S.A. Hesp

Influence of selected factors on the dietary compositions of three targeted and co-occurring temperate species of reef fishes: implications for food partitioning.

The dietary compositions of three medium to large targeted fish species, which co-occur over reefs in temperate waters of south-western Australia, were determined. These data were then used to ascertain statistically the extent to which body size, season and habitat influence the diets of these species and the degree to which food resources were partitioned among and within those species, and thus reduced the potential of interspecific and intraspecific competition. On the west coast, Bodianus frenchii (Labridae) and Epinephelides armatus (Serranidae) spent their whole life over prominent limestone reefs, as did Glaucosoma hebraicum (Glaucosomatidae) in all but juvenile life, when it lived over low-relief, limestone substrata. The dietary composition of each species changed with increasing body size, which, in G. hebraicum, was particularly pronounced at c. 300 mm total length (L(T)) and therefore at the size when this species shifts habitat. When the three species co-occurred over the same reefs, their dietary compositions were significantly different, with that of B. frenchii being by far the most discrete, reflecting a far greater contribution by sedentary taxa. Thus, the diet of B. frenchii was distinguished from those of the other two species in containing substantial volumes of bivalve and gastropod molluscs and echinoid echinoderms and essentially no teleosts. Although the diets of G. hebraicum and particularly E. armatus were dominated by teleosts, and especially for larger individuals, the former species ingested greater volumes of cephalopods and small crustaceans. The pointed jaws of B. frenchii, with their forwardly directed and interlocking anterior incisors, are ideally adapted for biting and retaining their invertebrate prey, which are attached to or reside within reef crevices. In contrast, the mouths of G. hebraicum and E. armatus are broader and rounder and contain numerous small, slender and inward-pointing teeth. These teeth, in conjunction with prominent backward-curved canines in E. armatus, facilitate the capture and retention of fish prey. Observations in situ indicate that G. hebraicum is a suction feeder, while E. armatus is predominantly a ram feeder. Although reef environments on the west and south coasts differ, the diet of B. frenchii on these coasts differed only slightly. Interspecific differences in diet, combined with size-related changes in dietary compositions and the occupation of different habitats by juvenile and adult G. hebraicum, reduce the potential for competition for food resources among and within B. frenchii, G. hebraicum and E. armatus and thus helps facilitate the coexistence of these species which historically have been abundant over reefs in south-western Australia.

Relationships between the movements, growth, age structures, and reproductive biology of the teleosts Sillago burrus and S. vittata in temperate marine waters

Sillago burrus and S. vittata both use sheltered, nearshore shallow waters (≤ 1.5 m) as nursery areas. However, the juveniles of the former species remain there for only a few months, before migrating into deeper waters (5 to 15 m) as they increase in size, whereas some juvenile S. vittata do not undergo a similar migration until considerably later. S. burrus rarely exceeded 2 yr of age and was never found beyond 4 yr of age. Although only a small number of S. vittata exceeded 2 yr of age, a few individuals of this species were caught between 4 and 7 yr old. The maximum and asymptotic lengths of S. burrus (251 and ∼180 mm, respectively) were far lower than those of S. vittata (325 and ∼320 mm, respectively), whereas the growth coefficients (K) were much higher for the former species, i.e. ∼2.4 vs ∼0.4. Virtually all S. burrus, and also those S. vittata that moved into deeper waters early in life, spawned at the end of their first year of life. Since relatively few S. burrus reached 2 yr of age, the attainment of almost full size by the end of their first year of life enables a relatively large number of eggs to be produced by fish at the end of their first year-for many, their only spawning period. Those individuals of S. vittata that remained in their shallow nursery areas until the end of their first year of life, did not reach maturity until the end of their second year of life. The proportions of mature gonads and the numbers of yolk-vesicle and yolk-granule oocytes and post-ovulatory follicles in ovaries were far higher in both S. burrus and S. vittata during December to February than in any other month, demonstrating that these two species spawn largely in these summer months. During this period, the ovaries of individual S. burrus and S. vittata often contained post-ovulatory follicles, as well as yolkvesicle and yolk-granule oocytes that ranged widely in size, strongly suggesting that both species are multiple spawners. During the spawning period of S. burrus, the ovaries possessed large numbers of hydrated oocytes and no post-ovulatory follicles or the reverse situation, and the oocytes tended to form several relatively discrete size groups. This indicates that S. burrus produces eggs in batches and that the spawning of the members of this species is synchronised. The presence of large numbers of yolk-granule oocytes with migrating nuclei in the ovaries of many S. vittata at certain times suggests that this species is also a batch- and synchronised spawner. Comparisons between the results of the present study and past work emphasise that the relationships between the timing of offshore movements and the sizes and ages at first maturity vary considerably amongst whiting species.

