (Cirripedia: Archaeobalanidae) reared in the laboratory Olga M. Korn* and Aleksey S. El¢movO *Institute of Marine Biology FEB RAS, Vladivostok, 690041, Russia. E-mail: [email protected]. ODepartment of Invertebrate Zoology, Moscow State University, Moscow, 119899, Russia In 1994 a warm-water barnacle Solidobalanus fallax was recorded for the ¢rst time in the UK (South- ward, 1995). The naupliar development of this immigrant species is now described, from larvae reared in the laboratory. The planktotrophic nauplii of S. fallax reached the cyprid stage 8 d after hatching, at 258C. Larval development includes six naupliar and one cyprid stage, following the typical pattern of the thoracican Cirripedia. Naupliar stages have a broad rounded convex cephalic shield without dorsal and marginal spines. Nauplii have a trilobed labrum with the small teeth found in other warm-water species. The abdominal process becomes nearly equal to the dorsal thoracic spine in stage VI. The arrangement of abdominal spines and larval setation are in the usual balanoid pattern. The signi¢cant di¡erence of the S. fallax larvae from those of the related temperate-water archaeobalanid species Hesperibalanus hesperius supports the classi¢cation of these species into separate genera. INTRODUCTION Balanus amphitrite (Clare et al., 1995). Newly hatched Solidobalanus fallax (Broch) (Archaeobalanidae) ranged nauplii I were concentrated with a beam of light and then from Angola through West Africa and Morocco to transferred to 1-l beakers containing ¢ltered seawater with added antibiotics. For this study, nauplii were cultured at Algeria, occurring at 7^220 m depth (Newman & Ross, 25 C on a mixed diet of Skeletonema costatum (approxi- 1976; Southward, 1995). In 1994 this species was recorded 8 mately 5 10 4 71 for the ¢rst time in Europe, in the English Channel o¡ cells ml ) and Thalassiosira weis£ogi (approximately 3 103 cells ml71). Larvae were preserved Plymouth, at 44^56 m depth on the valves of the queen  scallop, Aequipecten opercularis (Southward, 1995). The in 4% formalin. Drawings were made using a camera discovery was followed-up by studies of the ecology, lucida, and measurements were made with an ocular larval development and settlement of this immigrant micrometer. species. The following measurements were taken: total body The family Archaeobalanidae comprises three subfami- length, from the anterior margin of the shield to the tip lies: Archaeobalaninae, Semibalaninae and Elminiinae. of the dorsal thoracic spine; shield width (the greatest In the subfamily Archaeobalaninae the larval stages are width of the body behind the frontolateral horns); shield known for Hesperibalanus hesperius (Barnes & Barnes, 1959; length, from the anterior margin of the shield to the Korn & Ovsyannikova, 1981), Acasta spongites (Moyse, hind shield margin, excluding the posterior shield spines 1961), Chirona hameri (Crisp, 1962), and Conopea galeata in naupliar stages IV,V and VI; the length of the fronto- (Molenock & Gomez, 1972; Lang, 1979). Solidobalanus lateral horns and posterior shield spines in naupliar fallax is another member of this subfamily.The purpose of stages IV, V and VI. Measurements were also taken of this paper is to describe the larval development of cyprid length, from the anterior to the posterior cara- S. fallax under laboratory conditions and to use larval pace margins; and cyprid depth, the distance between morphology as additional information for comparative the dorsal and ventral margins of the carapace at the studies of the archaeobalanine group. deepest point. Alphabetical setal formulae follow the system of Newman (1965) and Lang (1979). MATERIALS AND METHODS Specimens of Solidobalanus fallax were obtained by RESULTS trawling in June^November 1995 from the Plymouth area Larval culture (inside the Eddystone, mostly between East and West Rutts). Adults obtained from the shells of Aequipecten Larval development of Solidobalanus fallax includes six opercularis, and from the carapace and limbs of the spider naupliar and one cyprid stage, following the typical crab, Maia squinado, also from shells of Buccinum inhabited pattern of development in the thoracican Cirripedia. by hermit crabs, were maintained in the laboratory as Development through six naupliar stages to the cyprid broodstock using techniques, previously developed for took 8 d at 258C(Table1). Journal of the Marine Biological Association of the United Kingdom (1999) Downloaded from https://www.cambridge.org/core. University of Athens, on 01 Oct 2021 at 09:39:57, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0025315499001277 1040 O.M. Korn