brisbane river

THE

BRISBANE RIVER

A SOURCE-BOOK FOR THE FUTURE

INVERTEBRATE AND INTERTIDAL COMMUNITIES OF THE ES-TUARY 1977; Stejskal1984; Stejskal and Chamberlain 1984; Davie 1986). They are dominated, at least on soft substrates, by polychaete worms, tiny bivalves, gastropod snails, and peracarid crustaceans (Amphipoda, Isopoda, and Tanaidacea). Planktonic communities, and in particular the crustacean component, have also received some attention. Bayly (1964, 1965, 1966) studied the taxonomy of copepods and the seasonal variation of their populations, and Greenwood (197 4, 1976, 1977, 1978, 1979) studied copepod communities of Moreton Bay, with one of his study stations being at the mouth of the Brisbane River. Kennedy (1975) reported on the planktonic communities of the Fitzroy River, and although it is a slightly more tropical estuary his results are relevant to the Brisbane River.

Estuaries and associated mangrove and wetland areas link the land and the sea. They are major sources of primary production and thus support and fertilize much of the coastal life of our shallow shelf waters. However an estuary, and particularly a sub-tropical or tropical one, is a most difficult environment for animals or plants to live in because of the sudden changes that occur. Environmental transitions in an estuary are a complex of interrelated factors including changes in temperature, salinity, osmotic pressure, pH, Eh or redox potential, surface and interfacial tension, partial pressure of oxygen and certain other gases, radiation, turbidity, suspended solids, turbulence or other movements of water, cations, anions find toxic substances, and nutrients (ZoBell1973). These conditions are cons antly changing from time to time and place to place. The Brisbane River is tidal for over 85 km although a measurable salinity is seldom discernible more than 70 km upstream. It is strongly poik.ilohaline seasonally, such that during the low rainfall / low flow periods of winter and spring salinities may exceed 25 p.p.t. over 30 km from the mouth. Conversely during the high rainfall / high flow periods of the summer months, particularly following the passage of cyclones, freshwater may extend to Within 20 km upstream of the mouth (Bayly 1965; Stephenson 1968).

Patterns in Diversity Davie (1986) and Stephenson and Campbell (1977) have reported on Serpentine Creek, a small estuary which used to lie just to the north of the Brisbane River, but which was destroyed by land reclamation for the Brisbane Airport. They found that the benthic communities followed the expectations of a model of physical control. The upstream areas had large salinity fluctuations and had fewer species, lower diversities (Standard Shannon and Gleason measures) and significantly lower

BENTHIC AND PLANKTONIC INVERTEBRATES

Peter Davie, B.Sc., M.Sc., is a marine zoologist and Curator of Crustacea at the Queensland Museum. He is Vice-President and a past President of the Australian Littoral Society, and has long been interested in the study of estuarine systems. He has researched estuarine invertebrate communities, and has published widely on the taxonomy of crabs.

The benthic invertebrate communities of the Brisbane River and nearby estuaries have been moderately well studied (Snelling 1959; Straughan 1967; Hailstone 1972, 1976; Campbell eta/. 1977; Park 1979; Stephenson and Campbell 131

The Brisbane River

153'00'E

•

Slllinity during low flow conditions

¢

Salinity during high flow conditions

27'30'E

Figure 1: A map showing the variation in salinities in the Brisba ne River during high and low flow conditions. The

11umbers refer to the stations sampled by Boesch (1977). salinity change of 30 down to 18 p.p.t. during low flow conditions but experiences salinities below 5 p.p.t. during high flow conditions. The third zone occurred between Jindalee and Westlake and marks a transition to high constancy and dominance of freshwater species, although several estuarine species are still common. Salinities are not more than 2 p.p.t. above Westlake. Several estuarine benthic species (peracarid crustaceans in particular) are found virtually to the limit of tidal influence, almost 20 km above Goodna and actually above the limit of tidal influence in some of the tributaries. Bayly (1965) and Stevens (this volume) have commented on the high chloride ion content (brought through the atmosphere from the sea) of the fresh-water runoff in the Brisbane drainage basin which apparently allows deep penetration of the estuary by some species. Freshwater species, chiefly tubificid oligochaetes, were dominants at Boesch' s Westlake and Goodna sampling stations. A number of very euryhaline marine species and estuarine endemics occur throughout the estuary and several estuarine endemics and fre shwater species are restricted to the u pper reaches of the estuary.

community stability than the downstream sites where salinity variations were much less extreme. Upstream however there were much greater total abundances which were due to a few species which were estuarine endemics or opportunists. Kennedy (1975) working in the Fitzroy River, similarly found that maximum plankton volumes occurred when diversity was loyvest, between salinities of 12 p.p.t. and 18

p.p.t. Zonation Boesch (1977) conducted a study on the zonation of subtidal benthic invertebrates within the Brisbane River. He found marked changes in faunal composition at three areas. The first is about 10- 15 km from the mouth (his Station 2) where although salinities are still high (34 p.p.t. for most of the year) the effect of summer flooding lowers it to around 10 p.p.t. He suggests that the accelerated coenoclinal change is mainly a response to the estuarine (salinity) complex-gradient. 'A large portion (58 %) of those species recorded from Station 1 were not found as far up-estuary as Station 2 and the attenuation of within-habitat species diversity and richness is greatest in this zone.' Between the City and Long Pocket there is a second zone of accelerated change, particularly between St. Lucia and Long Pocket as several euryhaline species abundant down-estuary are no longer found . This zone is coincident with a

Factors Affecting Diversity and Distribution In general the up-estuary limits of marine and estuarine organisms are set by tolerance of low 132

