The Queensland lungfish (Neoceratodus forsteri), also known as the Australian lungfish, Burnett salmon and barramunda, is a surviving member of the family Neoceratodontidae and order Ceratodontiformes. It is one of only six extant lungfish species in the world. Endemic to Australia, the Neoceratodontidae are an ancient family belonging to the class Sarcopterygii, or lobe-finned fishes.
Fossil records of this group date back 380 million years, around the time when the higher vertebrate classes were beginning to evolve. Fossils of lungfish almost identical to this species have been uncovered in northern New South Wales, indicating that Neoceratodus has remained virtually unchanged for well over 100 million years, making it a living fossil and one of the oldest living vertebrate genera on the planet.
It is one of six extant representatives of the ancient air-breathing Dipnoi (lungfishes) that flourished during the Devonian period (about 413–365 million years ago) and is the most primitive surviving member of this lineage. The five other freshwater lungfish species, four in Africa and one in South America, are very different morphologically from N. forsteri. The Queensland lungfish can live for several days out of the water, if it is kept moist, but will not survive total water depletion, unlike its African counterparts.
The small settlement of Ceratodus, Queensland, derives its name from that of the Queensland lungfish.
Distribution and habitat
The Queensland lungfish is native only to the Mary and Burnett River systems in south-eastern Queensland. It has been successfully distributed to other, more southerly rivers, including the Brisbane, Albert, Stanley, and Coomera Rivers, and the Enoggera Reservoir in the past century. The Queensland lungfish has also been introduced to the Pine, Caboolture, and Condamine Rivers, but current survival and breeding success are unknown. Formerly widespread, at one time at least seven species of lungfish were in Australia.
This species lives in slow-flowing rivers and still water (including reservoirs) that have some aquatic vegetation present on banks. It occurs over mud, sand, or gravel bottoms. Australian lungfish are commonly found in deep pools of depths between 3 and 10 m and live in small groups under submerged logs, in dense banks of aquatic macrophytes, or in underwater caves formed by the removal of substrate under tree roots on river banks. The lungfish is tolerant of cold, but prefers waters with temperatures between 15 and 25 °C.
The Queensland lungfish is incapable of surviving complete desiccation of its habitat, although it can live out of water for several days if the surface of its skin is constantly moist. Unlike the African species, Protopterus, it does not survive dry seasons by secreting a mucous cocoon and burying itself in the mud.
The Queensland lungfish is essentially a sedentary species, spending its life within a restricted area. Its home range rarely extends beyond a single pool or, occasionally, two adjacent pools. It does not follow a set migratory path, but may actively seek out suitable spawning habitats between July and December.
Queensland lungfish are olive-green to dull brown on the back, sides, tail, and fins, and pale yellow to orange on the underside. They have been described as having a reddish colouring on their sides which gets much brighter in the males during the breeding season. This colouration is the only distinguishing sexual characteristic of the lungfish. They have stout, elongated bodies and flattened heads with small eyes. The mouth is small and in a subterminal position. The lungfish can grow to a length of about 150 cm (4.9 ft), and a weight of 43 kg (95 lb). It is commonly found to be about 100 cm (3.3 ft) and 20 kg (44 lb) on average. Both sexes follow similar growth patterns, although the females grow to a slightly larger size. They are covered in slime when taken from the water.
The skeleton of the lungfish is partly bone, and partly cartilage. The vertebrae are pure cartilage, while the ribs are hollow tubes filled with a cartilaginous substance. The body of the lungfish is covered with large, bony scales. Ten rows occur on each side, grading to small scales on the fins. The scales are each embedded in their own pockets, and overlap extensively, such that vulnerable areas of the body are covered by a thickness of at least four scales. Two unusually large and thick interlocking scales cover the back of the head where the bony skull is thin. They have powerful diphycercal tails that are long and paddle-shaped. The pectoral fins are large, fleshy, and flipper-like. The pelvic fins are also fleshy and flipper-like and situated well back on the body. The dorsal fin commences in the middle of the back and is confluent with the caudal and anal fins.
The dentition of the lungfish is unusual: two incisors, restricted to the upper jaw, are flat, slightly bent, and denticulated on the hind margin. These are followed by dental plates on the upper and lower jaws.
Juveniles have different body proportions from mature adults. The head is rounder, the fins are smaller, and the trunk is more slender. The mouth is initially terminal, but shifts back as the fish grows. The dorsal fin typically reaches to the back of the head in young juveniles, and gradually moves caudally until it only extends to the mid-dorsal region in adults. They show a gradual change in body form as they develop, but no metamorphosis is externally detectable and no obvious point occurs at which they can be termed adult. As a juvenile, the lungfish is distinctly mottled with a base colour of gold or olive-brown. Patches of intense dark pigment will persist long after the mottling has disappeared. Young lungfish are capable of rapid colour change in response to light, but this ability is gradually lost as the pigment becomes denser.
The lungfish is reputed to be sluggish and inactive, but it is capable of rapid escape movements with the use of its strong tail. It is usually quiet and unresponsive by day, becoming more active in the late afternoon and evening.
A distinctive characteristic of the Queensland lungfish is the presence of a single dorsal lung, used to supplement the oxygen supply through the gills. During times of excessive activity, drought, or high temperatures (when water becomes deoxygenated), or when prevailing conditions inhibit normal functioning of the gills, the lungfish can rise to the surface and swallow air into its lung. More frequent air breathing is correlated with periods of greater activity at night when it uses the lung as a supplementary organ of respiration.
Unlike the South American and African lungfishes, the Australian species has gills on all the first four gill arches, while the fifth arch bears a hemibranch. It is also the only facultative air breather lungfish species, only breathing air when oxygen in the water is not sufficient to meet their needs. The lung is a single long sac situated above and extending the length of the body cavity, and is formed by a ventral outgrowth of the gut. Internally, the lung is divided into two distinct lobes that interconnect along its length, compartmentalized by the infolding of the walls. Each compartment is further divided to form a spongy alveolar region. Blood capillaries run through this region close enough to the air space in the lung to enable gas exchange. Lungfish breathe in using a buccal force-pump similar to that of amphibians. The contraction of smooth muscles in the walls of the lung results in exhalation.
The sound of the lungfish exhaling air at the surface prior to inhaling a fresh breath has been compared to that made by a small bellows. Young lungfish come to the surface to breathe air when they are about 25 mm long.
Reproduction and development
The Queensland lungfish spawns and completes its entire lifecycle in freshwater systems. The age of first breeding is estimated to be 17 years for males and 22 years for females. Males typically become mature at 738–790 mm and females at 814–854 mm. After an elaborate courtship, the lungfish spawn in pairs, depositing large adhesive eggs amongst aquatic plants. They spawn from August until November, before the spring rains, in flowing streams that are at least a metre deep.
Eggs are most abundant during September and October. The stimulus for spawning is believed to be day length. The lungfish is known to spawn both during the day and at night. The lungfish is selective in its choice of spawning sites. Eggs have been recorded on aquatic plants rooted in gravel and sand, slow- and fast-moving waters, in shade and in full sun, but never on aquatic plants covered with slimy algae, in stagnant water, or where loose debris was on the water’s surface.
Opposite of its South American and African relatives, the Australian lungfish does not make a nest or guard or care for its eggs. When spawning does take place, the pair of fish will lie on their sides or become entwined. They usually deposit their eggs singly, occasionally in pairs, but very rarely in clusters. The male lungfish fertilizes each egg as it emerges, and the eggs are deposited in dense aquatic vegetation. The newly laid egg is hemispherical, delicate, heavily yolked, and enclosed in a single vitelline and triple jelly envelope. The egg about 3 mm in diameter; with the jelly envelope, it has a total diameter of about 1 cm (0.39 in). The egg is sticky for a short while until silt and small aquatic organisms have covered it, but long enough for it to become attached to submerged vegetation. It is negatively buoyant and if it falls to the lake or river bed, it is unlikely to survive to hatching.
The female has a large ovary and the potential to lay many eggs, but in the wild only produces a few hundreds of eggs, at most, during her lifetime. In captivity, 200 to 600 eggs have been laid in a single event. The lungfish does not necessarily spawn every year. A good spawning season occurs usually once every five years, regardless of environmental conditions.
The eggs and young are similar to those of frogs, but the offspring differ from both frogs and other lungfishes by the absence of external gills during early development. Within the egg, head structures and pigmentation start to appear by day 17. They hatch after three to four weeks and resemble tadpoles. The young fish are slow-growing, reportedly reaching 27 mm (1.1 in) after 110 days, and about 60 mm (2.4 in)after 8 months. During the first week, it lies on its side, hiding in the weeds, and moving only when stimulated by touch. It will swim spontaneously, and often retreat back into the gelatinous envelope when disturbed. Newly hatched larvae develop a ciliary current over their skin and gill surfaces. This is believed to either provide respiratory exchange across the skin and gills without necessitating any movements of the jaw or brachial apparatus, or to keep the skin of the unprotected larvae free of debris, parasites, and predatory protozoans. Larvae are reported not to feed for two to three weeks while the yolk is still present. By the time the yolk is fully used, a spiral valve has developed in the intestine and the fish starts to feed. The young can grow about 2 inches per month under optimal conditions.
