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Ontogenetic Development of Lipophrys Trigloides (Pisces: Blenniidae), with Some Notes on the Spawning Behaviour

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Ontogenetic Development of Lipophrys Trigloides (Pisces: Blenniidae), with Some Notes on the Spawning Behaviour J. Mar. Biol. Ass. U.K. (2005), 85, 185^188 Printed in the United Kingdom Ontogenetic development of Lipophrys trigloides (Pisces: Blenniidae), with some notes on the spawning behaviour P O Cla¤udia Faria* ,Fa¤tima Gil and V|¤tor C. Almada* *Unidade de Investigac°a‹ o em Eco-Etologia, Instituto Superior de Psicologia Aplicada, R. Jardim do Tabaco 34, O P-1149-041 Lisboa, Portugal. Aqua¤rio Vasco da Gama, R. Direita, Dafundo, P-1495 Lisboa, Portugal. P Corresponding author, e-mail: [email protected] The full developmental sequence from egg to juvenile of Lipophrys trigloides in controlled conditions is described. In addition, some notes on the spawning behaviour of adults are provided. Embryonic develop- ment lasted 13^14 days at 21.00^23.008C, 16^17 day s at 17.25 ^18.508C and 17^18 days at 14.00^16.508C. Newly hatched larvae measured 4.8 mm, had the mouth and anus opened, pigmented eyes and almost no yolk. Most larvae settled between day 39 and 42 after hatching at 16.0^17.0 mm total length (TL) and showed full juvenile pigmentation patterns and behaviour 5 to 10 days later (17.0^18.0 mm TL). A summary of the characteristics of the eggs and larvae that may help in the distinction between L. trigloides and Lipophrys pholis, a very close related species, is also provided. INTRODUCTION fed twice a day with Brachionus sp. enriched with protein Selco (Artemia Systems) and algae, which were replaced Lipophrys trigloides (Valenciennes, 1836) is a relatively by Artemia sp. nauplii by day 58. Larvae were collected common rocky intertidal ¢sh species along the Portuguese daily, anaesthetized (ethylene glycol monophenyl ether rocky shores. Although there is some information about Merck) and photographed daily until metamorphosis. All the biology of this species (e.g. Nieder & Zander, 1993; larval measurements correspond to total lengths. Nieder, 1997), nothing is known about its breeding beha- viour and development. In this paper the full developmental sequence of RESULTS L. trigloides, from egg to juvenile, is described for labora- Captive males guarded batches from September to July. tory reared ¢sh. Some notes on the behaviour of the breeding pair are also presented. Breeding males presented a general black coloration with white swollen tips of the dorsal ¢ns. During the breeding period, both males established territories in crevices and MATERIALS AND METHODS under stones, where spawning took place. On some occa- sions males were observed excavating the substrate under Eggs and larvae were obtained from a captive group of the nest stone. Males defended and ventilated the devel- six ¢sh (2 males with 7.0 cm and 7.7 cm TL respectively; 4 oping eggs with undulating movements of the body and females between 6.6 and 8.7 cmTL), maintained sinceJune tail, until hatching. On one occasion it was possible to 2001 at Vasco da Gama Aquarium (Lisbon). The 600-l observe the spawning event. The female presenting tank was illuminated with £uorescent light (60 W), from normal coloration pattern and the black male rubbed the 0900 to 1900 hours. The bottom of the tank was covered nest wall with the genital papilla and performed pectoral with a sand layer and several large £at stones and shells were provided as shelter and breeding sites. ¢n beatings and high amplitude movements of the tail. The complete sequence of embryonic development is These movements were repeated several times, alternating based on three batches spawned on vertical stones and the tank wall (for temperature ranges see Table 1). A sample of Table 1. Temperature range of the batches used for (A) eggs was removed daily for description from the stone embryonic development and (B) for larval development. guarded by the male, by aspiration with a long pipette. The eggs were observed under a Nikon stereomicroscope, Mean (8C) Range (8C) SD N photographed by a Nikon FX-35DX camera and preserved in bu¡ered 5% formalin. The egg capsules A. Spawning. were opened and the embryos were distended to allow 17/02/2003 15.21 14.00^16.50 0.07 19 more detailed observations. 22/04/2003 17.96 17.25^18.50 0.47 19 Full larval development is described from one batch that 29/05/2003 21.71 21.00^23.00 0.51 14 B. Hatching. hatched on 15 December 2003.We used three other batches 14/02/2003 18.25 16.50^20.50 1.31 37 to con¢rm the patterns of larval development (see Table 1). 