-

Invertebrate and Intertidal Communities

species, euryhaline oportunists, estuarine endemics, and freshwater species. The stenohaline marine and freshwater species are severely restricted in their distribution by osmotic pressure and only enter the main body of the estuary during periods of drought or flooding when conditions are suitable for them. The euryhaline marine species may be more abundant in the lower reaches of an estuary than elsew ~ and equate with 'equilibrium' species (Gr~~s~~ and Grassle 1974). Euryhaline opportunists are apparently disfavoured in ' equilibrium' polyhaline habitats by biotic interaction. Estuarine endemics are normally restricted, in homoiosmotic estuaries, to upper low salinity areas, but Boesch (1977) believes that in poikilohaline estuaries such as the Btisbane River the effect of poikilohalinity is to displace this group and the other groups downstream thus allowing the domination of much of the estuary by estuarine endemics. Basically he suggested that while physiological tolerances set limits to upstream penetration, biological interactions largely set lower (down-estuary) distributional limits. The results of Davie' s (1986) study in the Serpentine Creek largely support this hypothesis. Although salinity variability appears to be the dominant physical control, other factors could play an interactive role. Straughan (1967) suggested that increased current flow combined with increased turbidity could have a 'sand-blasting' effect, and this may have been the reason for the mortality and lack of settlement of fouling organisms that he observed in the Brisbane River in 1963. Sediment characteristics have long been recognised as presenting a complex of limiting values influencing the distribution of benthic fauna (e.g. Driscoll and Brandon 1973; Kay and Knights 1975; Whittaker and Levin 1977; Coleman and Cuff 1980). Aziz and Greenwood (1982) found that juveniles of the prawn Metapenaeu s bennettae preferred to bury themselves in either silt, or a very fine or fine sandy substrate of less than 250 microns particle size. They felt that this may be a significant factor in allowing juvenile prawns to settle in the upper reaches of the Brisbane River. Coarser sediments with a greater range of grain sizes can also create a large number of potential niches (Gray 1974; Johnson 1974) and thus lead to greater community diversity. Related to this Davie (1986) suggested that more unstable sediments at one of his study

salinity. Similarly the down-estuary limits of f shwater species are generally set by t~~erance of high salinity . Little information is vailable on salinity tolerances of species in :outheastern Queensland estuaries. Barnes (1967) conducted salinity tolerance experiments on fivP. species of intertidal crabs in the Brisbane River. He found that the range of salinity that can be tolerated by a species is not always a reliable guide to the environmental conditions in w hich it is found- sometimes its upstream penetration is more governed by other fact ors such as lack of suitable substrates. Bayly (1965) fo und that populations of a species of copepod of the genus Pseudodiaptomus were invariably on the marine side of those of another, Gladioferen s. He concluded that Pscudodiaptomus is a euryhaline marine component of the brackish water planktor•. Gladioferans pcctinatus on the other hand, maintained po pulations in the upper reaches of the river during summer despite the fact that this is the tim e of maximum flow, but extended downstream into water of higher·salinity during the colder months . Local stu dies have shown that salinity limits distribution not only because of the adults ability to tolerate changes, but also because of the tolerances of larvae and juveniles, and the effect of salinity on reproduction. Hodge (1963) investigated the biology of two species of mysid shrimp that are present in the River in large numbers- Rhopalophthalmus brisbanensis and Gastrosaccus dakini. She found that populations of the two species differ in salinity preference; in breeding times; and in times of peak population . For breeding, G. dakini requires a salinity range of only 2- 10 p.p.t., while juveniles of R. brisbanensis require a range of 7- 17 p.p.t. Salinity has also been shown to be critical for other crustaceans. Fielder and Greenwood (1983) found that under laboratory conditions the larvae of the sesarmid crab Bresedium brevipes, which occurs as an adult only in the upper estuary, only metamorphosed to the megalopa stage at salinities of 10 p.p.t. Aziz and Greenwood (1981) reported that although juveniles of the prawn Metapenaeus bennettae can tolerate salinities from 1.0- 62.0 p.p.t. they are normally restricted to low salinity waters even though adults are not so restricted. Boesch (1977) from his River work gave a broad model of estuarine zonation based on fiv e species' distributional classes viz. stenohaline marine species, euryhalin~ marine 133

The Brisbane River

sites could be contributing to higher diversity. Intermediate levels of disturbance from both abiotic fluctuations and predation have been suggested as causes of increased diversity (Caswell1978; Connell1978) and as stabilising factors (Murdoch and Oaten 1975) by preventing monopolisation of competitively dominant species.

temperature fell to its minimum of 18° C. The types of bacteria present changed from bacilli in lower salinity, higher temperatures to streptococci during higher salinities and lower temperatures. Over the two summers of his study there was a significant 'population crash' of planktonic species in November- December when the inorganic Nand P levels hit their peak. This was accompanied by the occurrence of 'sewage fungus' organisms of bacterial, fungal, and algal types. Some of these organisms secreted mucous in sufficient quantity to change the viscosity of the w ater andtrap other planktonts.

Seasonality In the Brisbane River and Moreton Bay, approximately annual cycles in population size of benthic invertebrates have been commonly reported with minimum populations around March and maxima between September and December (Hailstone 1972; Park 1979; Stephenson eta/. 1974, 1976, 1978; Davie 1986). It seems that there are also recognisably ' different seasonal communities although the results of the studies done are not totally consistent. Davie (1986) found that there was a sequential pattern of changing community structure but within this a three-season year could be recognised : early summer (1 - 2 mths, c. Nov.- Dec.); summer/ early autumn (3- 4 mths, c. Dec. - Mar.); and ' mid-year' (6- 7 mths, c. Apr.- Oct.). Park (1979) working at the mouth of the Brisbane River, recognised only a two-season year in summer I autumn and winter I spring, while Stejskal (1984) working on intertidal fauna at Cribb Island, found three consecutive time groups over three years of sampling, with no indication of seasonality. Davie (1986) found that, to some extent, seasonal changes in community structure were due to several estuarine endemic or opportunistic species (in particular the crustacean Apscudes estuarius and the bivalve Notospisula trigonella) which apparently achieve large populations at times when other species are osmotically disfavoured, and decline sharply in numbers when conditions favour competition from other species. Not~sp~~ ula trigonella has been regarded as an en~ment opportunist by Pearson and Rosenburg (1978) and Stejskal (1984) observed that benthic populations of N. trigonella reproduce continually such that larvae are constantly available for settlement whenever abiotic conditions are suitable. Kennedy (1975) investigated the planktonic communities of the Fitzroy River estuary and found that there was a marked increase in bacterial count over the winter period although salinity rose from 3 - 12.5 p.p.t. and

