The Ecology of the Quokka (Setonix Brachyurus) (Macropodidae: Marsupialia) in the Northern Jarrah Forest of Australia
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The Ecology of the Quokka (Setonix brachyurus) (Macropodidae: Marsupialia) in the Northern Jarrah Forest of Australia. by Matt Hayward A thesis submitted in fulfillment of the requirements for the degree of Doctor of Philosophy School of Biological, Earth and Environmental Science, University of N.S.W. JULY 2002 To Mum 2 “…I saw my first quokka. It came shambling out of the low shrub that surrounded the knoll and stood regarding me insolently. It was a malicious-looking beast, with a rat-like cast of features and a small, mean mouth, but it was small and showed no immediate aggressive tendencies. I was not alarmed. I looked at it and it looked at me and the decencies seemed to require some sore of overture on my part. I broke off a piece of cheese and some apple and tossed them to the quokka. It leaned forward and sniffed the offering, then looked up at me suspiciously with no sign of gratitude. “Go on” I said amiably, “tuck in. Cheese and apple never hurt a quokka.” The quokka leaned forward again and picked up the offering with both its tiny paws. It sniffed again, delicately took the apple in its mouth, chewed for a moment, wrinkled its nose and spat the apple out. “All right,” I said. “So quokkas don’t like apples. Try the cheese.” It did. It popped the gorgonzola into its mouth and chewed experimentally. Then it fell on its back in a dead faint. I leaped [sic] to my feet in great agitation. I was then a convinced conservationist and thought creatures such as quokkas should be preserved at all costs… I emptied my haversack, picked up the small, quivering body and gently laid it inside. I carefully left the top flap open so the quokka could breathe, put the haversack on my back and leaped, or rather clambered, onto my bicycle… I went racing down the slope, pedalling hard. Then it became apparent that I was moving so fast that pedalling was beside the point. I stopped and sat in the saddle concentrating on steering the bicycle at what was, for me, a breathtaking speed… Then the quokka began to recover. I became aware of a violent stirring at my back and then obvious movements indicating that it was climbing out of the haversack. I was, for a moment, vaguely concerned that it might fall to the ground at this impossible speed and hurt itself, but all I could do was cling to the handlebars and try to keep the bicycle in the middle of the road. I felt the quokka reach the top of the haversack, then I felt its tiny paws on my neck. Even with the rushing wind of my passage I became aware of a strange smell, a combination of mildewed carpet and gorgonzola… There was something shrieking in my left ear. It was the quokka, panicking. It was climbing around my neck trying, it seemed, to get in front of me… Then the quokka sank its teeth into my left ear. I heard my own yelp of pain over the rushing wind and took my left hand off the handlebar to try and prise the quokka loose… Then it stopped … and just let its dead weight hang on my ear from its clamped jaws. 3 A quokka weighs only a kilogram or two, but you try riding a runaway bicycle down a steep hill with a couple of kilograms of quokka hanging onto your left ear… Tugging away at the quokka’s tail, bellowing in pain and fear, I swung into that bend, bouncing away at a hundred kilometres an hour at least, and I hadn’t the slightest chance in the world of getting around it. The machine bounced once at the beginning of the bend, once more at the edge of the cliff, and then sailed swiftly into the air and down into the sea five metres below… I wasn’t hurt and the quokka seemed to get dislodged in the flight. I poked my head out of the water and saw the wretched thing standing on the water’s edge, glaring at me, and snarling. I bought myself a drink at the pub and the barman asked me what had happened to my ear. I told him I had bumped into a tree. Extract from Wombat’s Revenge (Cook 1987) 4 Abstract The quokka (Setonix brachyurus Quoy & Gaimard 1830) is a medium-sized, macropodid marsupial that is endemic to the mesic, south-western corner of Australia. While being a tourist icon on Rottnest Island, the species is threatened with extinction. It has been intensively studied on Rottnest Island in the 1960s and 1970s, however very little is known of its ecology on the mainland. Additionally the insular and mainland environments are extremely different suggesting that ecological differences between the two populations are likely. Consequently, this study sought to determine the basic autecology of the quokka and identify what factors have attributed to its threatened conservation status. The northern jarrah forest of Western