(MOLLUSCA, NUDIBRANCHIA). by A. M. Ayling*
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Tane (1968) lh: 25-k2 25 THE FEEDING BEHAVIOUR OF ROSTANGA RUBICUNDA (MOLLUSCA, NUDIBRANCHIA). By A. M. Ayling* INTRODUCTION Most members of the order Nudibranchia are spec• ialised carnivores feeding on sessile and encrusting animals such as hydroids, polyzoans, Porifera, ascldians and alcyonarians. All have different means of feeding eg. scraping, tearing or sucking and differ• ent modifications especially in the buccal mass. The least specialised of these grazing carnivores are the members of the Doridacea which feed on sponges. Most are brightly coloured either for camouflage when on the food sponge or to serve as a warning for predat• ors. The bright red Rostanga rufescens of Britain feeds on the encrusting red sponge Microciona in the order Poecilosclerida (Morton) and the very similar R. pulchra (MacFarland) of North America feeds on a sponge of the same order Ophlitaspongia penata (Cook 1962). The New Zealand species R. rubicunda (Cheeseman) occurs commonly on Westmere reef, Auckland, in association with three very similar sponges. Microciona coccinea Holoplocamium neozelanicum Ophlitaspongia seriata The feeding of R. rubicunda in relation to these three sponges was investigated using a number of techniques. The food of carnivores and scavengers is frequ• ently local and specific and thus chemoreception from a distance is undoubtedly important in feeding behav• iour. R. pulchra is attracted to Ophlitaspongia pennata by chemotaxis (Cook 1962) and it was thought that the same was probably true of the relationship between R. rubicunda and one or more of the above ^Department of Zoology, University of Auckland. 26 mentioned sponges. The small size of this dorid and its ease of collection make it possible to attempt the type of feeding behaviour experiments used by Stehouwer (1952), Braams and Geelen (1953) and Cook (1962) to determine its food preferences and other aspects of its feeding behaviour. This method was substantiated by making spicule mounts of gut contents and faeces to determine the sponges eaten and also by observing the animal in the field. FIG. 1. ROSTANGA RUBICUNDA. EXTERNAL CHARACTERS Rostanga rubicunda (Cheeseman l88l) is a small bright scarlet nudibranch that seems to be very close to the British R. rufescens and the North American R. pulchra, MacFarland. The mantle is covered with minute, closely packed, erect tubercules and the foot 27 extends a short distance posteriorly to this. The rhinophores are completely retractable and have 12 broad laminae which run obliquely upwards. The apex is a small projecting flat topped style. The rhino• phores of nudibranchs were developed following the loss of ctenidia and the osphradium as a replacement sensory area and are clubbed and finely plicate to increase the sensory epithelium. They contain relat• ively large lobed ganglia associated with the recep• tors. In Rostanga the rhinophores seem to be used for detecting the presence of the food sponge at a distance ie. by chemotaxis (Cook 1962) although Augersberg (1922) after experiments with food extrac• ts concluded that the rhinophores were not involved with chemoreception at a distance. However, placed as they are in an anterior, elevated position it would be fair to suggest that the function of the rhinophores is similar to that of the osphradium they replace ie. to test the water flowing over them for contained chemicals, either favourable (food) or un• favourable. The gills (branchiae) are also retract• able, eight in number, erect and bipinnate. ECOLOGY Rostanga rubicunda can be found occasionally in the sublittoral fringe on most protected rocky shores in the North Auckland area, eg. Eastern beach, Waiwera, Takapuna, Bonaccord harbour, Leigh. It is, however, limited in numbers in most of these areas by the scarcity of its food. But on Westmere reef which straddles the main tidal stream of Auckland Harbour the detritus rich waters which surge twice daily across it provide ample nourishment for a prolific growth of sponges. The three food sponges of Rostan• ga rubicunda occur quite commonly amongst these and this slug occurs in considerable numbers at this lo• cality. In this area the slugs are not found as part of the under-stone fauna but prefer more open situa• tions even though their food sponges are often pres• ent under stones. Approximately 50% of the Rostanga population were found feeding on the red food sponges. The sponges eaten by 20 of these feeding slugs were 28 identified using spicule mounts. II were feeding on Ophlitaspongia 7 " " " Microciona 2 " " " Holoplocamium Rostanga was found in considerable numbers over the entire period during which they were