This article appeared in the 1986 issue Vol 1. No.1 of ATOLL

This is the first article I ever wrote!

The Biology of Marine Fishes

by J. Charles Delbeek, B.Sc., M.Sc., B. Ed. I have been asked to write an ongoing column on the biology of marine fishes and how this relates to our attempts to keep them in a closed-system environment. Hopefully, much of the information I will be presenting in this column will be new to the majority of you (at least that is what Scott has asked for), however, I'm sure that there will be those of you for which this may not be true. In any event, I would appreciate as much feedback from you as possible regarding the content of these articles and anything you would like to see covered in future columns. Of course, to completely cover the biology of marine fish in such a limited series of columns would be impossible but what I will attempt to cover is that information which will be of the most concern regarding keeping fish in the marine aquarium. Towards this end the following topics will be covered: fish form and its relationship to their ecology, the effect of light on fish behavior, coloration, osmoregulation, respiration, excretion, breeding behavior and feeding behavior plus anything else you might like covered (if space and time permit).

Fish Form and Ecology: Why Does That Fish Look That Way Daddy?

The great diversity of fish form is a direct reflection of their ecological diversity. Fishes can be found in almost any body of water, be it an underground cave, beneath the arctic ice, in the deepest marine trench, in a fast moving mountain stream or in a tropical reef. In each of these environments, fish have evolved body forms, fin shapes and behavior to help them survive. Much can be learned about the ecology of a fish simply by examining its anatomical features or by watching it move through the water (Moyle and Chech, 1982). Most fishes can be classified into one of six broad categories of body shape: rover-predator, lie-in- wait predator, surface oriented, bottom dwelling fish, deep-bodied fish and eel-like fish. The rover- predator type of body form is what comes to mind when one thinks of a fish: a streamlined (fusiform) body with the fins well spaced along the body providing stability and maneuverability. A fusiform body is often indicative of a fish which is constantly on the move, searching for prey which they capture through pursuit (Moyle and Chech, 1982). An example of a rover-predator often found in marine aquariums are wrasses (family Labridae) and members of the genus Pseudanthias (e.g. Purple Queen). Lie-in-wait predators are usually piscivorous (fish eating) in nature and their morphology is well suited for the capture of fast swimming prey from ambush (Moyle and Chech, 1982). The body is often elongated, the head flattened and the mouth large. The dorsal and anal fins are typically placed far back on the body and the caudal fin is large. This arrangement of fins, and their size, gives the fish the large amount of forward thrust needed to capture their prey. Their cryptic coloration and secretive behavior make them also less visible to their intended victims. Examples commonly found in the marine aquarium are the Serranidae (groupers) and Apogonidae (cardinal fishes). Surface-oriented fish are usually small in size with an upward pointing mouth, a dorsoventrally flattened head with large eyes and a posteriorly placed dorsal fin (Moyle and Chech, 1982). This morphology is well suited for the capture of small fish and/or invertebrates which live just below the water surface. I can't think of any marine aquarium fish that fall into this category but some examples would be flying fishes and halfbeaks (Exocoetidae) as well as the freshwater killifish (genus Fundulus) and the four-eyed fish (Anableps anableps). Bottom fishes come in a wide variety of shapes and sizes but the common characteristic is an adaptation to a benthic (bottom) existence. The swimbladder in bottom dwelling fish is generally reduced or absent. Bottom fishes can be classified either as bottom rovers or bottom clingers (Moyle and Chech, 1982). Bottom rovers have compressed bodies in one plane or another, a humped back, large pectoral fins, flattened heads and terminal or subterminal mouths. The sand tilefish (Malacanthus plumeri) is an excellent example of a bottom rover as are the goat fish (Pseudupeneus maculatus) and sharks (believe it or not). Bottom dwellers are small fish with flattened heads, large pectoral fins and modified pelvic fins that allow them to adhere to the bottom. Examples of bottom fishes include the popular blennies (family Blennidae), gobies (family Gobiidae) and the Mandarin (Synchiropus splendidus; ever noticed the size of those blue pectorals?!). Deep- bodied fishes are laterally flattened fish whose body depth is at least one-third of their overall length (Moyle and Chech, 1982). The dorsal and anal fins are typically long and the pectoral fins are high on the body directly above the pelvic fins. The mouth is usually small and protrusible, the eyes large and the snout short (Moyle and Chech, 1982). Most deep- bodied fish have developed stout spines in a bid to sacrifice speed for maneuverability and protection from predators. Common marine examples of deep- bodied fish include butterflies and angelfishes (family Chaetodontidae). Eel-like fish have elongated bodies, blunt or wedge shaped heads and tapered or rounded tails; pelvic fins are either reduced or absent (Moyle and Chech, 1982). Eel-like fishes are well adapted for entering small crevices and holes in the reef. The most obvious examples commonly encountered in marine aquariums are the moray