Factors Responsible For Seasonal Patterns High rainfall with concurrent reduced salinity has been considered a causal factor for fa unal seasonality in local studies, i.e. Park (1979) working on macrobenthos at the mouth of the River; Vohra (1965) and Stejskal (1984) w orking on intertidal fauna at Victoria Point and Cribb Island respectively; Davie (1986) working on the macrobenthos of Serpentine Creek; and Young and Wadley (1979) studying epibenthic fauna in Moreton Bay. Food availability may also be an important factor. Ruello (1973) noted the importance of seasonal freshwater run-off through mangrove and reed swamps in carrying enormous quantities of nutrients and plant detritus into the Hunter River estuary. This material plays a particularly important role in the nutrition of juveniles of the school prawn (Metapenaeus macleayz) which is also an important commercial species in Moreton Bay. Davie (1986) noted that sediment scouring by summer flooding could also be effecting the seasonal community structure; and as has been noted earlier, competition and predator I p rey relationships also appear to cause changes in community structure. Kennedy (1975) fou nd that plankton volumes in the Fitzroy River reached a minimum 6- 8 weeks after the annual invasion by the large rhizostome medusa Catostylus. He felt that this filter feeder must be responsible for substantial predation. The Effects of Major Floods Several studies have shown that major floods have a severe effect on the benthic fauna (Stephenson et al. 1977; Saenger et al. 1980; Davie 1986). The recovery from such events can be very slow and in the lower estuary appears to exceed three years (Davie 1986). The pattern 134

Invertebrate and Intertidal Communities

S~ARMINE

L.W.S.

ZONE

OCYPODINE ZONE

1--------------------

Figure 2: A profile of the river bank at about Bulimba showing the different crab zones described by Snelling (1959).

of recovery is typically one of a linear increase in species numbers accompanied by a logarithmic increase in populations (Saenger et a/. 1980; Davie 1986). Davie (1986) found however that in the upper estuary recovery was apparently very rapid with a' stable' community structure re-established well within the first year. This difference he attributed to the fact that the species inhabiting this part of the estuary suffer similar events on a seasonal basis and therefore must be more capable of rapid re-colonization.

desiccation during emergence time. Most species are more or less zoned from mean Low Water Spring tidal level (L.W.S.) to mean High Water Spring (H.W.S.) depending on their degree of terrestrialization. The crabs and molluscs are the best known and the most dominant of the intertidal surface animals so these will be briefly discussed. Crabs Snelling's (1959) study shows that although populations may be patchy in distribution they still generally fall into a recognisable zonation up the bank. Four zones can usually be identified, at least towards the mouth of the River. They are named after the predominant types of crab to be found in each zone. Species diversity decreases with increasing distance from the water. (1) Sesannine Zone: extends up the shore above High Water Neap (H.W.N.). The substrate is usually dry gravelly mud with loose stones. The most conspicuous crabs present are Sesarma erythrodactyla and Helice haswellianus although a number of other species of the family Grapsidae occur in varying abundances depending on the location.

INTERTIDAL FAUNA The intertidal fauna has been reported on by Snelling (1959), Moverley (1984) and for the n( earby Serpentine Creek by Campbell et al. 1 97~) . Although Snelling and Moverley f tud1ed only the crabs, it appears that the d~cto.rs governing the occurrence and thstnbution of the fauna as a whole are much d'e same as those that have already been lScussed for benthic invertebrates. For ~xample there are eighteen species at the mouth hut only two 37 kms upstream. There is owever the added dimension of avoiding

5

135

The Brisbane River

(2) Ocypodi11e Zo11e : from about mean High Water (H .W.N.) down to Mean Sea Level (M.S .L.). The most obvious species are Fiddler Crabs (Uca coarctata, Uca vomeris, Uca /ongidigitum) and the semaphore Crab (Hcloeciu s cordiformis). He/oecius prefers a firmer substrate and tolerates some gravel. (3) Upper Macrophthalmine Zone : from about M.S.L. down to mean Low Water Neap (L.W .N.) but requires soft fine mud. The dominant species are Australoplax tridentata in the lower reaches and Paracleistostoma mcneilli in the higher reaches. Other species occur but not so commonly. (4) Lower Macrophthalmine Zone : from L.W.N. down to L.W.S. The mud here is very soft with many shallow pools, and is inhabited in great numbers by Macrophthalmus setosus. As Snelling (1959 : 79) says: 'The zones d escribed are by no means constant. The observations suggest that the main factor limiting the vertical distribution is not the period of exposure but the water content of the soil. This would account for the fact that Euplax [= Australoplax] triden tata is often found above H .W.S., but is restricted to situations where the

mud is very wet. Similarly Heloecius cordiformis is only found where the mud is firm enough for its large burrows to be made. Thus the zonation pattern as outlined may become much modified.' Moverley (1984) reiterated the fact that the boundaries between the zones can become quite confused. Moverley (1984) found seasonal variation in populations of several common species and postulated that juvenile recruitment occurred only in the lower estuary, and upstream populations relied for their maintenance on the immigration of maturing crabs. He therefore considered seasonal increases at a particular locality to be either the result of recruitment or immigration, while seasonal decreases are the result of mortality of senile individuals or emmigration. Maximum species diversities occurred between February and March . Molluscs There are no published accounts of the ecology of the mollusca n fauna of the Brisbane River, however the report of Campbell et al. (1977) fo r Serpentine Creek gives a good indication of what would be found . The molluscs appear to

Figure 3: This large Fiddler Crab (Uca vomeris) is common on the banks of the River. Its claw is bright ora nge. 136