Australia was selected as the study region due to it being at the northern limit of extant quokka distribution and because it was thought that the factors threatening the quokka would be exacerbated there. Fossil deposits suggest that the quokka originally occupied an area of approximately 49,000 km2 in the south-western corner of Australia. Historical literature show that they were widespread and abundant when Europeans colonised the region in 1829 but a noticeable and dramatic decline occurred a century later. The arrival of the red fox to the region coincided almost exactly with this decline and so it was probably ultimately responsible. Continued predation by both it and the feral cat are likely to have continued the decline, along with habitat destruction and modification through altered fire regimes. Specific surveys and literature searches show that since the 1950s, the area occupied by the quokka has declined by 45% and since 1990 by 29%. Based on the criteria of the IUCN (Hilton-Taylor 2000), the conservation status of the quokka i should remain as vulnerable. An endangered status may be more applicable if the quokkas restriction to patches through its existence as a metapopulation is considered. Trapping of eight sites supporting quokka populations in the mid-1990s revealed three sites now locally extinct despite the ongoing, six year old, fox control programme. Another three are at serious risk of extinction. Extant population sizes ranged from one to 36 and population density ranged from 0.07 to 4.3 individuals per hectare. This is considered to be below the carrying capacity of each site. The overall quokka population size in the northern jarrah forest may be as low as 150 adult individuals, of which half are likely to be female. Even the largest extant populations are highly susceptible to stochastic extinction events. This small size was surprising considering the six year old, introduced predator control programme. Historically, the restriction to discrete habitat patches, the occasional inter-patch movement, the lack of correlation between the dynamics of each population and reports of frequent localised extinctions and colonisations suggest that the quokka population once existed as part of a classic metapopulation. The massive decline of the quokka in the 1930s pushed the metapopulation structure into a non-equilibrium state such that today, the extant populations are the terminal remnants of the original classic metapopulation. Wild mainland quokkas breed throughout the year. A significant reduction in the number of births occurs over summer and this coincides with a decline in female body weight. Despite this, the mainland quokka is relatively fecund and is able to wean two offspring per year. The level of recruitment from pouch young to independence was low and this may explain the apparent lack of population increase following the initiation of fox control. ii A total of 56 trapped quokkas were fitted with a radio collar. Mean home range size for quokkas was 6.39 ha with a core range of 1.21 ha and this was negatively related to population density. Male home ranges were larger than females but not significantly when the sexual size dimorphism was considered. Nocturnal ranges were larger than diurnal ranges reflecting nocturnal departures from the swamp refugia. Home range sizes varied seasonally, probably due to changes in the distance required to move to obtain sufficient nutrients and water over the dry summer compared to the wet winter and spring. Telemetry confirmed trapping results that showed no movement between swamps or populations. Home range centres shifted to the periphery of the swamp following the winter inundation and this may increase the species susceptibility to predation. The lack of dispersal is probably caused by quokka populations existing below carrying capacity and following selection for philopatry under the threat of predation for dispersing individuals. Without dispersal to recolonise or rescue unpopulated patches, the collapse of the original quokka metapopulation appears to have occurred. On a macrohabitat scale, the quokka in the northern jarrah forest is restricted to Agonis swamp shrubland habitats that form in the open, upper reaches of creek systems on the western side of the forest. This restriction was probably initially due to the high water requirements of the quokka but is likely to have been exacerbated by increased predation pressure since the arrival of the fox. On a microhabitat scale, the quokka is a habitat specialist, preferring early seral stage swamp habitats, probably for foraging, as part of a mosaic of old age swamp that provides refuge. Despite the six year old, introduced predator control programme, foxes and cats are still the major cause of mortality to quokkas.