24/04/2003 21.55 18.50^24.75 1.83 35 Upon hatching, larvae were reared in glass 30-l tanks, 29/05/2003 16.98 16.00^18.00 0.57 20 illuminated with £uorescent light (18 W) 24 h per day. A 15/12/2003 17.25 17.00^17.50 0.24 41 constant £ow of seawater was maintained. Larvae were Journal of the Marine Biological Association of the United Kingdom (2005) 186 C. Faria et al. Early ontogeny of Lipophrys trigloides Table 2. Ontogenetic events of embryonic development of Lipophrys trigloides inorderof¢rstappearance(indays):(1)embryo recognizable; (2) cephalic and caudal dilatation; (3) eye lens; (4) brain; (5) notochord di¡erentiation; (6) brain lobes; (7) notochord; (8) myomeres; (9) heart beatings; (10) pigmented eyes (beginning); (11) auditory vesicles; (12) otoliths; (13) tail bud free of the yolk; (14) gut di¡erentiation; (15) median ¢nfold; (16) embryo movements; (17) pectoral ¢n buds; (18) mouth di¡erentiation; (19) hatching glands; (20) anus visible but closed; (21) mouth visible but closed; (22) opercula di¡erentiation; (23) pectoral ¢ns developed; (24) anus opened; (25) mouth opened; (26) mandibles di¡erentiation; (27) opercula opened; (28) gas bladder; (29) hatching. 8C 1234567891011121314151617181920212223242526272829 14.00^16.50d1d2d3d3d3d4d4d4d5d5d5d7d7d7d7d7d8d8d9d9d10d10d10d10d11d11d13d14d18 17.25^18.50d2d3d3d3d3d3d4d4d4d4d4d5d5d5d5d5d7d8d9d9d10d10d10d10d11d11d13d14d17 21.00^23.00d1d2d2d2d2d2d3d3d3d3d3d3d4d4d4d4d5d6d6d6d7d7d7d7d9d9d9d12d14 A B C Figure 1. Eggs collected at di¡erent developmental stages: (A) day 6: beginning of the eyes pigmentation; (B) day 9: embryo with developing pectorals; (C) day 14: embryo prior to hatching (from batch spawned at 14.00^16.508C). with resting periods. In general, both individuals alter- to about a half, a white and opaque cup-like structure is nated their movements over the nest wall. present. Both males guarded multiple clutches. From February The main ontogenetic events of embryonic development to June, one of the males was observed guarding eggs at di¡erent temperatures are shown in Tables 2 and 3 for seven distinct periods of time, separated by 1^12 respectively. Figures 1 and 2 present eggs and larvae days without eggs. During each guarding period collected at di¡erent developmental stages. The length of eggs spawned at di¡erent times were present. Some the embryonic period varied with temperature: 13^14 days batches were placed adjacent to recently laid eggs and at 21.00^23.008C, 16^17 days at 17.25^18.508C and 17^18 others ¢lled the areas left vacant by hatching. days at 14.00^16.508C. Recently laid eggs are golden brown and transparent. Newly hatched larvae (Figure 2) measured 4.8 mm TL They are spherical with a somewhat £attened surface (SD¼0.19, range: 4.5^5.0 mm TL, N¼10). The anus and (Figure 1). The major axis is 1.3 mm (SD¼0.16, range: mouth were opened, with formed lips and di¡erentiated 1.12^1.50 mm, N¼51), and the minor axis is 0.86 mm jaws. The yolk was almost fully absorbed or absent. The (SD¼0.06, range: 0.80^0.92 mm, N¼51). From the base liver was developed, the eyes were fully pigmented and the gas bladder was formed, but not completely ¢lled. Table 3. Ontogenetic events of larval development of Lipophrys trigloides in order of ¢rst appearance (days after hatching): (1) exogenous feeding; (2) ¢lled gas bladder; (3) caudal ¢n rays; (4) nothocord starts to £ex; (5) ventral ¢n buds; (6) anal ¢n rays; (7) 2nd dorsal ¢n rays; (8) notochord £exion completed; (9) ¢rst dorsal ¢n rays; (10) ventral ¢n rays; (11) median ¢n fold reabsortion; (12) ossi¢ed vertebrae; (13) larvae begun to contact the aquarium bottom; (14) most larvae settled on the bottom; (15) ¢rst development of juvenile pigmentation. Size-ranges are also indicated. 8C 123456789101112131415 16.00^18.00 d2 d2 d11 d13 d17 d17 d17 d17 ^^^^^^^ 4.8^6.9 mm 7.0^8.9 mm 17.00^17.50 d2 d2 d9 d10 d10 d15 d15 d16 d16 d20 d21 d21 d31 d42 d53 5.0^6.9 mm 7.0^8.9 mm 11.0^12.9 mm 416.0 mm 16.50^20.50 d2 d2 ^^^^^^^^^^d24 d39 d40 5.0^6.9 mm 416.0 mm 18.50^24.75 d2 d2 ^^^^^^^^^^d36 d40 d42 5.0^6.9 mm ??? Journal of the Marine Biological Association of the United Kingdom (2005) Early ontogeny of Lipophrys trigloides C. Faria et al. 187 A B C D Figure 2. Larvae collected at di¡erent developmental stages: (A) day 1: newly hatched larva (5.0 mm TL); (B) day 14: 8.5 mm TL; (C) day 35: 12.8 mm TL; (D) day 41: juvenile (16.5 mm TL). Pectoral ¢ns were di¡erentiated and the sagittae and the caudal peduncle), on the throat, and at the cephalic lapilli otoliths were visible. The opercula were opened. region where melanophores extended from between the Larvae presented heavy peritoneal pigmentation and eyes to the dorsal region and opercula. At day 3 (5.8 mm), 8^10 rows of melanophores on the pectoral ¢ns. Ventrally, di¡use yellowish pigmentation, which subsequently there were 1^2 melanophores on the base of the pectoral extended all over the head and body, was present.
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