Invertebrate and Intertidal Communities

fall into three broad categories (a) species of the mangrove / saltmarsh zone, (b) species occurring on muddy banks, mud flats and sand flats, and (c) log and stump infauna . (a) In the mangrove / saltmarsh zone species are, in the main, distributed according to factors such as suitable substrate, shade, moisture, salinity etc., although the seaward mangrove fringes at the mouth of the river may have some species not found further back in the drier areas and vice-versa. A coarse subdivision into surface fauna, subsurface fauna, and fauna of tree trunks is possible with some overlap. (b) The mud bank and mud / sand flat species are all limited to areas below Mean Sea Level. Polinices sordidus, Pyrazus ebeninus, Na ssarius spp. and Vclacumantu s australis are all common surface dwellers but les ~-fre.quent in sandier areas. Infaunal bivalves hke Tellina species and Sanguinolaria donacioides are similarly less abundant in sand. Presumably the amou nt of organic matter present or the moisture holding capacity affect the distribution. (c) Logs, dead stumps, mangrove trunks and exposed roots have a small specialized fauna . Species involved are bivalves like Saccostrea commercia/is, Crassostrea sp ., and Modiolu s pulex; the gastropod Bembicium auratum; and the 'ship worms' (teredinids). Most require well inundated, relatively saline sites.

as far as Hamilton (Steele 1976) however one species Aegiceras corniculatum (the River Mangrove), now at least, penetrates about 64 kms upstream . This change has been attributed to greater intrusion of salt water following dredging of the river. Despite this upstream penetration the upper mangroves do not support the wide variety of organisms that they do in the lower estuary as the salinity regime is too severe for the greater number of animals. Also Aegiceras is restricted to only patchy narrow fringes. Five species are represented in the estuary. The two most common are Avicennia marina (Grey or White Mangrove) and Aegiceras corniculatum (River Mangrove). Excoecaria agallocha (Milky Mangrove) is well represented particularly in the upper estuary, but is nowhere abundant. Rhizophora stylosa (Red Mangrove) and Bruguiera gymnorhiza (Orange Mangrove) have a limited distribution in the lower estuary (Hegerl1975) . Mangroves are of immense importance in maintaining bank stability and normal shoreline gradients, but they are also susceptible to destruction by being undercut by wash from shipping, particularly on steeper banks at low tide. There is also evidence to suggest that they are sensitive to smothering by silt. Both Watson (1928) and Heger! (1975) attributed mangrove deaths to the covering of their pneumatophores (air roots) by silt after heavy flooding. Another significant threat to mangrove communities is interference with normal patterns of freshwater runoff via channelisation. Mangrove species are varyingly dependent on freshwater and the balance between fresh and saline water is critical to their distribution on the shore (Saenger et al. 1983).

FLORA The productivity of an estuary is dependent on a variety of plants including macrophytes such as mangroves, seagrasses, sedges etc; microand macroepiphytic algae; benthic micro- and macroalgae; and phytoplankton . There have be~n virtually no studies on any of these ih the Bnsbane River !

Mangroves are now well known to play an essential role in high riverine and shallow coastal productivity. Degradation of leaves and organic detritus derived from plant material are directly utilized by the benthos (Heald 1971; Hughes and Sherr 1983; Poore and Rainer 1974). Data are not available for the Brisbane River, but Miller (1976) working in a small mangrove catchment in North Queensland found that soluble forms of organic carbon and particulate matter of less than 1-2 microns derived primarily from the mangrove belt, appeared to contribute a considerable proportion of the total organic carbon available

Mangroves There are no maps available of the distribution ~ ma~groves in the Brisbane River although owhng (1986a) has provided an excellent re · B Vtew of the mangrove vegetation of Moreton ay. h The mangrove forests of the Brisbane River (~ve been reduced by approximately 50% enry et al. 1987) through dredging, c. hann ertsatton . and urban development. There tssom e evt'd ence to suggest that before E uropean settlement mangroves extended only 137

The Brisbane River TABLE 1: Algal species recorded from the Serpentine Creek estuary by

Atherton and Dyne (1977) DIVISION Cyanophyta Oscillatoriaceae Chroococcaceae Nostocaceae DIVISION Chlorophyta CLADOPHORALES Cladophoraceae ULOTRICHALES Ulvaceae Monostromaccae SIPHONALES Caulerpaceae Codiaceae DIVISION Phaeophyta ECTOCARPALES Ectocarpaceae DIVISION Rhodophyta GIGARTINALES Gracilariaceae Rhabdoniaceae CERAMIALES Delesseriacea

Rhodomelaceae

Microcoelus lyngbyaceus Schizothrix sp. Anacystis marina Chroococcus turgidus Scytonema rhizophorae

Cladophora social is Rhizoclonium implexum Ulva lactuca Enteromorpha clathrata Blidingia minima Caulerpa fastigiata Boodleopsis pus ilia

Ectocarpus sp.

Gracilaria verrucosa Catenella nipae Caloglossa leprieurii Caloglossa ogasewaraensis Caloglossa ad nata Polysiphonia variegata Polysiphonia macrocarpa Bostrychia tenella Bostrychia kelanensis Bostrychia moritziana (?) Bostrychia flagellifera

for export and biological metabolism . It is therefore of utmost importance that the remaining mangroves in the Brisbane River and its environs be nurtured, and that regrowth be encouraged. Dowling (1986b) has summarised the work that was conducted on re-vegetation of the Serpentine Creek Ana branch, and concluded that replanting can be done successfully.

and he suggested this was the normal period of highest planktonic productivity- this coincided with a standing crop of zooplankton which was much greater than in Moreton Bay. A blqom of another diatom Melosirag ranulata was observed in the upper estuary during February and March 1960. Atherton and Dyne (1977) have made a comprehensive intertida.l survey of Serpentine Creek and the species they found could be considered to also probably occur in the Brisbane River. They recorded 24 species (Table 1) most of the which were associated with mangrove pneumatophores, roots or trunks; with rotting logs; or on the mud surface. They stress that the factors controlling the presence or absence,

Algae There are almost no accounts of the algal flora of the Brisbane River. Bayly (1965) reported that from August to September during 1960 and 1963 there were enormous numbers of the diatom Coscinodiscus centra/is in the estuary, 138

Invertebrate and Intertidal Communities

distribution, abundance, and seasonal variation are highly numerous and complex. Species have different insolation requirements and for many these are only met within the complex environment of a mangrove community. Atherton and Dyne (1977) believe that destruction of mangroves would produce a situation where species diversity would plummet and a very few dominant species prevail. Eastwell (1987) and Moss (this volume) report on studies which show the Brisbane River to be considerably nutrient enriched with nitrogen and phosphorus washed into the river by rainfall runoff or discharged into the river at specific points mostly from sewage treatment plants and wastewater treatment of industrial effluents: In normal circumstances these nutrients would lead to massive algal blooms which would remove the oxygen from the water and cause severe problems, however the River is so turbid that light becomes the limiting factor preventing such occurrences. As Eastwell (1987) points out, a reduction in turbidity th rough cessation of dredging and other controls would likely require great improvements in the quality of some effluent discharges if algal problems are not to occur.

of dredging there must be concomittant improved standards of waste water and sewerage inputs, so as to avoid biological pollution caused by enrichment effects.