The Queensland lungfish has very complex courtship behavior made up of three distinct phases. The first is the searching phase, when the fish will range over a large area, possibly searching for potential spawning sites. A pair of fish will perform circling movements at the surface of the water close to beds of aquatic plants. They breathe air more frequently and more noisily than normal, possibly reflecting a greater physiological requirement for oxygen. Individual fish have been observed to breathe air at regular intervals of about 20 minutes, with air breathing accompanied by a distinct loud burp made in the air. The noisy breathing may be a form of a mating call. The lungfish seem to do their noisy breathing in concert, even responding to each other, but never in close vicinity of where the eggs are laid.
The next phase involves behavior, similar to “follow-the-leader”, during which one fish, the male, shows interest in the female and nudges her with his snout. Up to eight individuals may be involved in follow-the-leader behavior. The male lungfish may occasionally take a piece of aquatic plant into its mouth and wave it around. In the third phase, the fish dive together through aquatic vegetation, the male following the female and presumably shedding milt over the eggs.
Adults have a high survival rate and are long-lived (at least 20–25 years). A Queensland lungfish named "Granddad" at the Shedd Aquarium in Chicago was the oldest living fish in any Aquarium, and was already an adult when he was first placed on display in 1933; Granddad was estimated to be at least in his eighties, and possibly over one-hundred, at the time of his death on February 5, 2017.
The Queensland lungfish has an unusually large karyotype, very large chromosomes and cells, and a high nuclear DNA content relative to other vertebrates, but less than what is reported for other lungfishes. In spite of this, it displays low genetic diversity between populations from the Mary, Burnett, and Brisbane catchments. This low level of genetic variation could be attributed to population “bottlenecks” associated with periods of range contraction, probably during the Pleistocene, and in recent times during the periods of episodic or prolonged drought that are known to reduce some reaches of these river systems.
Diet and feeding habits
The Queensland lungfish is primarily nocturnal, and is essentially carnivorous. In captivity, it will feed on frogs, earthworms, pieces of meat, and pelleted food. In the wild, its prey includes frogs, tadpoles, fishes, a variety of invertebrates, and plant material. No quantitative dietary data are available, but anecdotal observations clearly indicate the diet of the lungfish changes with development. This is proven to be correlated with a change in dentition.
Lungfish larvae are bottom feeders. They eat microcrustaceans and small Tubifex worms, occasionally supplementing their diets with filamentous algae. Soft foods such as worms and plants are partially crushed with a few quick bites and then swallowed. In the adult lungfish, movement of the prey in and out of the mouth is accompanied by strong adduction of the jaws. This crushing mechanism is coupled with hydraulic transport of the food, achieved by movements of the hyoid apparatus, to position the prey within the oral cavity. The Queensland lungfish exhibits the most primitive version of these biomechanical feeding adaptations and behaviors.
Although the status of the Queensland lungfish is secure, it is a protected species under the Queensland Fish and Oyster Act of 1914 and capture in the wild is strictly prohibited. It was placed on the CITES list in 1977. The lungfish is currently protected from fishing, and collection for education or research purposes requires a permit in Queensland, under the Fisheries Act of 1994, and from the Commonwealth Government. It is included on the list of “vulnerable” species, as studies have failed to show it meets the criteria needed to be considered a threatened or endangered species.
Human activities currently threaten the Queensland lungfish, particularly water development. It is potentially at risk in much of its core distribution in the Burnett and Mary Rivers, as 26% of these river systems are presently impounded by weirs and dams. Barriers to movement and altered flow regimens downstream of dams for irrigation purposes could lead to the disruption of existing population structure and cause even more loss of genetic variation.
Queensland lungfish can be very fast-growing, yet with a delayed first breeding age. For a long-lived species with naturally low mortality rates, successful spawning and juvenile recruitment is not essential every year and may only occur irregularly in medium to long cycles, even in natural environments. The length of these cycles could easily mask the potentially deleterious impacts on recruitment for many years. Additionally, large adults could remain common for decades and give no indication of a declining population in the longer term.
The Mozambique mouth brooder, or tilapia, has been declared a noxious and threatening alien species to the lungfish in Queensland.
Proposed 2006 damming projects on both the Mary and Burnett rivers threaten the habitat of the remaining lungfish. The dams would change the flow of the rivers, eliminating the slow, shallow areas the fish need for spawning. Scientists worldwide have become involved in saving the habitat for these lungfish, citing their evolutionary importance.
- ^"Part 7- Vertebrates". Collection of genus-group names in a systematic arrangement. Retrieved 30 June 2016.
- ^Haaramo, Mikko (2007). "Ceratodiformes – recent lungfishes". Mikko's Phylogeny Archive. Retrieved 3 July 2016.
- ^Froese, R.; Pauly, D. (2017). "Neoceratodontidae". FishBase version (02/2017). Retrieved 18 May 2017.
- ^ abcdeLake, John S. Australian Freshwater Fishes. Nelson Field Guides. Melbourne: Thomas Nelson Australia Pty. Ltd., 1978. p. 12.
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- ^ abcdefghijklmnopAllen, G.R., S.H. Midgley, M. Allen. Field Guide to the Freshwater Fishes of Australia. Eds. Jan Knight, Wendy Bulgin. Perth, W.A.: Western Australia Museum, 2002. pp. 54–55.
- ^ abcFrentiu, F.D., J.R. Ovenden, and R. Street (2001). "Australian lungfish (Neoceratodus forsteri: Dipnoi) have low genetic variation at allozyme and mitochondrial DNA loci: a conservation alert?". Conservation Genetics. 2: 63–67. doi:10.1023/A:1011576116472.
- ^ abcdefgWhitley, G.P. (1960). Ed. Jack Pollard, ed. G.P. Whitley’s Handbook of Australian Fishes. Victoria: Wilke and Company Ltd. p. 334.
- ^Martin F. Gomon & Dianne J. Bray, 2011, Australian Lungfish, Neoceratodus forsteri, in Fishes of Australia, accessed 07 Oct 2014, http://www.fishesofaustralia.net.au/home/species/1988
- ^Kemp, A. (1995). "Threatened Fishes of the World: Neoceratodus forsteri (Krefft, 1870) (Neoceratodontidae)". Environmental Biology of Fishes. 43: 310. doi:10.1007/bf00005863.
- ^ abcdefghijklmPusey, Brad, Mark Kennard, and Angela Arthington (2004). Freshwater Fishes of North-eastern Australia. Nathan, QLD: CSIRO Publishing. pp. 49–59.
- ^Joss, J. (2002). "Australian Lungfish, Neoceratodus forsteri". Fishes of Sahul. 16: 836–844.
- ^ abBrooks, S.G. & P.K. Kind. "Ecology and demography of the Queensland lungfish (Neoceratodus forsteri) in the Burnett River, Queensland with reference to the impacts of Walla Weir and future water infrastructure development". Queensland Department of Primary Industries, Brisbane, Report No. QO02004 (2002).
- ^ abcdeKemp, A. (1986). "The biology of the Australian lungfish, Neoceratodus forsteri (Krefft 1870)". Journal of Morphology. 190: 181–198. doi:10.1002/jmor.1051900413.
- ^ abKrefft, G. (1870). "Description of a gigantic amphibian allied to the genus Lepidosiren from the Wide-Bay district, Queensland". Proceedings of the Zoological Society. 16: 221–224.
- ^ abKemp, A. "The embryological development of the Queensland lungfish, Neoceratodus forsteri (Krefft)". Memoirs of the Queensland Museum. 20. (1982): 553–597.
- ^Kemp, A. (Autumn 1990). "A relic from the past – The Australian lungfish". Wildlife Australia: 10–11.
- ^Grigg, G.C. (1965). "Studies on the Queensland lungfish, Neoceratodus forsteri (Krefft) III. Aerial respiration in relation to habits". Australian Journal of Zoology. 13: 413–421. doi:10.1071/zo9650413.
- ^"Freshwater Fish Distribution".
- ^Grigg, G.C. (1965). "Studies on the Queensland lungfish, Neoceratodus forsteri (Krefft)". Australian Journal of Zoology. 13: 243–253. doi:10.1071/zo9650243.
- ^ abcdKemp, A. "Spawning of the Australian lungfish, Neoceratodus forsteri (Krefft) in the Brisbane River and in Enoggera Reservoir, Queensland". Memoirs of the Queensland Museum. 21. (1984): 391–399.
- ^Merrick, J.R.; Schmida, G.E. (1984). Australian Freshwater Fishes Biology and Management. Sydney: Griffin Press. pp. 46–51. ISBN 0-9591908-0-5.
- ^Whiting, H.P. & Q. Bone (1980). "Ciliary cells in the epidermis of the larval Australian dipnoan, Neoceratodus". Zoological Journal of the Linnean Society. 68: 125–137. doi:10.1111/j.1096-3642.1980.tb01922.x.
- ^ abGrigg, G.C. (1965). "Spawning behaviour in the Queensland lungfish, Neoceratodus forsteri". Australian Natural History. 15: 75.
- ^Johnson, Steve (2017-02-06). "Australian lungfish 'Granddad,' the oldest zoo animal in Chicago, dies". Chicago Tribune. Chicago. Retrieved 2016-02-07.
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- ^Neoceratodus forsteri — Australian Lungfish, Queensland Lungfish. Biodiversity: Species Profile and Threats Database. Department of the Environment, Australian Government.