examined, ie. from early March to September. Powell (1957) states that it also occurs commonly over the months September to November. Thus it is probable that this animal may be found throughout the year with equal abundance. The smallest slugs collected were about 10 mms. in length and the largest 33 mms. with an average of 15-20 mms. FEEDING BEHAVIOUR Materials and Methods Rostanga rubicunda3 as has been pointed out, is always found where its food sponges are present and as sponges occur in regions where there is consider• able water movement it should be possible for the animal to trace its food by detecting secretions from the sponge and following them through the current to their source, ie. by chemotaxis. As Rostanga feeds on at least three sponges the response to each sponge would be expected to increase with the slugs' prefer• ence for that sponge as food. To test this an arti• ficial current flow was set up in the laboratory and arranged so that the slugs could be given a choice between two sponges. To achieve this seawater was run into two bowls in which the various sponges could be placed and then siphoned out into a single shallow tray in which the slugs to be tested were placed. The water was allowed to overflow from the tray and run to waste as it could not be recirculated without mixing the sponge "odours". The apparatus use is shown in Fig. 6, and diagramatically in Fig. 2. (i) Using this apparatus five fairly similar sponges from Westmere were tested against blank controls and against each other to find out which were eaten by Rostanga and the order of preference 29 MCOMINO 5CAWATER SLUG TRAY 'lilt OVF»FLOW * INFLOW FIG. 2. CURRENT FLOW APPARATUS. amongst these. These sponges were: - Ophlitaspongia seriata Holoplocamium neozelanicum Microciona coccinea Hymeniacidon perleve Suberites cupuloides Other aspects of Rostanga's feeding behaviour were tested using this apparatus, viz. (ii) Gregariousness - a few of the slugs themselves were placed in one of the sponge howls to see the slugs in the tray were affected at all. 30 (iii) The "fouling" effects of other encrusting org• anisms and algae found associated with the sponges on the reaction of the slugs was inves• tigated to see if the presence of secretions from other organisms masked those from the sponge and confused the slugs' sensory mechan• isms. (iv) The effects of varying the current speed were tested by regulating the siphoning height be• tween the bowls and the tray. (v) The effects of light or dark On the slugs' reactions were tested to see if feeding activ• ity was greater during the day than at night or vice versa. (vi) The minimum amount of sponge needed to give a definite reaction was investigated. To see if the slugs could detect the sponges without water movement another apparatus was set up in which the sponge secretions could only reach the slugs by diffusion. In this the two sponge bowls were placed at either end of the tray containing the slugs; the three linked by water bridges. These bridges were made as short as possible to minimise the diffusion path (see Fig. 3. ). After five 3 hr. experiments using sponges that evoked a considerable reaction in the current flow apparatus and 35 speci• mens of Rostanga in the tray none of the animals had responded. It is thus probable that water movement is necessary to enable Rostanga to detect and seek out its food. Results Each experiment was run for three hours and at the end of that time the animals in each sponge bowl were counted. The number of animals remaining in each tube was also noted and both figures recorded as percentages. The slugs were removed from the bowls as soon as they reached them to prevent feeding on the sponges. 31 SIPHON *—z^T^ SPONGE SPONGE BOWL •OWL FIG. 3. DIFFUSION APPARATUS Table 1. Controls Run no. Bowl Contents of bowl % in bowl %in tube I. A. nil 2. 7 0 B nil 1. 1+ 0 2 A nil 0 0 B nil 0 0 Feeding preference experiments 3 A nil 0 1. 1+ B l+. 55gm. Ophlitaspongia 26 13. 7 1+ A l+. 55gm. Ophlitaspongia 31. 5 5-5 B nil 0 0 5 A 2. 25gm. Ophlitaspongia 20. 6 l+. l B 2. 3gm. Ophlitaspongia 15-1 9. 6 6 A nil 1. 1+ 1. 1+ B lgm. Holoplocamium 6. 9 7 A 3-71gm. Holoplocamium 2. fk 6. 9 B nil 0 1. 1+ Feeding preference experiments. Gregariousness "Fouling" effects 33 (Table 1. continued) Run no. Bowl Contents of bowl % in bowl % in tube "Fouling" effects 22 b Microciona 23. 3 10 Variation of current speed 23 A Microciona with current 0 0 velocity of 0. 86 m. p. h. B Microciona with current 26 10 velocity of O. 58 m. p. h. 2k A Microciona with current 3 5 velocity of 0. 75 m. p. h. B Microciona with current 25 8 velocity of 0. 58 m. p. h. 25 A Microciona with current 27 3 velocity of 0. 7 m. p. h. B Microciona with current 7 1 velocity of O.