ACKNOWLEDGEMENTS I am grateful to my colleagues at the Queensland Museum for help in checking the accuracy of the species lists: in particular Thora Whitehead, Kevin Lamprell, and Darryl Potter for their help with molluscs; and Dr Patricia Mather for her assistance with ascidians. Dr Patricia Hutchings of the Australian Museum, Sydney, is particularly thanked for her help with the polychaete list, and where not indicated she is the authority for nomenclatural changes. The manuscript has greatly benefited by the careful reading and criticism of Bruce Campbell and John Short of the QM, and Dr Jack Greenwood of the Zoology Department, University of Queensland. Ms Julie Taylor is also thanked for her assistance in preparing the figures.

REFERENCES Atherton, G. and Dyne, G., 1977. Survey of the algal flora of the Serpentine Creek area. Brisbane Airport

Development Project Environmental Study. Vol. IV. Marine Study Factor Reports pp. 83- 109. (A.G.P.S. :

FUTURE DIRECTIONS Some of the broad ecological principles that control the distributions of animals and plants within the River are slowly being understood but much remains to be done. Long-term monitoring studies are vital before it will be possible to construct any sort of predictive model which can separate inherent instability from changes brought about by human agencies.

Canberra). Aziz, K.A . and Greenwood, J.G., 1981. A laboratory investigation of temperature and salinity tolerances of juvenile Metapenaeus bennettae Racek and Dall (Crustacea: Penaeidae). journal of Experimental Marine Biology and Ecology 54: 137 - 47. Aziz, K.A. and Greenwood, J.G., 1982. Response of juvenile Metapenaeus bennettae Racek and Dall, 1965 (Decapoda, Penaeidae) to sediments of different particle size. Crustaceana 43(2): 121 - 6. Barnes, J.L., 1969. The osmotic behaviour of a number of gra psoid crabs with respect to their different penetration of an estuarine system. journal of Experimental Biology 42: 535- 51. Bayly, I. A. E., 1964. A new species of Isias (Copepoda: Calanoida) from the Brisbane River estuary and a comparison of the Australasian centropagid genera. Australian journal of Marine and Freshwater Research 15: 239- 47. Bayly, I.A. E., 1965. Ecological studies on the planktonic Copepoda of the Brisbane River estuary with special reference to Gladioferens pectinatus (Brady) (Calanoida). Australian journal of Marine and Freshwater Research 16: 315- 50. Bayly, I. A. E., 1966. A new species and new records of Pseudodiaptomus (Copepoda: Calanoida) from the Brisbane River estuary, Queensland.

. We remain almost totally ignorant of th~ b1ology of particular species - even the most common. This is an area requiring urgent attention. h Replanting ma ngroves, and nurturing those t at have survived, is important for many reasons including: bank stabilization; estuarine broductivity; provision of faunal corridors for both aquatic and terrestrial organisms; and just ecause they look good. The hull shape and the speed of river vessels mus~ be controlled so as to minimise bank erosion caused by increasing river traffic. . Pollution must continue to be even more ngorously controlled. Along with the reduction 139

The Brisbane River

Proceedings of lire Royal Society of Queensland 78(5): 49-58. Boesch, D.F., 1977. A new look at the zonation of benthos along an estuarine gradient, In Coull, B.C. (Ed.) Ecology of the Marine Benthos (University of South Carolina Press: Columbia). Campbell, B.M., Wallace, C. and King, H., 1977a. The sublittoral macrobenthos of the proposed Brisbane Airport extension area. Brisbane Airport Development Project E1Jvironmental Study. Vol. IV. Marine Study Factor Reports. pp. 20- 48. (A.G.P.S. : Canberra). Campbell, B.M., Wal.lace, C. and King, H ., 1977b. Field study of marine littoral invertebrate macrofauna from the proposed Brisbane Airport extension area. Brisbane Airport Development Project

Environmental St udy. Vol. IV. Marine Study Factor Reports. pp. 49- 77. (A.G .P.S. : Canberra). Caswell, H ., 1978. Predator-mediated coexistence: a non-equilibrium model. American Naturalist 112: 127- 54. Coleman, N. and Cuff, W., 1980. The abundance, distribution and diversity of the molluscs of Western Port, Victoria, Australia. Malacologia 20: 35-62. Connell, J.H ., 1978. Diversity in tropical rainforests and coral reefs . Science 199: 1302 -1 0. Davie, P.J.F., 1986. Patterns in time and space in sub-tropical estuarine soft-bottom macro benthic communities in Serpentine Creek, Southeast Queensland. Unpubl. M.Sc. Thesis, University of Queensland, Brisbane. Driscoll, E.G. and Brandon, D. E., 1973. Mollusk sediment relationships in northwestern Buzzards Bay, Massachusetts, U.S.A. Ma/acologia 12: 13-46. Dowling, R.M., 1986a. The Mangrove Vegetation of Moreton Bay. Queensland Botany Bulletin 6: 45pp, 3 maps. Dowling, R.M. , 1986b. Summary report on mangrove revegetation trials undertaken by Botany Branch, Department of Primary Industries, Queensland, for Department of Housing and Construction. Eastwell, J., 1987. Water Quality, pp. 27-32, In , The

Brisbane River. A Strategy for our Future. Supporting TeciJIJica/lnformation. (Brisbane City Council: Brisbane). Fielder, D.R. and Greenwood, J.G ., 1983. The zoeal stages and megalopa of Bresedium brevi pes (De Man, 1899) (Crustacea : Decapoda : Grapsidae), reared in the lab ora tory . Journal of Plankton Rescarch5(4): 585- 98. Grassle, J.F. and Grassle, J.P., 1974. Opportunistic life histories and genetic systems in marine benthic polychaetes. journal of Marine Research 32: 253- 84. Gray, J.S., 1974. Animal-sediment relationships.