For information to assist regulatory considerations, refer to Policy Statements and Guidelines, the Conservation Advice, the Listing Advice and/or the Recovery Plan.
|EPBC Act Listing Status||Listed as Vulnerable|
|Approved Conservation Advice|| Department of the Environment (2014). Approved Conservation Advice for Neoceratodus forsteri (Australian lungfish). Canberra: Department of the Environment. Available from: http://www.environment.gov.au/biodiversity/threatened/species/pubs/67620-conservation-advice.pdf. In effect under the EPBC Act from 11-Apr-2014. |
|Listing Advice|| Threatened Species Scientific Committee (2003). Commonwealth Listing Advice on Neoceratodus forsteri (Australian Lungfish). Available from: http://www.environment.gov.au/biodiversity/threatened/species/n-forsteri.html. In effect under the EPBC Act from 06-Aug-2003. |
|Recovery Plan Decision|| Recovery Plan required, included on the Commenced List (1/11/2009). |
|Adopted/Made Recovery Plans||There is no adopted or made Recovery Plan for this species|
|Adopted/Made Threat Abatement Plans||No Threat Abatement Plan has been identified as being relevant for this species|
|Scientific name||Neoceratodus forsteri |
|Species author||(Krefft, 1870)|
The distribution shown is generalised from the Departments Species of National Environmental Significance dataset. This is an indicative distribution map of the present distribution of the species based on best available knowledge. Some species information is withheld in line with sensitive species polices. See map caveat for more information.
The current conservation status of the Australian Lungfish, Neoceratodus forsteri, under Australian and State/Territory Government legislation is as follows:
National: Listed as Vulnerable under the Environment Protection and Biodiversity Conservation Act 1999.
Queensland: In Queensland, the Australian Lungfish is not listed as threatened under the state's Nature Conservation Act 1992, however, the taking of Australian Lungfish has been prohibited since it was declared a protected species under the Queensland Fish and Oyster Act 1914. The species is currently protected from fishing, and collection requires a permit under the Fisheries Act 1994.
Scientific name: Neoceratodus forsteri
Common name: Australian Lungfish
Other common name: Queensland Lungfish
The Australian Lungfish (herein referred to as the 'Lungfish') is a long, heavy-bodied freshwater fish. The species has five pairs of gills and its fins resemble flippers. Adult Lungfish can weigh up to 48 kg and grow to around 2 m (Environment Australia 2003ab; Grigg 1975), but commonly reach 1.3 m. In the Burnett River, the mean length was 906 ± 199 mm, and the mean weight 7573 ± 4563 g (Brooks & Kind 2002). Adult Lungfish are dark brown or olive brown on the back and pink on the belly (Kemp 2016, pers. comm.), with some whitish colour on the belly and underside of the head. Lungfish in Lake Samsonvale are less pink and more yellow on the belly, probably owing to the fact that the environment is so depauperate that the lungfish are unhealthy (Kemp 2016, pers. comm.).
They have large, overlapping scales and a small mouth with large, crushing teeth on the palate and lower jaw (Allen 1989a; Grigg 1975). Juveniles are dark olive, brown or yellow with a mottled pattern above and a dull pink belly (Kemp 1995; Kind 2002). The species is able to breathe aquatically using its gills, and aerially using its single lung. It usually uses its gills, but surfaces to breathe when it is active and requires more oxygen. For example, it breathes air more often at night while foraging, when swimming in floodwaters, and when spawning (Grigg 1964, 1975; Kemp 1984; Merrick & Schmida 1984).
The Australian Lungfish is endemic to Australia and restricted to south-eastern Queensland (Wager 1993). The species' natural distribution is the Mary, Burnett and Brisbane River systems and (possibly) the Pine River system (Kemp 2014). The species has been translocated to many other locations and translocated populations persist in the Coomera, Condamine, Albert and Logan Rivers (Kemp 2014).
The Lungfish occurs in a number of water body types, ranging from relatively undisturbed streams to highly altered environments, such as Lake Samsonvale and Lake Wivenhoe (Brooks & Kind 2002; Johnson 2001; Kemp 2014). However, heavily altered water bodies are unlikely to be suitable for the species in the long-term, evidenced by the local extinction of the species from the Enoggera Reservior, following translocation to the area (Kemp 2014).
The distribution of the Lungfish within the Burnett River is from Ben Anderson Barrage to a point around 10 km upstream of the John Golby Weir (around 335 km from the mouth of the river). It is also known to occur in tributaries of the Burnett River, including (Brooks 1995; Brooks & Kind 2002):
- Three Moon Creek to around Mulgildie;
- 20 km up the Boyne River from its junction with the Burnett River, as far as Boondooma Dam, including the waterhole directly below the dam wall;
- Barambah Creek, but not in the upper reaches above Barambah Gorge. Brooks (1995) suggested that they may occur above the gorge to at least the Joe Sippel Weir, but Books & Kind (2002) found no evidence of the species above Barambah Gorge;
- The lower reaches of the Auburn River below the falls/gorge.
Kind (2002) caught and observed the Lungfish on the Burnett River at sites including Ben Anderson Barrage, Bingera Weir, Lee's Waterhole, Drinan's Crossing, Walla Guaging Weir, Booyal Crossing, Mingo Gorge, Grey's Waterhole, Booyal Crossing (immatures), and Jones Weir. Grigg (1975) observed Lungfish spawning at Bon Accord Crossing over Barambah Creek, in 1964.
The distribution of the Lungfish within the Mary River is from the Mary River Barrage (59.3 km from the mouth of the river) to the town of Conondale (220 km from the mouth of the river). It also occurs in large tributaries of the Mary River, including:
- Yabba Creek, which flows into the Mary River between Kenilworth & Gympie;
- Tinana Creek, Coondoo Creek, Wide Bay Creek, Obi Obi Creek, and Munna Creek (Brooks and Kind, as cited in Kind 2002; Simpson 1994).
Kind (2002) caught and observed the Lungfish in the Mary River at sites including Kenilworth Homestead, Kenilworth Quarry, Moy Pocket, Traveston Crossing, and Widgee Crossing at Gympie. He also caught them at Imbil Bridge and near Imbil Weir (a subadult) on Yabba Creek, Bungawatta on Tinana Creek, and in Condoo Creek. O'Connor (1897) translocated 78 Lungfish from the Mary River to a number of locations. Details of the release were provided by Welsby (1905), O'Connor (1902) and Kemp (2016, pers. comm.):
- the North Pine River (eight fish taken to the site, three survived for release);
- a lagoon near the Albert River (five fish taken to the site, four survived for release);
- a dam near Cressbrook on the upper Brisbane River (five fish were released; it is unclear whether the fish dispersed to the Brisbane River, however, dead lungfish were observed in a fish kill event in the Brisbane River two years later);
- Enoggera Reservoir (18 fish, at least some fish survived but, following clearing of water hyacinth from the reservoir in 1973, recruitment at the site would have ceased);
- the Condamine River (21 fish);
- the Coomera River (16 fish taken to the site, 14 fish survived for relase and were observed at the release point for many months); and
- the Botanic Gardens in Brisbane (two fish).
The most successful captive breeding facility is in Queensland and operated by Jindalee Internation (Kemp 2016, pers. comm.), however, captive reared fish cannot be released into the wild because the environment is now so poor, and they will have difficulty in finding adequate food, or even know how to find it (Kemp 2016, pers. comm.).
The Lungfish was comprehensively surveyed in the Burnett and Mary River systems between 1997 and 2002, but the status and distribution of the translocated populations is based on anecdotal information and sporadic records (Brooks & Kind 2002, Kind 2002).
After floods of 2009-2013 and after the destruction of lungfish that have fallen over the walls of water impoundments, the population of the species is likely to be much less than 10 000 individuals, even in their natural environments (Kemp 2016, pers. comm.). Sunwater and SEQwater may not be recording the number of fish killed by falling over the walls of water impoundments (Kemp 2016, pers. comm.).
There are three known populations of the Australian Lungfish, occurring naturally in the Burnett, Mary and Brisbane River systems (Kemp 1986; Lissone 2003).
There are 'moderate numbers' in the North Pine River, and Lungfish are often caught and reported by anglers downstream at Young's Crossing after water flows over the top of the dam. It is uncertain whether this population is natural or a result of translocation (Kemp 2014). In the 1980's and 1990's, Lungfish were regularly found at the base of the Enoggera dam wall after floods (however, this population is considered to be locally extinct) (Kemp 2014).
Translocated populations of Lungfish are still present in the Condamine River, Logan River and Coomera River (DPI information, cited by Kemp 2016, pers. comm.). They probably still occur in Lake Manchester, but are probably extinct in Gold Creek Reservoir, where they have not been seen since 1975 (Kemp 2016, pers. comm.).
The Australian Lungfish requires still or slow-flowing, shallow, vegetated pools with clear or turbid water in which to spawn and feed (Allen 1989a; Merrick & Schmida 1984). The species is restricted to areas of permanent water (Brooks & Kind 2002) and cannot live in saline waters or migrate through sea water (Arthington 2009). Emergent or submerged vegetation are essential for successful deposition of eggs and for providing refuges for juveniles (Kemp 2014).