Growth and reproductive biology of the foxfish Bodianus frenchii, a very long‐lived and monandric protogynous hermaphroditic labrid

Samples of the foxfish Bodianus frenchii, collected over reefs on the lower west and south coasts of Western Australia, contained individuals ranging up to 78 years old. Although B. frenchii is far smaller than many other species within the Labridae, its maximum age is the greatest yet recorded for this highly speciose family and, together with Achoerodus gouldii, provides an example of a temperate hypsigenyine with exceptional longevity. Length and age compositions of females and males and the histological characteristics of gonads of a wide length range of individuals demonstrated that B. frenchii is a protogynous hermaphrodite. Furthermore, as, on both coasts, the length of the smallest male was greater than that at which all females had become mature, B. frenchii is a monandric protogynous hermaphrodite, i.e. all of its males are derived from functional females. Attainment of maturity by females is related more to length than age, whereas the reverse is true for sex change. On the basis of Schnute growth equations and length-to-body mass regression equations, the predicted length at age and body mass at length of fish on the south coast were greater than those on the west coast throughout life. Although B. frenchii spawns daily during the main spawning season, which extends from October to February on both coasts, its fecundity at any given length is substantially greater on the south than on the west coast. The more rapid growth of juveniles and earlier attainment of maturity by B. frenchii on the south coast than on the warmer west coast, together with maturation at a similar size on both coasts, run counter to the trends observed in many species and certain ecological theories regarding the relationships between life-cycle traits and latitude and temperature. The attainment by B. frenchii of a larger body length at age, of greater body mass at length and of greater fecundity at both length and body mass in fish on the south than on the west coast strongly suggests that conditions on the former, cooler coast are more favourable for this labrid, which belongs to a sub-genus whose other species typically live in cool, deep, temperate waters.

Do the maximum sizes, ages and patterns of growth of three reef‐dwelling labrid species at two latitudes differ in a manner conforming to the metabolic theory of ecology?

The size and age data and patterns of growth of three abundant, reef-dwelling and protogynous labrid species (Coris auricularis, Notolabrus parilus and Ophthalmolepis lineolata) in waters off Perth at c. 32° S and in the warmer waters of the Jurien Bay Marine Park (JBMP) at c. 30° S on the lower west coast of Australia are compared. Using data for the top 10% of values and a randomization procedure, the maximum total length (L(T) ) and mass of each species and the maximum age of the first two species were estimated to be significantly greater off Perth than in the JBMP (all P < 0.001) and the maximum ages of O. lineolata in the two localities did not differ significantly (P > 0.05). These latitudinal trends, thus, typically conform to those frequently exhibited by fish species and the predictions of the metabolic theory of ecology (MTE). While, in terms of mass, the instantaneous growth rates of each species were similar at both latitudes during early life, they were greater at the higher latitude throughout the remainder and thus much of life, which is broadly consistent with the MTE. When expressed in terms of L(T), however, instantaneous growth rates did not exhibit consistent latitudinal trends across all three species. The above trends with mass, together with those for reproductive variables, demonstrate that a greater amount of energy is directed into somatic growth and gonadal development by each of these species at the higher latitude. The consistency of the direction of the latitudinal trends for maximum body size and age and pattern of growth across all three species implies that each species is responding in a similar manner to differences between the environmental characteristics, such as temperature, at those two latitudes. The individual maximum L(T), mass and age and pattern of growth of O. lineolata at a higher and thus cooler latitude on the eastern Australian coast are consistent with the latitudinal trends exhibited by those characteristics for this species in the two western Australian localities. The implications of using mass rather than length as the indicator variable when comparing the maximum sizes of the three species and the trends exhibited by the instantaneous growth rates of those species at different latitudes are explored. Although growth curves fitted to both the L(T) and masses at age for the males of each species lay above those for their females, this would not have influenced the conclusions drawn from common curves for both sexes.