eels (genus Gymnothorax). Fin shape and placement, perhaps more than body shape, can tell you an awful lot about the habitat in which a certain fish has developed. Fin type can be placed into two main categories: paired fins (pectorals and pelvics) and unpaired fins (dorsal, anal and caudal). The pectoral fins are important in low speed maneuvering and their size and position in a particular fish give an indication as to their function in that species. Pectoral fins tent to be high up on the sides of fish which depend on precise movements for capturing prey such as the Long- nosed Butterfly (Forcipiger flavissimus). Pectoral fin length has often been assumed to increase with the importance of low speed maneuvering in the behavior of a fish (Gatz, 1972). Therefore, the longer the pectoral fins, and the more rounded they are, the more important low speed movement is in the behavior of a fish. The same can be said of the overall size of the pectoral fins. Often fish which spend a great deal of time on or near the bottom, have large pectoral fins relative to the size of the fish. Pectoral fins are often used by benthic species for support and for moving amongst bottom debris such as in the hawkfishes (i.e., Cirrhitichthys sp.) and Blennidae. In some fish the pectoral fins have lost much of their role in swimming and have taken on more of a behavioral role. For example the spreading of the pectorals as a warning display exhibited by various species of lionfish (family Scorpaenidae). Pectoral fin length is often associated with the habitat preference of a fish species being longer in rocky habitats and shorter in free swimming species. Pelvic fin size is related also to the amount of braking they perform; the larger the fin the more they are involved in braking and hovering (Gatz, 1972). Again, in some fish, the pelvic fins have become highly specialized. For example, in many of the Gobiidae, the pelvic fins have become fused to form a suction device for clinging to rocks and coral. The dorsal and anal fins are generally longer on rover-predators and deep- bodied fishes, for stability while swimming (Moyle and Chech, 1982). Short fins are often a trait of bottom-dwelling or surface-oriented fish. In some fish species, the dorsal fin has become more involved in display behavior than in swimming (e.g. Blennidae and Gobiidae). The area and shape of the caudal fin is directly proportional to the amount of swimming a fish does (Gatz, 1972). The fastest swimming fishes, such as tuna and marlin have a stiff, quarter-moon shaped fin attached to narrow caudal peduncle (Moyle and Chech, 1982). Most rover-predator fish requiring frequent sustained swimming have forked tails; the deeper the fork, the more active the fish. Deep-bodied fish and most surface- and bottom-oriented fishes have tails that are rounded, square or only slightly forked. Mouth shape, size and position are all good indicators of preferred habitat and diet in fish. In general, bottom feeding fish have mouths which point downward (subterminal) such as in the goat fish and some blennies, while surface oriented fish have mouths which point upwards (supraterminal) such as in Fundulus spp. The majority of fish, however, have mouths situated at the anterior point of the head (terminal mouths). Within the various types of coral reef fish often kept in the home aquarium, there is a host of different mouth shapes, ranging from the long tube-like mouths of many Chaetodontidae to the large, gaping mouths of the Serranidae. The size of the mouth is usually related to the size of prey consumed by a fish. Therefore, fish which feed on small invertebrates by suction often have a small, highly protrusible mouth, lined with many small teeth (Moyle and Chech, 1982). Beyond these general mouth types, fish exhibit a wide variety of mouth shapes which often reflect specialized modes of feeding and nowhere is this more evident than in reef fishes. The stability of the reef environment is often cited as the mechanism which allows such a highly diversified community to exist and to allow such highly specialized forms to evolve. Examples range from the Long Nosed Butterfly with its ridiculously elongated snout designed to reach into the smallest crevice where it can find the microcrustaceans it requires to the anglerfishes (order Lophiiformes) with their modified dorsal spine which acts as a lure and its huge mouth capable of rapid movement (an anglerfish capturing film is one of the fastest and shortest moments in nature!). This concludes the first article on fish biology. I hope that you have gained a new appreciation for what highly specialized beasts fish really are. When you next purchase a marine fish (or any fish for that matter) try to apply what you've read in this column and you won't be totally surprised when that new fish, which looked so nice and happy and fat all by himself in the dealer's tank, mercifully begins to dine on the inhabitants of your tank and soon begins to look just as happy as before. . . , but fatter! Seriously though, when purchasing a fish, try to take into account the native habitat from which it came and its normal lifestyle. Imagine being a marathon distance runner confined to 100 m track - would you be happy running that tiny track for the rest of your life? Try to buy fish best suited to the tank you have and don't expect a fish which normally roams wide areas of the reef to totally prosper in a 30 gallon aquarium. It just isn't natural for the fish. Now its time to get off the old soapbox and start thinking about the next article: "The Effect of Light on the Behavior and Well Being of Marine Fish: Who Shut Off the Lights?"