Burnett River and tributaries
The Burnett River extends around 420 km from its source to the sea. Brooks & Kind (2002) reported that Lungfish were 'often' found in freshwater reaches of the Burnett River up to 310 km inland from the river mouth, and less frequently found upstream of this point. At sites between 50 and 350 km upstream, the abundance of Lungfish was negatively correlated with the distance from the mouth of the river (fewer lungfish further away from the sea), except for the relatively small population in Ben Anderson Barrage pond near the river mouth. Based on the catch rate per unit effort, Brooks and Kind (2002) concluded that Lungfish were less likely found in impoundments than in flowing stretches of the Burnett River. In the lower Burnett River, Ben Anderson Barrage pool is the closest water storage to the sea (26 km away). It usually consists of 41 km of impounded slow-flowing water surrounded by steep banks (Kind 2002). Bingera Weir is 42.5 km from the sea, so it is usually inundated by Ben Anderson Barrage pond. When it is not submerged, it impounds 19 km of the river above Ben Anderson Barrage. There is a section of the river upstream of Ben Anderson Barrage pond consisting of 7 km of flowing water with shallow runs up to 32 m wide, and riffles around 4m wide, up to Ned Churchward Weir (formerly called Walla Weir). Ned Churchward Weir (74.5 km from the river mouth) impounds water 34.5 km upstream to a point 109 km from the mouth of the Burnett River. Within the Ben Anderson Barrage pond, there were more Lungfish in the headwater section, where there were shallow areas and more complex habitat (Berghuis & Broadfoot 2004a; Brooks & Kind 2002). The central Burnett River area contains three irrigation storages that are impassable to lungfish in normal flow conditions (John Goleby Weir, Jones Weir and Claude Wharton Weir). Unimpounded sections of the river include many shallow pools and runs with a width of less than 10m, containing submerged logs and branches (Brooks & Kind 2002). Rocks are more common in the Burnett River than the Mary River, and are used as shelter by the Lungfish (Kind 2002). Between Claude Wharton Weir (202.4 km from the river mouth) and the Burnett River Dam (131 km from the river mouth, with an impoundment extending to a point 176 km from the river mouth) is a section of flowing river, riffles, shallow runs and pools within the Goodnight Scrub, which approaches the natural state of the river (Kind 2002; Sinclair Knight Mertz 2001).
Mary River and tributaries
The Mary River flows for around 250 km from its source in the Conondale Range to the sea. In the Mary River, Kind (2002) found that shallow pools, runs and riffles were common, and there were few waterfalls. Areas where Lungfish were studied consisted of 67% pools, 21% runs, 11% riffles and glides, and 1% backwaters. The average pool depth is 1.8m, the average run depth 1.2 m, and the average stream width less than 30 m. In pools where adult Lungfish were studied, macrophytes (water plants that are large enough to be seen with the unaided eye) covered 37 to 55% of the habitat (mean 47%), around 40% was open water, and the rest was stream bank vegetation (including grass growing into the water), rocks, eroded banks and woody debris. The substrate was typically sandy. Macrophytes were most abundant at depths of 0.9 to 2.1m. Yabba Creek is regulated by water releases from Borumba Dam (31.1 km from the junction with the main channel of the Mary River) and Imbil Weir (10.9 km from the junction). The average depth of Yabba Creek is 1.7 m and there is abundant overhanging vegetation, submerged macrophytes and rocky cover. Lungfish in the Mary River catchment were also less abundant in the impounded waters than further upstream. Tinana Creek is 140 km long. It flows into the Mary River downstream of the Mary River Barrage. The lower section is regulated by Tallegalla Weir (37.5 km from the junction), Teddington Weir (15.8 km from junction), and Tinana Barrage (1.5 km from the junction). The average depth of Tinana Creek is 1.7 m, and it is generally less than 10 m wide. It comprises a series of pools, runs and riffles, and there is abundant overhanging vegetation and submerged woody debris, but little macrophyte vegetation in the water (Kind 2002). Coondoo Creek is 70 km long, and flows into Tinana Creek 75 km from its junction with the main channel of the Mary River. Adult Lungfish in the Mary River are associated with overhanging riparian (riverside) vegetation, woody debris in the water, and dense macrophyte beds. They shelter in complex, shaded habitat. They most prefer habitat with overhanging vegetation and macrophytes in relation to their availability, and often use habitat with woody debris, although adults are not as reliant on submerged branches as some other Australian freshwater fish. The species avoids open water, and very seldom uses rocky habitat and eroded banks, which are uncommon in the Mary River. Lungfish are usually inactive when located in overhanging riperian vegetation and woody debris, especially during the day. They occur in water that is 1.86 m ± 0.61m deep on average, and in deeper water in winter than in summer (2.07 ± 1m m versus 1.81 ± 0.82 m). They use shallower water in the spawning season than at other times (1.77 ± 0.95 m), and slightly deeper water during the day than at night. Adult lungfish use water depths of 2 to 3 metres. Two macrophytes, Vallisneria (Vallisneria gigantea) and Hydrilla (Hydrilla vertillata) were used by all Lungfish radio-tracked in the Mary River, but Kind (2002) did not find any adult Lungfish using the macrophytes Baby Tears (Bacopa monniera), Nitella (Nitella sp.), water milfoils (Myriophyllum spp.), or hornwort (Ceratophyllum demersum). In relation to their availability, Lungfish preferred some species of macrophytes over others; they preferentially used two species that form very dense submerged banks and occur in a range of water depths; dense water weed (E. densa) and Hydrilla. They also prefer multi-species mixtures of floating and submerged macrophytes, and two species with floating leaves; Water Primrose (Ludwigia peploides) and Waterlilies (Nymphoides sp.) (Kind 2002).
The upper reaches of the Brisbane River consist of two water storages, the Somerset and Wivenhoe dams (Kemp 1987). Below these dams, the river forms wide, slow-flowing permanent reaches with a maximum depth of around 4.5 m, and occasional shallow riffles. The riverbed consists mainly of sand, gravel and rocks, and the water is clear to turbid (after rain), with a nearly neutral pH (8). Red Bottle-brush (Callistemon saligna) and She-oak (Casuarina sp.) trees overhang the banks, and their root masses grow in the water, contributing to spawning and shelter habitat for Lungfish. Aquatic macrophytes are abundant (Kemp 1987).
Enoggera Reservoir is a permanent body of water around 60 ha in area and 18 m deep. It is filled by springs in the D'Aguillar Ranges and drains into Enoggera Creek, which drains into the Brisbane River estuary. The bottom of the dam is mainly mud, covered with detritus from water lilies and Eucalyptus leaves. The water is clear, but stained brown by tannins from decomposing leaves, and slightly acidic (pH 5). It has steeply sloping sides with Para grass (Brachiaria mutica) growing out from the edges to a water depth of around 2 m, forming a dense floating mat along with some aquatic macrophytes. Macrophytes are generally absent from the reservoir except at the edges. Water hyacinth (Eichornia crassipes) has been controlled in the Reservoir since 1974 (Kemp 1987). Kemp (1987) noted that Enoggera Reservoir was a poor environment for the Lungfish. In 2016, "a few surviving lungfish had been reported in Enoggera Creek below the reservoir (Kemp 2016, pers. comm.).
Habitat requirements for juvenile and immature Lungfish
With one exception (a juvenile caught in 1 m of clear water, Kind et al. 2005), juvenile Lungfish have always been found in dense cover such as beds of macrophytes in water 500 mm deep or less. The chosen habitat of juveniles has been well known since the work of Semon (1899) and Bancroft (1924, 1928). Observations of captive-bred juveniles showed that they prefer macrophytes over other shelters, and use the densest macrophyte cover available, especially if it is shaded. It appears that wild juveniles remain in the same type of cover as that used for spawning for many months or years after hatching (Kind 2002). One was found sheltering amongst river macrophytes (Semon 1899, as cited in Brooks & Kind 2002), and at least seven were recovered from the roots of Water Hyacinth plants removed from Enoggera Reservoir (Longman 1928). Brooks & Kind (2002) caught 21 juveniles and saw nine others at Drinan's crossing on the Burnett River, in macrophytes near where they found 293 hatched Lungfish eggs. They also caught two juveniles at Gayndah on the Burnett River, where they found three hatched eggs. All of these juveniles were in dense macrophytes in water less than 500 mm deep. Kind et al (2005) caught two juvenile Lungfish in dense aquatic vegetation and debris in 300 mm of water in the upper reaches of Jones Weir. Brooks & Kind (2002) caught two juvenile lungfish at Traveston Crossing on the Mary River amongst Hydrilla and Nymphaena plants in 500 mm deep water, and one at Twin Bridges on the Brisbane River amongst Vallisneria gigantea and woody debris. All juveniles smaller than 100 mm were caught in habitat suitable for spawning. The largest juveniles were found in a macrophyte species that are rarely used for spawning (H. verticillata), in the largest macrophytes, and in the deepest water. Larger juveniles inhabited larger macrophytes at greater depths. Immature Lungfish between 300 and 700 mm long were found most often associated with overhanging streamside vegetation, in areas of dense woody debris, undercut banks and dense macrophytes. The average water depth used by immature Lungfish during the day was around 1.5 m (Kind 2002).
Brooks & Kind (2002) reported that Lungfish gathered in deeper water within the Burnett River in autumn, possibly in preparation for the dry season. Unlike the African and South American Lungfish, the Australian Lungfish does not use stagnant mud as a refuge during dry seasons or droughts. It cannot live in stagnant water or bury itself in mud. Surveys conducted before major impoundments were built showed that even during prolonged drought, parts of the Mary and Burnett Rivers inhabited by Lungfish still flowed slowly and pools were well-oxygenated (Grigg 1965).