Occanograpl1y and Marine Biology Annual Revue 12: 223-62.

Greenwood, J. G., 1974. Calanoid copepods of Moreton Bay: a taxonomic and ecological account. Ph.D. Thesis, Zoology Department, University of Queensland. Greenwood, J. G., 1976. Calanoid copepods of Moreton Bay (Queensland). I. Families Calanidae, Eucalanidae, and Paracalanidae. Proceedings of the Royal Society Queensland 87: 1 - 28. Greenwood, J. G., 1977. Calanoid copepods of Moreton Bay (Queensland). II. Families Calocalanidae to Centropagidae. Proceedings of the Royal Society Queensland 88: 49- 67. Greenwood, J. G., 1978. Calanoid copepods of Moreton Bay (Queensland). III. Families Temoridae to Tortanidae, excluding Pontellidae. Proceedings of the Royal Society Queensland 89: 1 - 21. Greenwo·od, J. G., 1979. Calanoid copepods of Moreton Bay (Queensland). IV. Family Pontellidae. Proceedings of the Royal Society Queenslm1d 90: 93-111. Hailstone, T.S., 1972. Ecological studies of subtidal be nthic macrofauna at the mouth of the Brisbane River. Unpubl. Ph.D. Thesis, University of Queensland, Brisbane. Hailstone, T. S., 1976. Delimitation of subtidal macrobenthos associations at the mouth of the Brisbane River. Australian Journal of Marine and Freshwater Research 27: 217- 38. Heald, E.J., 1971. The production of organic detritus in a sou th Florida estuary. Sea Grant Technical Bulletin 6. University of Miami. 110 pp. Heger!, E.J ., 1975. The effects of flooding on Brisbane River mangroves. Operculum 4(2-3): 156- 57. Henry, K., Moore, G., Richardson, E. and Harris, 1., 1987. Ecological processes in the Brisbane River, pp. 33-41, In The Brisbane River. A Strategy for our Future. Supporli11g Technical Information . (Brisbane City Council: Brisbane). Hodge, D., 1963. The distribution and ecology of the mysids in the Brisbane River. University of Queenslm1d Papers, Department of Zoology 2(5): 91104. Hughes, E.H. and Sherr, E.B ., 1983. Subtidal food webs in a Georgia estuary:l\ 13 C analysis. Journal of Experimental Marine Biology and Ecology 67: 227- 42. Johnson, R.G., 1974. Particulate matter at the sediment-water interface in coastal environments. Journal of Mari11e Research 113: 313- 30. Kay, D.G . and Knights, R.D ., 1975. The macro-invertebrate fauna of the intertidal soft sediments of so utheast England. Journal of the

Marine Biological Association of the United Kingdom 55: 811- 32. Kennedy, G.R., 1975. Plankton of the Fitzroy River estuary, Queensland: (1) the upper reaches. Proceedings of the Royal Society of Queensland 86: 163-9.

Invertebrate and Intertidal Communities Miller, G.)., 1976. Export and production of organic detritus from north Queensland mangroves on a summer's day. Operculum 5(2): 56- 60. Mover ley, ).H., 1984. Population studies on mangrove crabs of Moreton Bay estuaries. Unpubl. M.Sc. Thesis, University of Queensland. Murdoch, W.W. and Oaten, A., 1975. Predation and population stability. Advances in Ecological Research 9:1 -131. Park, L.H., 1979. Macrobenthos in the vicinity of Luggage Point outfall, mouth of the Brisbane River. Unpubl. M.Sc. Thesis, University of Queensland, Brisbane. Pearson, T.H. and Rosenberg, R., 1978. Macrobenthos succession in relation to organic enrichment and pollution of the marine environment. Oceanography and Marine Biology Annual Revue 16: 229 -311. Poore, G.C.B. and Rainer, S., 1974. Distribution and abundance of soft-bottom molluscs in Port Phillip Bay, Victoria, Australia. Australian journal of Marin e and Freshwater Research 25: 371 - 411. Ruella, N.V., 1973. Burrowing, feeding, and spatial distribution of the school prawn Metapenaeus macleayi in the Hunter region (Australia). journal of Experimental Marine Biology and Ecology 13: 187204. Saenger, P., Hegerl, E.). and Davie, j.D.S., 1983. Global Status of Mangrove Systems. Commissions on Ecology. Paper 3. International Union for Conservation of Nature and Natural Resources. Saenger, P., Stephenson, W. and Moverly, )., 1980. The estuarine macobenthos of the Calliope River and Auckland Creek, Queensland. I. Analyses of 'pre-thermal' da ta. Memoirs of the Queensland Museu m 20: 143- 61. Snelling, B., 1959. The distribution of intertidal crabs in the Brisbane River. Australian journal of Marine and Freshwater Research 10: 67 - 83. Steele, ).G., 1976. The Brisbane River (Rigby: Adelaide), 63 pp. Stejskal, I.V., 1984. The spatial and temporal patterns of the macrofauna of a subtropical sand/ mud flat m southeast Queensland, Australia. Unpubl. Ph.D. Thesis, University of Queensland, Brisbane. St ejs · kal~ LV. and Chamberlain, D., 1984. The Impact of Atrport Construction on the Intertidal