Life cycle characteristics of the Blue Morwong Nemadactylus valenciennesi, compared with those of other species of Cheilodactylidae

The demonstration by this study that the life cycle characteristics of the reef fish Nemadactylus valenciennesi differed markedly from those of other cheilodactylids led us to compare these differences and hypothesise on their evolutionary implications. The distribution, length and age compositions, growth and the length and age at maturity of N. valenciennesi in south-western Australian waters have been determined. The maximum age (21 years) of N. valenciennesi (maximum total length = 984 mm) is far less than those of the much smaller Cheilodactylus fuscus, Nemadactylus macropterus (both ∼40 years) and Cheilodactylus spectabilis (97 years). Yet N. valenciennesi, N. macropterus and C. spectabilis mature at a similar young age (3-7 years). While, on the basis of data for other species, all three species mature at lengths consistent with their asymptotic lengths, the last two species mature at a far earlier age than would be predicted from their long life spans. The early attainment of maturity by N. macropterus and C. spectabilis may reflect a response to high juvenile mortality or constrained adult growth, whereas their long life spans could reflect a response to variable recruitment in the past or compensation for limited annual reproductive output due to very constrained adult growth.

Implications of the divergent use of a suite of estuaries by two exploited marine fish species

Biological characteristics of the marine species King George whiting Sillaginodes punctatus and Australian herring Arripis georgianus in three seasonally open estuaries (Broke, Irwin and Wilson Inlets), one permanently open estuary (Oyster Harbour) and one normally closed estuary (Wellstead Estuary) on the south coast of Western Australia have been determined and compared. Sillaginodes punctatus enters the seasonally and permanently open estuaries early in life and reaches total lengths (L(T)) >280 mm at which it can be legally retained and thus contributes to commercial and recreational fisheries in these systems. This sillaginid almost invariably emigrates from these estuaries before reaching its typical size at maturity (L(T50)) and does not return after spawning in marine waters. In contrast, virtually all female A. georgianus (≥ 98%) in the three seasonally open estuaries and the majority in the normally closed (89·5%) and permanently open estuaries (83%) exceeded the L(T50) of this species at maturity, reflecting the fact that the nursery areas of this species are predominantly located much further to the east. Although adult females of A. georgianus in seasonally open and normally closed estuaries had developed mature ovaries by autumn, at which time they were prevented from migrating to the sea by closure of the estuary mouths, this species did not spawn in those estuaries. The oocytes in their ovaries were undergoing extensive atresia, a process that had been incipient prior to oocyte maturation. As the adult females of A. georgianus in the permanently open Oyster Harbour at this time all possessed resting gonads, i.e. their oocytes were all previtellogenic, the adults that were present in that estuary earlier and were destined to spawn in autumn must have emigrated from that permanently open estuary to their marine spawning areas prior to the onset of gonadal recrudescence. The body masses at length of A. georgianus, which were almost invariably higher in summer and autumn than in winter and spring, were greater in the very productive environments of the seasonally open and normally closed estuaries than in the less productive and essentially marine environment of Oyster Harbour and coastal marine waters. In general, the same pattern of differences between water bodies was exhibited by the growth of A. georgianus and by the more restricted data for body mass at L(T) and growth of S. punctatus. Despite an increase in anthropogenic activities in Wilson Inlet over the last two decades, the growth of both species was very similar to that recorded 20 years earlier. The fisheries implications of the results for the two species are discussed.

Timing of growth zone formations in otoliths of the snapper,Chrysophrys auratus, in subtropical and temperate waters differ and growth follows a parabolic relationship with latitude

&NA; Chrysophrys auratus was collected from one sub‐tropical and two temperate regions spanning >2400 km along the coast of Western Australia (˜23.5‐35.5° S). Marginal increment analysis demonstrated that, while a single opaque zone is formed in the otoliths of C. auratus each year, the period of deposition varies among regions. An opaque zone was formed in May to early September in the sub‐tropical upper west coast, and thus when water temperatures were declining to their minima. In contrast, opaque zone formation occurred 3 months later in August to December in the temperate lower west and south coasts, when water temperatures were rising from their minima. The length and age distributions differed markedly among populations of C. auratus, with the strongest year classes varying among the three regions. Thus, it is likely that year class strength of C. auratus throughout its distribution along the coast of Western Australian is mostly related to local environmental conditions. Chrysophrys auratus grew far less rapidly and attained a smaller size in the warmer upper west coast than in the cooler temperate regions of the lower west and south coasts. A collation of data on C. auratus from ten populations in Australia and three in New Zealand showed that growth is greatest towards the mid‐latitudes of its geographic range, i.e. at ˜31° S. Estimates of mean lengths at specified ages thus exhibit a parabolic relationship with latitude, with reduced growth (i.e. edge‐of‐range effects) occurring towards the latitudinal margins of the distribution of this sparid.