The Australian Lungfish first breeds at around 15 years of age in males and 20 years in females (Kind 2002). One female Lungfish spawning for the first time was 810 mm long, and one was 820 mm long (Brooks & Kind 2002).
Hatchlings and juveniles are vulnerable to predatory insect larvae, shrimps, fish and wood ducks (Bancroft 1928; Illidge 1894; Kemp 1987). Adults have few or no natural predators (Kind 2002) and the species is considered long-lived, with estimates of life spans from 50 to 100 years (Arthington 2009). Lungfish are difficult to age, and the otoliths, even in large fish, are not easy to extract. They have only one incremental line (Retzius 1881). Methods of ageing using carbon dating of the scales (Kelly et al. 2010) are unlikely to give accurate ages, because the tissues extracted from the scale and used for measurements come from a part of the scale that is laid down over many years (Kemp 2015; Kemp et al. 2015). Modifications suggested by Fallon et al. (2015) are not an improvement, as they still use many layers of the scale tissue, and they do not have an accurate understanding of the carbon residence time of the areas from which the scales were obtained (Kemp et al. 2015).
Timing of spawning
The Australian Lungfish spawns at night between August and December, with peak activity in late October. In the Burnett River, spawning occurs between mid August and early November (Brooks & Kind 2002). In the Brisbane River, eggs have been found between mid August and December (Kemp 1984). The breeding season in the Enoggera reservoir was usually shorter than that in the Mary, Burnett or Brisbane River (Kemp 1987). Lungfish delay or skip breeding if their spawning habitat is disturbed. Following severe winter flooding on the Brisbane River, spawning was reduced and did not commence until October in one year, and there was no evidence of spawning after a second flood the next year (Kemp 1993). Spawning is triggered by increasing daylength, and is not related to rainfall or water chemistry (Kemp 1984).
Conditions required for spawning
Lungfish pairs spawn amongst aquatic macrophytes. In the Brisbane river, eggs have also been recorded on submerged roots of Red Bottle-brush (Callistemon saligna) (Kemp 1984). Although water depth, substrate, macrophyte species, macrophyte density and macrophyte height may be important for suitability for Lungfish spawning in the Burnett River (Brooks & Kind 2002), it has been suggested that they are not particular about where they spawn (Kemp 2016, pers. comm.) and, although unusual, spawning has been observed over bare substrates (Roberts et al. 2014).
Lungfish will spawn in both still and flowing water, but their spawning behaviour is different in the two environments. Brooks (1995) found that in areas of still water, egg densities were highest at water depths of 50 to 100 mm, and fewer eggs were found in deeper water. In still water, Lungfish prefer to spawn where the river bed is sandy, in water that is very shallow (less than 100mm deep). In flowing water, which contains more oxygen, Lungfish eggs may be found in depths of more than a metre. They usually use sites between 200 and 600 mm deep. In flowing water, they will spawn over a variety of river bed substrates. Lungfish spawned in shallow river sections between impoundments. No spawning was observed in Jones Weir or Ned Churchward Weir pools between 1997 and 2000, but fish in reproductive condition were caught once near an island in Ben Anderson Barrage Pool. Spawning Lungfish choose clear water and avoid turbid (muddy) water.
Lungfish eggs die before completing development at temperatures below 10° or above 30° C (Kemp 1981). A clutch consists of between 50 and 100 eggs (Kemp 1987). The eggs are around 3 mm in diameter, with an adhesive jelly coating around 1 cm wide. They sink when first laid (Kemp 1987). Eggs adhere to the surfaces of submerged macrophytes, and occur singly, in pairs, or very rarely in clumps (Brooks 1995). If suitable macrophyte species are not available, Lungfish will not spawn (Kemp 1984). Suitable macrophytes grow in a dense mass in shallow water in a variety of substrates from gravel to mud, and contain a complex community of algae, protozoa, worms, small molluscs and crustaceans (Brooks 1995; Kemp 1993). Suitable species include Vallisneria gigantea (the most commonly used), Hydrilla verticillata, Nitella sp., Potamageton perfoliatis, Milfoil (Myriophylum sp.), Najas tenuifolia, Water Couch Grass (Paspalum distichum) (a terrestrial grass that grows in a dense mat from the water's edge across the surface), Baby Tears (Bacopa monniera), Curly Pondweed (Potamageton crispus), Chara sp., Carex sp., Slender Knotweed (Persicaria decipiens), Ludwigia peploides, Cladophera sp., various species of filamentous algae, and the introduced Water Hyacinth (Eichhornia crassipes), which floats on the surface (Bancroft 1911, Brooks 1995; Grigg 1965a; Kemp 1993; Roughley 1951). Brooks & Kind (2002) also collected Lungfish eggs from partially submerged Para grass (Urochloa mutica) in the Brisbane River and the Mary River.
Spawning always occurs in river sections where the cover of macrophyte algae is very high (usually exceeding 90%), and Lungfish choose the densest macrophytes available in which to lay their eggs. The highest densities of eggs were on macrophytes that were 160 mm tall, and fewer eggs occurred on taller water plants. Lungfish use macrophyte species with complex branching or leaf whorls rather than strap-like leaves, perhaps because eggs that detach from the surface of these are less likely to fall to the bottom (Brooks & Kind 2002).
In 1999 and 2000, Lungfish spawning was observed on the lower Boyne River, upper Burnett River at the junction with the Auburn River, at several riffle and glide sections between the Ned Churchward Weir wall and the Isis pumping station, and an area of Ben Anderson Barrage. Spawning on the Burnett River was recorded within a 7 km riffle and glide section of the river between Ned Chruchward Weir and the impoundment of Ben Anderson Barrage, where there was a steady flow of water, and abundant macrophytes. Spawning habitat on the lower Boyne River and the area 7 km downstream of Ned Churchward Weir was improved by releases of water from the weir and Boondooma Dam during 1999 and 2000. Large numbers of Lungfish congregated there during spawning, e.g. more than 500 individuals were counted within the 7 km on one day in October 2000. This area is the only remaining spawning habitat in the lower 80 km of the main channel of the Burnett River (Kind 2002). Lungfish also spawned above the influence of Ned Churchward Weir in the Goodnight Scrub area (which was then a flowing reach of the river between 80 and 200 km from the river mouth). Suitable macrophyte density for Lungfish spawning was rare in the Mary river between 1999 and 2002, as a result of a record high flow event that scoured the banks in 1999 (Brooks & Kind 2002).
Recruitment levels, and conditions required for successful reproduction
Lungfish eggs hatch after 30 days (Kemp 1981, 1987). Egg survival is best in shallow water that has a dense cover of macrophytes. Brooks & Kind (2002) found that the proportion of live Lungfish eggs increased with the density of macrophytes at the spawning site, up to a density of 40% (all macrophyte densities above 40% are equally suitable for egg survival). A lower proportion of live eggs occurred on taller macrophytes and in deeper water. The highest egg survival is between 200 and 800 mm water depth. Kemp (1994) reported that around 5% of eggs laid in the wild are unfertilised.
Young larvae look like tadpoles and are initially poor swimmers, resting on the bottom of the water on their sides until they have digested the yolk (Kemp 1995). Hatchlings and juveniles need refuges in water plants (Kemp 2016, pers. comm.). Newly hatched Lungfish retreat from light (J. Joss, as cited in Brooks & Kind 2002). They first breathe air when they are 27 mm long and around 110 days old, and resemble adult fish when they are six or seven months old (Kemp 1981). They begin to feed four to six weeks after hatching (Kemp 1995), and reach 6 cm after eight months and 12 cm after two years (Allen 1989). Growth curves of wild Lungfish on the Burnett River show that they reach around 40 cm at five years of age (Brooks & Kind 2002). Juvenile Lungfish smaller than 300 mm long have been collected from the Burnett and Mary Rivers and Mt Crosby Weir, 62 km downstream from Wivenhoe Dam in the Brisbane River (Longman 1928, as cited in Kind 2002; Johnson 2001; Kemp 1987; Semon 1899). One was found in the Boyne river (a tributary of the Burnett River) in 1892, and 20 in Enoggera Reservoir between 1928 and 1932. Six juvenile Lungfish were caught in the strainer of the water treatment works at Mt Crosby Weir in the Brisbane River in 1961, and eight in 1982 (Grigg 1965c; Kemp 1987). 'Numerous' juveniles were caught in the Burnett and Mary Rivers in 1981 and 1982. Brooks & Kind (2002) caught two juveniles at Traveston Crossing on the Mary River in 1998, and one at Twin Bridges in the Brisbane River in 1999. They also caught 23 juveniles in 1997 in the Burnett River. The size distribution of Lungfish between 1997 and 2001 indicated that recruitment of juvenile fish into this population was extremely poor, and there was a lack of fish under ten years old (fish up to 500 mm long were virtually absent from samples). This lack of successful recruitment was probably due to poor breeding conditions for several years (Brooks & Kind 2002). Three juveniles less than 200 mm long were caught in the upper reaches of Jones Weir in the Burnett River in 2004, and must have hatched in 2003 or 2004 (Brooks et al. 2005). Data on the frequency of different size classes of 2770 Lungfish in the Burnett River were collected by the Queensland Department of Primary Industries. They confirmed that recruitment for at least the five years prior to 2003 was lower than previously, and that there was a long period in the recent past where few juveniles survived. Kemp (1987) and Brooks & Kind (2002) suggested that Lungfish recruitment may be successful only in a small minority of years, when breeding conditions are good throughout much of the river (i.e. there are abundant macrophyte algae in shallow water, and high concentrations of microcrustaceans and invertebrates for juveniles to eat (Kemp 1977).