Macrofauna of Bramble Bay, South East Queensland. A report for the Department of Housing and Construction. Stephenson, W., 1968. The effects of a flood upon salinities in the southern portion of Moreton Bay. Proceedings of the Royal Society of Queensland 80: 19 34. Stephenson, W. and Campbell, B.M., 1977. The macrobenthos of Serpentine Creek. Memoirs of the Queensland Museum 18: 75- 93. Stephenson, W., Cook, S.D. and Newlands, S.j., 1978. The macrobenthos of the Middlebanks area of Moreton Bay. Memoirs of the Queensland Museum 18: 185. 212. Stephenson, W., Cook, S.D. and Raphael, 1., 1977. Th~ effect of a major flood on the macrobenthos of · Bramble Bay, Queensland. Memoirs of the Queensland Museum 18: 95- 118. Stephenson, W., Raphael, Y.l. and Cook, S.D., 1976. The macrobenthos of Bramble Bay, Moreton Bay, Queensland. Memoirs of the Queensland Museum 17: 425.47. Stephenson, W., Williams, W.T. and Cook, S.D., 1974. The benthic fauna of soft bottoms, southern Moreton Bay. Memoirs of the Queensland Museum 17: 73. 123. Straughan, D., 1967. Intertidal fouling in the Brisbane River, Queensland. Proceedings of the Royal Society of Queensland 79(4): 25- 40. Vohra, F.C., 1965. Ecology of intertidal Zostera flats of Moreton Bay. Unpubl. Ph.D. Thesis, University of Queensland, Brisbane. Watson, C.).)., 1928. Notes on the growth of the Grey Mangrove (A vicennia) in the upper Brisbane River. The Queensland Naturalist july 1928: 83-4. Whittaker, R.H. and Levin, S.A., 1977. The role of mosaic phenomena in natural communities. Theoretical Population Biology 12: 117-39. Young, P.C. and Wadley, V.A., 1979. Distribution of shallow-water epibenthic macrofauna in Moreton Bay, Queensland, Australia. Marine Biology 52: 8397. . ZoBell, C.E., 1973. Microbial and environmental transitions in estuaries, In Stevenson, L.H. and Colwell, R.R. (Eds) Estuarine Microbial Ecology (BelleW. Baruch Coastal Research Institute: University of South Carolina Press), 536 pp.

141

APPENDIX 1: A List of Estuarine Invertebrate Species for the Brisbane River. This list is an update of that given by Hailstone et al. (1978) but is restricted to species occurring in the estuary only - freshwater species are listed separately by Arthington et al. elsewhere in this volume. Taxonomic changes have been made where necessary but names used in the original references have been included in parentheses below the current name. Taxonomic changes follow advice from experts acknowledged earlier. A'?' after the species name signifies that there is some doubt as to the accuracy of the identification. QM indicates that a specimen from the Brisbane River is lodged under that species name in the collection of the Queensland Museum. This list should not be considered to be exhaustive, and the author would appreciate knowing of any additions or corrections that may be found. The numbers to the right of each species name refer to the numbered references given in APPENDIX2. PHYLUM BRYOZOA

PHYLUM PROTOZOA CILIA TEA PERITRICHIDA

Vorticella sp. Zoothamnion sp.

88 88

......

"""

N

(38) 88 (38) 88 (38) (38) 88 (38) 88 88 (38)

SCYPHOZOA

Aurelia labiata Catostylus mosaicus Charybdea rastoni Cyanea capillata Pelagia noctiluca

(38) 38 (38) 38 (38)

ANTHOZOA

Edwardsia sp. Plexaura sp.

86 47,48

PHYLUM CTENOPHORA

Beroe ovata Pleurobrachia pile.us

(38) 38

88 88 88

PHYLUM PHORONIDA

Phoronopsis albomaculata

PHYLUM CNIDARIA HYDROZOA

Aequoria australis Bimeria francisciana Bougainvillea sp. Cordylophora locustria Diphyes chamissonis Eutima curva Halopteris diaphana Liriope tetraphylla Obelia longicyatha Obelia nodosa Octophialucium sp.

Bugula neritina Conopeum reticulium Cryptonita pallasiones

14

PHYLUM MOLLUSCA POL YPLACOPHORA

Delicatoplax sp. Rhyssoplax sp.

-l

47,48 47,48

:r

85

::l rt>

...

u;· 0""

GASTROPODA

Acteocina fsiformis Bedeva paivae (= B. hanleyz) Chemnitzia darnleyensis Cyllene sp. (= Radulphus sp.) Dendrodoris nigra Diala sp. Diodora sp. (= Eligidion sp.) Epitonium sp. (= Folaceiscala sp.) Epitonium helicornua (?) (= Limiscala helicornua) Epitonium jukesiana (?) (= Acutiscala faba) Etrema capillata Euchelus atratus Nassarius dorsatus Nassarius burchardi (= Parcanassa ellana)

rt> c:l 0>

:::0

86, 47, 48, 88 47,48 47,48 47,48 85 47,48 47,48 47,48 47, 48, 47,48 47,48 47,48 14, (20), 85, 47

:;:·

...rt>

Nassarius jonasii (= Parcanassa mange/aides) (= Reticunassa paupera) (= Reticunassa pilata) Natica sp. No toseila sp. Polinices conicus (= Conuber conicum) Polinices sordid us (= Conubersordida) Pupafumata (? = P.solidula or P.alveola) Pupa sp. Pyrazus eben in us Ringicula sp. Scutus unguis Stiliger boodlese Thalotia marginata (= Prothalotia marginata) Velacumantus australis

..... w """

BIVALVIA Acrosterigma recueanum (= Regozara oxygonum) A nadara trapezia Anomia sp. Antigona chemnitzi (= Proxichione materna) Area sp. Arthritica helmsi Barbatia sp. (mis-spelled as Abartia) Callanaitis sp. (? Placamen tiara) Chama sp. Chama fibula Circe nana (?) Corbiculina sp. Corbicu/ina baronialis Corbicu/ina prolongata Corbula sp. Crassostrea commercia/is Crassostrea sedea Cycladicama sp. Electromactra sp.