and A.S. El¢mov Larval development of Solidobalanus fallax Table 1. Time taken at 258C for the appearance Larval morphology of the larval stages of Solidobalanus fallax. Mean sizes of the larval stages are given in Table 2. Body shape and setation of appendages are illustrated in Larval stage Day of appearance Figures 1^6. Setal formulae (after Newman, 1965) are given inTable 3. I 1 Nauplius larvae have a broad rounded convex cephalic II 1 III 2 shield with a pair of fairly long posterior spines at stages IV 3 IV^VI and without any other (dorsal or marginal) spines. V 4 The shield is only slightly longer than broad. Frontolateral VI 5 horns are of medium length, directed forward from stage Cyprid 8 Figure 2. Solidobalanus fallax. Labras (ventral view) of Figure 1. Solidobalanus fallax. Body outlines (ventral view) of naupliar stages I^VI (A^F) and thoraco-abdominal processes naupliar stages I^VI (A^F) and cyprid stage (G). (lateral view) of naupliar stages II^VI (G^K). Scale bar: 100 mm. Scale bar: 100 mm. Table 2. Measurements (mm) of the cultured larvae of Solidobalanus fallax. SWor CD Stage N TL (Mean SD) SL (Mean SD) SP (Mean SD) (Mean SD) FL (Mean SD) Æ Æ Æ Æ Æ I 4 253 3.7 127 6.1 58 5.2 II 10 428 Æ10.6 190 Æ6.1 99 Æ4.2 III 10 502 Æ8.6 250 Æ4.7 91 Æ5.0 IV 10 595 Æ21.0 346 14.5 91 7.3 321 Æ7.3 104 Æ10.9 V 10 727 Æ20.5 451 Æ10.9 127 Æ9.3 415 Æ11.1 122 Æ7.8 VI 10 885 Æ33.7 562 Æ15.6 156 Æ9.1 500 Æ13.2 125 Æ14.5 Cyprid 10 688 Æ21.4 Æ Æ 323 Æ10.5 Æ Æ Æ TL, total length; SL, shield length; SP, posterior shield spine length; SW, shield width or CD, cyprid depth; FL, frontolateral horns length; N, number; SD, standard deviation. Journal of the Marine Biological Association of the United Kingdom (1999) Downloaded from https://www.cambridge.org/core. University of Athens, on 01 Oct 2021 at 09:39:57, subject to the Cambridge Core terms of use, available at https://www.cambridge.org/core/terms. https://doi.org/10.1017/S0025315499001277 Larval development of Solidobalanus fallax O.M. Korn and A.S. El¢mov 1041 Figure 4. Solidobalanus fallax. Antennae of naupliar stages I^IV (A^D). Scale bar: 100 mm. Nauplius I (Figures 1A, 2A, 3A, 4A, 6A) Body shape is typical of stage I balanoid nauplii. The cephalic shield is pear-shaped, with well-developed fron- tolateral horns folded at angles to the long axis of the body. Thoraco-abdominal processes are short and poorly Figure 3. Solidobalanus fallax. Antennules of naupliar stages developed. Trilobed labrum is devoid of teeth and hairs. I^VI (A^F). Scale bar: 100 mm. All setae are simple. Nauplius II (Figures 1B, 2B, G, 3B, 4B, 6B) II and de£ected ventrally from stage IV. The labrum is Body shape is rounded. The frontal shield margin is trilobed, and the median lobe projects well beyond the convex. Frontolateral horns are long, slender and lateral ones. From stage II, the median labral lobe bears de£ected forward. Labrum has slender hairs. Two small several (1^3 pairs) of small teeth. Thoraco-abdominal teeth appear on the medial labral lobe. The abdominal processes are fairly long. In stage VI the abdominal process process is about 3/4 of the dorsal thoracic spine. The nearly matches the dorsal thoracic spine. The arrangement dorsal thoracic spine is barbed. The abdominal process of spines on the abdominal process is in the usual balanoid bears one pair of series-1 spines and some barbs proxi- pattern. As usually in balanoid larvae, antennae have one mally and distally. Some setae are plumose. cuspidate and one plumodenticulate seta, mandibles have two cuspidate setae. The mandibular plumodenticulate Nauplius III (Figures 1C, 2C, H, 3C, 4C, 6C) setae are poorly developed. The number and type of setae The cephalic shield has increased in size. The frontal varied little within each stage. shield margin is straighter. Frontolateral horns are shorter The diagnostic features of each larval stage can be than in stage II. The antennule bears the ¢rst preaxial summarized as follows. setae. Table 3. Setal formulae for the nauplii of Solidobalanus fallax. Antennule Antenna Mandible Naupliar stage Exopod Endopod Exopod Endopod VI S:SP:PSPP:SP:PPS:S 4P:8P PPPSP:SPP:PD:PSCP:G P:5P SSSS:PSPS:sPCP:PCP:G V S:SP:PSPP:SP:SP:S 3P:7PS PPPSP:SPP:PD:PSCP:G P:4PS SSSS:PSPS:sPCP:PCP:G IV SP:PSPP:SP:P:S 3P:6P PPPSS:SPS:PD:PSCP:G P:3PS SSSS:PSP:sPCP:PCP:G III S:PSPP:SP:P:S 2P:5P PPP:SP:PD:PSCS:G P:3PS SSS:PSS:PCP:PCP:G II SSPS:SP:P:S SP:4PS PPS:SP:PD:PSC:G P:3Ps SSS:PS:PCS:PC:G I SSSS:SS:S:S S:4S SSSS:SS:SS:SS:G S:3S SSS:SS:SS:SS:G Setal types: S, simple; P, plumose; C, cuspidate; D, plumodenticulate; G, gnathobase; s, short seta.
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