Kemp (1987) stated that successful breeding seems to have occurred at intervals of around twenty years since before 1900. Juvenile lungfish are found only sporadically, despite the fact that effective sampling techniques have been used at times when none were caught. For example, Bancroft (1911) failed to find any juveniles in the Burnett River despite exhaustive liming, dynamiting and dredging of spawning habitat (Kemp 1987). None were caught in the Mt Crosby water treatment works filters between 1961 and 1982, and none were caught in Enoggera Reservoir by electrofishing in the early 1980s (Kemp 1987). Brooks & Kind (2002) failed to find juvenile Lungfish in the Burnett River between 1998 and 2001, despite intensive and widespread sampling of spawning habitat using frame nets and electrofishing, which were previously effective.There is no recent evidence of successful Lungfish breeding in impoundments outside the Burnett River, Mary River, and the Brisbane River downstream of Wivenhoe Dam. All records of Lungfish in Lake Samsonvale and Lake Wivenhoe are of mature adults. In some years Water Hyacinth dies back during the winter (Kemp 1984), so it would not be available as juvenile habitat in those years. When Lungfish bred in the Enoggera Reservoir, it may have provided better spawning habitat than do impoundments on the Burnett River, because it supplies drinking water, so the water level changes slowly compared with the rapid changes in flow associated with irrigation supplies for sugar cane and other seasonal agriculture on the Burnett River (Brooks & Kind 2002).
Due to the variation in spawning conditions and food resources, successful recruitment of lungfish is not likely to occur every year. This can affect age classes in populations, with a result that seemingly secure populations may decline catastrophically once older individuals die, and with lower or no replacement breeding stock (Walker 2008).
Adult Lungfish are suctorial feeders, and ingest masses of material that they do not digest (Spencer 1892), particularly plant matter and sand. In fact they rely on molluscs and other small animals for food. They are low level benthic carnivores in practice (Kemp 2016, pers. comm.). Hatchling and juvenile lungfish feed on small invertebrates (Kemp 1981, 1996) and are active predators, at least when young.
Lungfish forage mainly at night. In the Mary River, adults forage predominantly where there are macrophyte beds in shallow water (less than 2 m deep). Juveniles ambush their prey in the structurally complex habitat of aquatic plants, catching and holding them with sharp, cone-shaped teeth (Kind 2002). Lungfish detect the vibrations of prey, and are capable of perceiving the weak electric fields generated by animals. Experiments have shown that they can accurately locate buried prey using electroreception (Watt et al. 1999). Juvenile lungfish in captivity do not behave as they would in the wild. They are aggressive to other small Lungfish when they are living in a confined space, which often has limited food (Kemp 2016, pers. comm.).
In rivers with natural flows of water, the Australian Lungfish is largely sedentary. In flowing (unimpounded) sections of the Burnett River and the Mary River, adults usually move around one or two pools at night and return each day to a certain habitat feature, such as a submerged log, rock or patch of macrophytes in one particular pool, where they rest. Individuals are routinely found resting in the same daytime retreat over many consecutive months. Movements exceeding one kilometre are rare, and only four of the 20 Lungfish radio-tracked were ever found more than 5 km from their original site (Kind 2002). Berghuis & Broadfoot (2004a) captured 42 of the same individuals that were originally tagged by Brooks & Kind (2002), and most were still in the same areas as they had been five years previously. In 2002, Lungfish movements in the (then) flowing section of the Burnett River (the Goodnight Scrub) were independent of temperature and water flow (Brooks & Kind). During a record flood on the Mary River, Lungfish remained in their home ranges and retreated to the shallow banks. After the river broke its banks, several left the channel to shelter amongst flooded vegetation, on the downstream side of large trees, or in a deep vehicle track perpendicular to the flow direction (Kind 2002). Despite their large size, they are capable of traversing very shallow riffle zones (e.g. 120 mm deep) into other pools to find food and spawning habitat (Kind 2002; Brooks & Kind 2002). Immature Lungfish (n = 3) moved up to 200 m from their daytime retreats while foraging at night, but remained close to cover (Kind 2002).
The movements of Lungfish are restricted by natural and man-made barriers. Natural barriers include waterfalls and gorges (e.g. on the Auburn River and Barambah Creek, which are tributaries of the Burnett River), and ephemeral (temporary) water (e.g. the Nogo and Perry Rivers, tributaries of the Burnett River which are devoid of Lungfish). Man-made barriers include dams, weirs, barrages and culverts. For example, Lungfish apparently inhabited the Boyne River above the Boondooma Dam before the dam was constructed. They no longer occur there, although they are abundant downstream of the dam to the junction of the Burnett River. Claude Wharton Weir at Gayndah and Jones Weir at Mundubbera block the passage of Lungfish, and the culverts at Booyal Crossing apparently also form barriers to Lungfish movements, except during particularly high flows (Brooks & Kind 2002). In the Burnett river, Lungfish in established impoundments moved much more than those in flowing sections, and Lungfish movements became much more variable at reproductive maturity. Adult Lungfish radio-tagged downstream of the Ned Churchward Weir (in the established Ben Anderson Barrage pond and the shallow zone upstream of this) moved regularly between the weir and the barrage. For example, one individual traversed the 48 km from the weir to the barrage at least four times between 1998 and 2001. There is an annual cycle of movement to and from spawning grounds in this section of the Burnett River, when fish abandon their usual home ranges. Lungfish spawned in the series of shallow riffles and runs immediately below Ned Churchward Weir wall. Some individual females moved upstream to spawn here, then returned to their usual home ranges in the impoundment more than once in a season (Kind 2002). Increases in water discharge from the weir caused Lungfish to move downstream, and downstream movements were significantly related to the river flow rate. These fish were probably not passively washed down the river, but took advantage of the flow to return more easily to their non-breeding season home ranges. Two Lungfish that had been tagged upstream of the weir apparently swam over the top of the wall during flooding in February 2003, and moved 25 and 36 km downstream. They were detected by Berghuis & Broadfoot (2004a) attempting to return upstream in April and May the same year, but they did not successfully pass through the fishlock. Some Lungfish tagged in the Jones Weir moved upstream during the spawning season into shallow sections of the Boyne and Auburn Rivers, which are tributaries of the Burnett River (Brooks & Kind 2002).
During the non-spawning season, fish in impoundments showed strong site fidelity to a restricted area, similar to fish in flowing parts of the river. Lungfish in the Mary River (which contained no substantial impoundments) did not undertake spawning migrations (Kind 2002). Lungfish that were caught and radio-tagged within the Ned Churchward Weir in the Burnett River made only local movements, similar to fish in the unimpounded section of the river. When the weir was filled, they moved into the flooded areas. Several fish then became stranded in a pool when the water level subsequently dropped, and had to be moved overland to the river before the pool dried. With the exception of one individual that moved out of the impoundment and travelled 11 km upstream, they did not move away from the inundated area to find a spawning site. During the spawning season, large groups of Lungfish were seen milling around the water surface over previous spawning sites that were now flooded by the weir. Brooks & Kind (2002) suggested that searching for spawning habitat further afield is a behaviour learned by Lungfish in established impoundments over time.
The mean linear range of Lungfish in the Burnett River was 11 714 ± 16 044 m (n = 29 fish). There was no difference between the sexes. Lungfish in the lower reaches of the river (in the Ben Anderson Barrage pond) downstream of Ned Churchward Weir had larger home ranges and their ranges were proportional to body length, unlike those within the impoundment and in flowing sections of the river (Brooks and Kind 2002). The mean linear range of Lungfish in the Mary river was 4770 m, including two outlying individual males, with ranges of 26.1 km and 30.5 km. The average linear range of the Lungfish in the flowing section of the Burnett River was 1758 m. Some individuals in flowing river habitat had linear ranges of 300 to 500 m. The linear ranges of Lungfish in the Mary River and the Goodnight Scrub (then flowing) area of the Burnett River were not significantly different, but those in the impounded sections of the Burnett River were larger. In adults, home ranges of both sexes overlap, and daily ranges are smaller in winter. Three immature fish had linear ranges of 900 to 2400 m. Unlike adults, juvenile Lungfish appear to be territorial, and aggressive to one another. During the day, larger juveniles in captivity pushed and bit smaller ones to exclude them from preferred shelter sites (Kind 2002).
Adult Lungfish are largely nocturnal. They forage in the early morning, late afternoon, and night, and their activity peak during foraging is around midnight. They are inactive between 10am and 3pm. They forage for shorter periods during winter than in summer. It is sometimes possible to observe them (e.g. during spawning), and to hear the gurgling sound of air rushing into the lung as they surface to breathe (Grigg 1975; Kind 2002), but systematic surveys to confirm their presence or absence at a site involve capturing fish (Brooks & Kind 2002; Kind 2002). Lungfish eggs are distinctive and can be surveyed using systematic searches of macrophytes in shallow water (Kind 2002). Juvenile Lungfish are difficult to detect between the period just after hatching, and when they reach 300 mm long, because they have a cryptic appearance and behaviour, remaining in dense cover and shade when young. They have been found amongst the roots of water hyacinth plants (Kemp 1984), amongst other macrophytes suitable for Lungfish spawning (Brooks & Kind 2002; Kind 2002), and in water treatment works (McDougall 2001, as cited in Johnson 2001).