47,48 86 47,48 47,48 47,48 86 47,48 47, 48 85 47,48 88 47,48 47, 48

47,48 86,47,48 86 47,48 86 20,85 47,48 85 47,48 86 47,48 14 78 78 20 47,48, 88 47, 48 86 47,48

Epicodakia·sp. Exotica donaciformis (= Macoma donaciformis) Ge/oina coaxans Leptomya pura Limea fragilis (= Promantellum parafragile) Macomasp. Malleus a/bus Modiolus sp. Modiolus agripeta Modiolus ostentatus Modiolus pulex Musculus cumingianus Musculus sp. Mysellasp . Notospisula trigonella (= N. parva) Nucula astricta Ostrea bresia Paphia sp. Pinctada sugillata Placamen sydneyense (?) Plagiocardium setosum (= Regozara setosum) Pteria lata (= Austropteria saltata) Sanguinolaria donacioides Semelesp. Spondylus sp. Tapes dorsatus (= Tapes watlingz) Tapes subrugata (= Paphia subrugata) (= Paratapes scordalus) Tel/ina sp. Tel/ina (Homalina) deltoidalis Tel/ina solenel/a Tel/ina texturata (?) Teredo poculifer Theora lata Theora sp. Trichomya hirsuta Trisidos tortuosa

20 86 14 86 47, 48 14 47,48 20,85 47,48 86 88 47,48 47,48 20,85 14, 20, 85, 86 47,48 86 47,48 47,48 47,48 86 47,48 47,48 14, 20,85 47, 48 47,48 86 86 86 47,48 85 47,48 86 88 86 86 86, 47, 48 47, 48

5"'

< "'..., ;;r

...,0"

-~

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Venerupis sp. Venus sp. Volachlamys singaporina

47,48 86

(= V. cumingz) Wallucina jacksonensis Zemysia sp.

47,48 47,48 47,48

PHYLUM ANNELIDA POLYCHAETA

.....

""'""'

Amaeana tri/obata Amphitrite pachyderma (? = A. rubra) Armandia intermedia ( = A.lanceolata) Australonereis ehlersi Baranto/la lepte Boccardia sp. Ceratonereis erythraensis Chaetopterus variopedatus Cistena antipoda ( = Pectinaria antipoda) Cirriformia sp. Dasybranchus caducus (?) Diopatra sp. Diopatra dentata Eunice sp. Eupolyninia koorangia Eurythoeparvecarunculata (?) Ficopomatus enigmaticus ( = Mercerie/la enigmatica) Glycera sp. Glycera prashadi (?) Hadrochaeta aspeta Haploscoloplos sp. Hydroides norvegica lso/da pulche/la Lanassa sp. Laonice sp. Laonome sp. Leitoscoloplos norma/is (? = Scoloplos armiger) Leon nates jousseaumei Leon nates stephensoni Lepidonatus carinulatus

86 60 86 62 14,20,85 20, 85,80 14 14,20 14, 20, 85, 88, 80 86 86,47,48 86, 47,48 86 47,48 14 47,48 (60) 86 88 20 86 (60) 85 47,48 86, 47,48 47,48 85 14, 47,48 20,48 (64)

86 47, 48

.

Loimia medusa (? = L. ba Iilla or L. ingens) Lumbrinereis sp. Lumbrinereis latreilli Lysilla pacifica Mage/ana sp. Marphysa sp. Marphysa sanguinea Mesochaetopterus sp. Mesochaetopterus capensis (?) Mesochaetopterus sagittarius ( = M. minutus) Namalycastis abiuma Neanthes uncinu/a Nephtyssp . Nephtys australiensis Nereis sp. Nereis jacksoni (?) Nereis persica (?) Notomastus sp. Oenonesp. Onuphis sp. Ophelina gigantea Owenia fusiform is Perinereis calmani Petaloproctus terricola Phyllodoce malmgreni Pista sp. Pista Irina Pista trunca Polycirrus nephrosus Polycirrus octoseta Polydora sp. Polyophthalmus sp. Prionospio sp. Protula paliata Pseudostreblosoma serratum Serpula vermicularis Sigambra parva Spio sp. Sthenolepis yhleni ( = Lean ira yhlem) Terebella sp.

86 (60) (60) 47,48 14,85,86 (58) 14, 20, 85, 47, 48 20, 85,48 14,86 86 86 86 80 (80) 47,48 20,85 20 86 47,48 14 47,48 86 86 14,20,85 47,48 86 14 86 (57) (60) (58) (57), (58) 14, 20, (86) 47,48 14,47,48 47,48 (61) 88 14 14 86 47, 48

..., :r' "'0:: ::!.

"'0'"

"' :l

"':;;:1 :::·

.

11)

Terebellides stroemi Thelepus robustus Thelepus sp. Websterinereis puncta/a

86 (59) 47,48 (64)

OLIGOCHAET A Branchiura sowerbyi Limnodrilus hoffmeisteri Peloscolex sp.

14, 17 14, 17 14

PHYLUM CRUSTACEA CLADOCERA

Evadne tergestina Evadne nordmani Pen ilia avirostris

......

"'" til

COPEPODA CALANOIDA Acartia sp. Acartia baylyi) Acartia pacifica Acartia tranteri ( = Acartia clausi) Arcocalanus gibber Bestiola simi/is Calanopia sp. Calanopia australica Calanopia elliptica Candacia discaudata Canthocalanus pauper Centropages furcal us Centropages orsinii Clausocalanus furcatus Clausocalanus minor Eucalanus sp. Euchaeta concinna Gladioferens pectinal us Isias uncipes Labidocera sp. Labidocera rrzoretoni Mecynocera clausii Paracalanus aculeatus Paracalanus crassirostris Paracalanus parvus Pontellopsis tasmanensis Pseudodiaptomus aurivilli

(38) (38) (38)

16 38, 41, QM 38,41 41 38 38,39 38, 39 16 38,42 38,42 (38) (41) 16 16,38,40 (38) (38) (40) (38) (40) 16 (38) (40) 16,38,40 16,38,40 16 38,42 (38) (40) 38,39 38,39 38,39 (38) (42) 16, 38,40

Pseudodiaptomus colefaxi Pseudodiaptomus marin us Pseudodiaptomus mertoni Sulcanus conflict us Temora turbinata Tor/anus sp. Tor/anus barbatus CYCLOPOIDA Kelleria queenslandica Oithona brevicornis Oithona nana Oithona robusta Oithona (Dioithona) rigida Oncaea clevei Paramacrochiron australis THORACICA Acasta sp. Balanus amphitrite Balanus trigon us MYSIDACEA Gastrosaccus dakini Gastrosaccus queenslandensis Rhopalophthalmus brisbanensis

38,40 38,40 16, 38,40 16 16, 38,41 16 38,41 90 90 90 90 90 90 90

5"