The Burnett River basin is one of the most developed river catchments in Queensland. It contains 26 water storages, including seven on the main channel of the Burnett River. In 2002, the Mary River system contained 11 water storages, and most barriers to fish movement were on tributaries of the river. Water infrastructure on the Mary River itself is limited to the Gympie Control Weir, a small weir 179.5 km from the river mouth that is regularly inundated by flows, and the Mary River tidal barrage at Tiaro, 59 km upstream from the river mouth (Berghuis & Broadfoot 2004a; Kind 2002). Dams can change the water quality downstream, because they often release poorly oxygenated water, increase sediment and cause bank erosion through flow regime changes (especially in the first five years after construction) (Walker 1985).
Flooding as a result of dams and weirs removes the breeding habitat of Lungfish (shallow water containing dense macrophytes), and dam walls block the movement of adult lungfish to the remaining breeding sites (Brooks & Kind 2002; Environment Australia 2003ab). The shallow, heavily vegetated areas required as spawning habitat are patchy, so Lungfish need to move between different parts of the river if their usual home range lacks adequate spawning habitat in a given year (Brooks & Kind 2002). The cumulative effect of multiple weirs and dams progressively removes Lungfish breeding habitat and disrupts movements. The closer impoundments are placed together, the more severe this effect is (Boardman 1996a; Brooks 2002; Brooks & Kind 2002). In 2002, 41% of the known range of the Lungfish within the main channel of the Burnett River (128km) had been inundated by impoundments (Brooks & Kind 2002). When the Burnett River (Paradise) dam has filled, this will increase to 55% (173 km). In 2003, it was estimated that there was a 26 per cent loss or reduction in the amount of breeding and nursery habitat for the Lungfish overall (Environment Australia 2003ab). This has now increased, as a further 14 of the remaining 35 riffle sections of the lower Burnett River were impounded by the Burnett River dam, so that now only 26 km of the formerly flowing reach in the Goodnight Scrub remains (Sinclair Knight Mertz 2001).
As a result of breeding habitat reductions, the population is likely to undergo a substantial decline within the next three generations (Environment Australia 2003ab). Impoundments are likely to have negative effects on eggs and juvenile Lungfish, because macrophyte beds of the density required rarely have time to establish, there is a risk of stranding, and the diversity of macroinvertebrates (eaten by juvenile Lungfish) is lower than in the unimpounded river (AWT 2001; Brooks & Kind 2002). Brooks & Kind (2002) reported that the extensive macrophyte beds that Lungfish used for breeding disappeared after Walla Weir on the Burnett River (now called Ned Churchward Weir) flooded the area. Flooding washed away macrophyte beds, and turbid (muddy) flows smothered recovering macrophytes. Construction of impoundments results in initial loss of spawning habitat as shallow areas upstream are inundated. For example, when Walla Weir was created, macrophytes died and decomposed within six weeks. After the water level upstream became constant, new growth of water plants established after a further six to nine weeks, but these plants died when the water level fluctuated again (Duivenvoorden 1998). Macrophytes that have been scoured or destroyed by flooding may take years to re-establish and grow into dense beds. Brooks & Kind (2002) found that extensive macrophyte beds in the Burnett River that were used by Lungfish during the 1995 breeding season and were then destroyed by high flows, had not fully re-established by the 2000 breeding season. At Walla Weir, spawning habitat was also exposed and adult Lungfish were stranded by changes in water level below the weir (Berghuis et al. 2000). Brooks (1995) recorded Lungfish spawning within the Ben Anderson Barrage on the Burnett River in 1995 when water levels were very low throughout the catchment, but eggs were killed by subsequent changes in water level. Juveniles rely on dense macrophyte beds in very shallow water for many months or years after hatching, so they are vulnerable to stranding, loss of shelter and loss of food from fluctuations in water level. Brooks & Kind (2002) and Kind (2002) noted that spawning habitat on the lower Boyne River and the area 7 km downstream of Ned Churchward Weir on the Burnett River was improved by releases of water from Boondooma Dam and Walla Weir during 1999 and 2000. However, they doubted that impoundments can be operated to provide long term habitat for recruitment of Lungfish.
Although Lungfish sometimes form spawning aggregations in unregulated sections of the Burnett River (Johnson 2001), Brooks & Kind (2002) considered that as the amount of suitable spawning habitat decreases and Lungfish in impoundments are unable to move to spawning sites outside, crowding effects are likely to increase beyond natural levels. These might include increased predation of eggs and juveniles, increased competition between adults for spawning sites, and between juveniles for habitat and food, and mortality from disease. Hatchlings and eggs are vulnerable to bacterial infections, both in the wild and when raised in captivity (Kemp 1994).
A recent study of differences in the development of larvae taken from impoundments and relatively unaltered stream reaches has drawn a link between embryonic abnormality and altered watercourses (Kemp 2014). Fertilised eggs were removed from various spawning sites and raised in a controlled environment, to eliminate the potential effects of varying substrate or macrophyte quality. Abnormalities observed from eggs taken from water impoundments ranged from those which caused mortality of the embryo, those which resulted in abormal body sturcture or development and those which resulted in abnormal sense organs or hatchlings with a lower body condition (Kemp 2014). Water contamination was not considered to be a likely causal factor; however, age or lack of genetic condition in adults are potential contributors (Kemp 2014). Although abnormalities in lungfish embryos and hatchlngs have been recorded sporadically and lungfish hatcheries report up to 70% mortality in some years, occurrence of lethal abormalities at the scale observed in impounded water bodies is not likely to be normal or sustainable for the species (Kemp 2014).
Lungfish can be injured or killed when they pass over the top of dam walls during high water flows. They swim over the top of Ben Anderson Barrage at high tide, and into the estuary. If not rescued, these fish are stranded and will be killed by the salt water or by sharks, because they are too large to use the vertical slot fishway in the barrage (Berghuis et al. 2000; Brooks & Kind 2002; Kind 2002; Stuart & Berghuis 2002). Stuart & Berghuis (2002) collected 44 lungfish from the bottom of the barrage between December 1997 and April 1999. A similar process occurs at Mary River tidal Barrage (Berghuis 2001) and Kemp (1995) also reported that Lungfish were stranded in unsuitable habitat when they descended over the Enoggera Reservoir wall. Johnson (2001) reported that around 60 individuals were found dead or injured at the base of the spillway of Lake Samsonvale dam wall after a rapid release of water there. He suggested that many Lungfish are likely to be injured or killed passing over the dam and weir walls in the Burnett River during severe floods, and very large floods could potentially kill hundreds of individuals. Berghuis & Broadfoot (2004b) also emphasized that most passage of fish downstream at Ned Churchward weir on the Burnett River happens over the spillway during high flows. Large fish such as Lungfish are most likely to be injured or killed by abrasion and shearing against the spillway face as they are washed over the wall. Some fish also pass over the wall during low flows. Conditions during low flows are particularly dangerous because water and fish drop vertically before hitting the slope of the spillway face.
- Recruitment in Lungfish is naturally erratic. In the early 1990s, a substantial proportion of the Lungfish in the Burnett River were small (200 to 600 mm long, Brooks 1995; Brooks & Kind 2002). There was spawning, but no detectable recruitment of Lungfish between 1995 and 1999. Juveniles were found in the Burnett River in 1997, but these were lost during a high flow event that scoured their macrophyte habitat from the river, and no further juveniles were found by Brooks & Kind (2002) until 1999, despite intense sampling using the same methods. During the many months that juvenile lungfish remain in beds of macrophytes after hatching, they are vulnerable to displacement and predation during flow events. The distribution of size classes of fish in the Burnett River indicates that there have also been periods of several years without recruitment in the past. Reducing the amount of suitable spawning habitat is likely to increase the number of years in which recruitment fails due to natural events such as droughts and floods. Because adult Lungfish are very long-lived, population decline due to increasingly frequent recruitment failure will not be detectable for several decades (TSSC 2003). As the species shows strong affiliation to spawning sites, there is also the threat of low recruitment from impacts on known spawning sites (Catalyst 2006).
- Lungfish are caught by recreational anglers targeting other fish (Kemp 1995; Kind 2002). Of 52 radio-tagged fish studied by Kind (2002) in the Burnett and Mary Rivers, four may have been caught by anglers. The aerial of one transmitter had apparently been cut by a tool, but the fish was released and continued to live in the river. One transmitter that had been implanted in a fish was recovered away from the water, and two were recovered next to campgrounds. Johnson (2001) noted that Lungfish are often caught accidentally by anglers in the North Pine River.
- Exotic or translocated native fishes, such as the exotic Tilapia (family Cichlidae), are believed to prey on lungfish eggs and young and compete with adults for breeding habitat (Environment Australia 2003).
- Widespread clearing of riverbank vegetation has reduced habitat for Lungfish, because they often shelter or spawn amongst parts of terrestrial plants that overhang the water, particularly in the Mary River. Clearing also increases the damage caused by floods and removes cover that Lungfish use to prevent themselves from being displaced by fast-flowing floodwater (Johnson 1997, as cited in Kind 2002; Kind 2002).
- It has been suggested that the number of water snails, clams, filamentous algae and macrophytes have declined in the Brisbane River in recent years, due to the flushing effects of water releases, and that cyanobacteria now dominate the microflora because of high nutrient levels from fertiliser runoff (TAPSF n.d.). These changes have reduced food and breeding habitat for Lungfish. Runoff from cotton cultivation, citrus crops and cane production is also a potential problem in the Burnett River (Tucker et al. 1999). Water quality in the Mary River in the vicinity of Gympie is affected by discharge from a sewerage treatment plant, which increases nutrients and decreases oxygen in the water. Meatworks effluent, pesticide and herbicide runoff also affect the river bank and water quality in sections of the Mary River (EPA 2001).
Lungfish in the Burnett River, the Mary River and the Brisbane River have low genetic diversity, probably because of past population bottlenecks combined with the long generation time and low juvenile survival. This could exacerbate the effects of other threats, so it has been noted that further hindering gene flow between impoundments is likely to have substantial affects on the species (Frentiu et al. 2001).
A research project is testing artificial environments for eggs and young lungfish in the Brisbane River. Long strands of acrylic wool tied in bundles and attached to Callistemon roots or floating platforms are rapidly colonised with protozoa and small clams (potential food for juvenile Lungfish). Despite low levels of spawning recently in the Brisbane River, these 'mops' were used by spawning Lungfish, and healthy eggs were recovered from them (TAPSF n.d.). Artificial spawning sites will only work if placed in an area where lungfish normally spawn, and they require maintenance (Kemp 2016, pers. comm.). They are no substitute for a natural environment (Kemp 2016, pers. comm.).
Johnson (2001) recommended that in order to reduce harm from existing impoundments:
- Water releases from weirs and dams should be gradual and carefully timed, and macrophyte abundance monitored.
- Passage opportunities should be constructed at causeways, bridges, and other crossings. Kemp (2016, pers. comm.) notes that fishways allow lungfish and other species to move freely in an impounded river, however, design improvements are required.
- At Ben Anderson Barrage, the vertical slot fishway might be improved by replacing the current 1: 15.8 gradient with a more gradual slope, such as 1: 18 to 20, and increasing the slot width. Until an appropriate fishway exists, staff should check for Lungfish in the estuary after major flows and manually relocate them above the barrage. The departure angle of the wall could be adjusted and energy-dissipating measures put in place at the base of the wall to minimise Lungfish injury during spillover at the barrage.
- Some Lungfish found spawning in very crowded sections of the Burnett River could be translocated to suitable habitat that is currently blocked by impoundments, such as Boondooma dam and Bjelke Petersen dam.
Devices to move fish upstream of dam walls and weirs (fishways)
The design of fishways depends on the height of the wall. Vertical-slot fishways are used for medium-sized weirs up to 6 m high. They consist of a concrete channel extending from the top of the weir (headwater) to the base of the weir (tailwater). Concrete walls or baffles are then inserted along the length of the channel to slow the flow of water. Vertical-slot fishways usually have a low slope (1:20 gradient) with slow water velocities (1 metre per second) and low turbulence (Larinier et al. 2002; QDPI 2004). The Vertical-slot fishway at Ben Anderson Barrage has a 1: 15.8 gradient and the slots are 150 mm wide, which is too narrow for adult Lungfish to swim up (Stuart & Berghuis 2002). However, adult Lungfish up to around 800 mm long have been observed moving through a 200 mm wide vertical slot fishway at the Tinana Creek Barrage (on a tributary of the Mary River) (Berghuis 2001).
Fishlocks are computerised devices to transfer fish over dam walls which are usually 8 to 10 m tall. The fishlock consists of a downstream channel at the tailwater (base of the weir) connected to a vertical chamber which extends to the top of the weir. There is an exit chamber at the top of the vertical chamber. Fish attempting to find a route upstream search for areas of high flow. Water is discharged from the fishlock to attract fish into the downstream channel and then into a cage at the bottom of the vertical chamber. The gate shuts behind them, the vertical chamber is filled with water, and the cage floats to the top, where another attraction flow entices fish to swim into the exit chamber (Larinier et al. 2002; QDPI 2004).
Lungfish use fishlocks infrequently. There is a 15 m high fishlock on Ned Churchward Weir in the Burnett River, and Berghuis et al. (2000) recorded that 12 lungfish used this. However, Brooks & Kind (2002) observed that radio-tagged Lungfish at the same weir approached but did not enter the fishlock, and large numbers of Lungfish gathered downstream prior to spawning seasons, but did not pass through. Berghuis & Broadfoot (2004a) used remote monitoring of Lungfish that had passive transponder tags to determine if they entered this fishlock. In 2003, they tagged 1285 Lungfish upstream and downstream of the weir. 41 Lungfish were detected approaching the fishlock in the downstream channel, and seven used the fishlock. These fish were 385 to 1170 mm long. Most Lungfish approaching downstream swam in and out of the channel several times before leaving the area. There was no difference in the size range of Lungfish in the downstream channel and the size range tagged in the River generally. More Lungfish congregated at the downstream channel (attempting to swim upstream from the direction of Ben Anderson Barrage) than the upstream one.
Berghuis & Broadfoot (2004a) suggested several reasons why so few Lungfish used the fishlock. Fish attempting to find a route upstream could previously swim anywhere across the width of the river. At a natural barrier, fish moving upstream were able to scan the constantly-flowing open water below the barrier to find the best place to pass. They are now expected to cross a turbulent channel that may be several hundred metres wide in high flows, in order to find a single fishlock entrance. The pattern of flow beneath the spillway and the fishlock entrance differ because the two are separated by a wall. The attraction flows at the fishlock are discontinuous, and insignificant compared to the rate of water flowing over the spillway during high flows, when large numbers of Lungfish congregate, probably attracted away from the fishlock by the higher water flow over the spillway. Additionally, fish entering the fishlock must wait in the chamber for up to 40 minutes before the gate closes behind them, and must exit at the top before the chamber empties. Several Lungfish entered the chamber at the bottom but then left again before the gate closed. Some fish took 17 minutes to pass through the fishlock, but one took more than five hours, and one took two days to leave. A potential problem for small individuals using fishlocks is that large, predatory fish sometimes exploit them as shelter in which to ambush prey (Berghuis & Broadfoot 2004b).
Berghuis & Broadfoot (2004a, 2004b) suggested the following improvements to the design of Ned Churchward Weir fishlock, and other fishways, to enable Lungfish passage:
- Water flows to attract Lungfish to the downstream fishlock entrance should be increased and varied to compete with the changing spillway flow. The time that water flows to attract Lungfish to the upstream entrance should be increased, as there is no attraction flow for 68% of each cycle. Constant attraction flows upstream and downstream could be accomplished by drawing water from the upstream fishlock channel, together with the present system of using surface water from the variable offtake outlet works at the base of the wall.
- A better entrance design.
- Mechanical restraint of fish in the chamber without preventing downstream movement. This restraining device could be removed during the appropriate part of each cycle to allow fish to leave.
- Mechanical devices to encourage fish to leave at the top of the fishlock.
- Management of overcrowding (e.g. multiple fishlocks), as Lungfish and other species migrate upstream and downstream at the same times of year.
- Extending the times that the fishlock operates by starting it as soon as possible after a peak flood flow has passed. It currently shuts down during floods to avoid damage.
- Because Lungfish need to be able to move safely both upstream and downstream, but the fishlock is designed for upstream passage, a dedicated downstream fishway should be considered. For example, a notch in the wall with a deep stilling basin below it to dissipate energy as fish pass over the weir, or a low gradient bypass that regulates flow velocity downstream.
Monitoring in the vicinity of Burnett River Dam
Six Lungfish sampling sites have been established between 119 and 201 km from the mouth of the Burnett River, in order to monitor the effect of the newly-constructed Burnett Dam. These are at Figtree, Cherelly Orchard, Kalliwa Hut, Mingo Gorge, Gray's Waterhole, and Claude Wharton Weir. The sites at Kalliwa Hut and Mingo Gorge are within the impoundment. Seven spawning survey sites have also been established. These are at Fig Tree, River Road Crossing, Kalliwa campground, Mingo crossing, Yenda Benyenda, downstream of Gayndah and in the Gayndah township. Kalliwa campground, Mingo crossing, and Yenda Benyenda are within the impoundment. Kind et al. (2005) collected baseline information on water quality, habitat, condition and size of Lungfish and the distribution of Lungfish eggs in August 2004 and January 2005, as part of a ten-year monitoring program. Lungfish were often found in the area of the Burnett River Dam, and occurred at similar density to areas downstream. There was no spawning at the two Gayndah sites.
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Department of the Environment and Heritage (DEH) (2006pk). Neoceratodus forsteri in Species Profile and Threats (SPRAT) database. Unpublished species profile. Canberra, ACT: DEH. Available from: http://www.environment.gov.au/cgi-bin/sprat/public/publicspecies.pl?taxon_id=67620.
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Citation: Department of the Environment (2018). Neoceratodus forsteri in Species Profile and Threats Database, Department of the Environment, Canberra. Available from: http://www.environment.gov.au/sprat. Accessed Wed, 14 Mar 2018 17:17:50 +1100.