MACROPODS

Macropods are classified as browsers, grazers or intermediate feeders (Table 14.4), correlating with their dentition and gastrointestinal anatomy (Vogelnest and Portas 2008). They are foregut fermenters (Fig.

14.1a), however the anatomy of the macropod stomach is quite different to domestic ruminants and has been described in detail (Freudenberger et al. 1989; Dawson 1995; Hume 1999; Tyndale-Biscoe 2005; Vogelnest and Portas 2008). The foregut microbial communities of macropods are dominated by Bacteroidetes and Firmicutes phyla which are involved in fibrolytic digestion (Chong et al. 2020). At the genus level, however, macropods harbour carbohydrate hydrolysing microbes which are distinct from that of the bovine rumen (Pope et al. 2010).

Managed care diets should reflect the nutritional composition of the natural diet with sufficient and appropriate fibre and digestible energy. The diets of browsers and intermediate grazer/browsers tend to be lower in fibre and higher in digestible energy, whereas those of grazers are higher in fibre and lower in digestible energy. Obesity, dental and GI diseases are concerns when animals are fed diets with excess digestible energy (Vogelnest 2015). Macropods are often fed hay, leafy greens, vegetables, fruits and commercial pellets or kibble. Dog and cat kibbles (which may also have excessive levels of vitamins A and D) should be avoided in favour of pellets designed for herbivores with low non-structural carbohydrate (NSC) content (< 15%) and all items high in sugar and/or starch (> 15%) should be restricted. Specific diets for macropods are presented in Sela and Mills (2010). Some macropods cannot be classified as obligate grazers or browsers. The frugivorous musky rat-kangaroo (Hypsiprymnodon mos- chatus) relies on a diet extremely low in fibre, consisting of fruits, soft-shelled seeds and invertebrates.

Their digestive system is simple and unspecialised, showing minimal fermentation ability (Dennis 2002). Potoroids are mycophagous. They are best described as concentrate selectors, selecting diets low in fibre. Hypogenous fungi constitute between 70-90% of the diet of eastern (Bettongia gaima- rdi) and brush-tailed (B. penicillata) bettongs and poto- roos (Potorous spp.) (Claridge et al. 2007). Other dietary items include browse, tubers, seeds and invertebrates which contribute in greater proportions to the diets of the northern (B. tropica), burrowing (B. lesueur) and rufous (Aepyprymnus rufescens) bettongs (Table 14.4).

Tree-kangaroos (Dendrolagus spp.) consume a diet dominated by leaves, but will also eat the fruits, seeds and flowers of trees. There is anecdotal evidence of free- ranging tree-kangaroos consuming birds and bird eggs (Martin 2005), however, it is advised a diet high in fibre including a variety of browse and some vegetables is more appropriate for zoo-housed animals to avoid obesity (Table 14.4) (Edwards and Ward 2001).

Macropod progressive periodontal disease (MPPD) is a significant cause of morbidity and mortality in macropods held in managed care. Inappropriate diets (e.g. low- fibre, high sugar), feeding hay with a high proportion of stalk compared to leaf and poor hygiene around feeding stations are all diet and feeding-related risk factors for development of MPPD (see Chapter 32). Providing long grass hay or fibrous tree bark has been shown to reduce the incidence of MPPD (Hume 2005). This also provides behavioural enrichment where animals can consume the leaves and strip the bark (Johnson-Delaney 2014).

Tammar wallabies (Notamacropus eugenii), rednecked pademelons (Thylogale thetis), quokka (Setonix brachyurus), nail-tailed wallabies (Onychogalea spp.), tree-kangaroos and musky rat-kangaroos are prone to developing a syndrome of weakness and wasting of the hind limbs due to myopathy which can be prevented with vitamin E supplementation, but selenium supplementation has no prophylactic effect (Kakulas 1961, 1963; Speare et al.

1989). The myopathy is not entirely diet related with studies showing a correlation with stress factors such as smaller habitat size and incidence of myopathy. A balanced diet containing a complete pelleted feed to supplement browse, hays and vegetables, should provide dietary levels of vitamin E > 100 IU∕kg dry matter which is sufficient to reduce this risk.

Calcium based uroliths have been reported in red kangaroos (Osphranter rufus), eastern grey kangaroos

Table 14.4. Feeding strategy, natural diet, managed care diet, mean retention times and animal models for Australian mammals.

Animal species	Feeding strategy³	Natural diet	Captive diet^b	Mean retention time (hrs)	Domestic animal model	Comments^c (see explanation of superscripts below table)
Order Monotremata
Short-beaked echidna (Tachyglossus aculeatus)	Cl, FG	Terrestrial invertebrates'²	Properly balanced in-house meat mixes and/or commercial Insectivore mixes, supplemented with insects, fibre sources, termites/mounds	48³	Understudy	Vitamin AandD toxicity if avian supplements are used Meat-based^c Avoid sugar Fibre may be necessary
Platypus (Ornithorhynchus anatinus}	Cl	Benthic macroinvertebrates (i.e. crayfish, beetles, midges, snails, crustaceans, caddisfly, mayfly larvae) from over 55 Familiesand 16 orders^3,4'⁵'⁶	Yabbies, earthworms, mealworms, fly pupae, maggots, freshwater shrimp, black worms, prawns, goldfish, tubifex worms, trout fingerlings and aquatic insect larvae	5³	Cat Dog	Low Tiskofnutritional deficiency if fed variety of live prey
Order Notoryctemorphia
Marsupial mole (Notorcytes spp.)	Cl	Termites, seed-eating ants, weevil larvae, beetles, arthropods⁷	Variety of insects, hard-boiled egg, cat/dog foods⁸, Insectivore mixes	-	Cat Dog	Insect-based diet^d
Order Dasyuromorphia
Numbat (Myrmecobius fasciatus)	Cl	20,000 termites from > 25 species daily¹⁴	Properly balanced in-house mixes that Outritionallycomplement termites (primarily Nasutitermes and Schedorhinotermes spp.), other insects as available	24-27¹⁵	Cat Dog	Insect-based diet^d
Antechinus spp. Parantechinus spp.	O	Variety of invertebrates and vertebrates, plants and nectars seasonally, fungi (some species)¹⁶	Dog food, hard-boiled egg, various insects, mice, chicks, Insectivore mixes	-	Dog Pig Rat Mouse	Low risk if fed variety of feed items that includes a complete feed⁸
Kaluta (Dasykaluta rosamondae)	O	Primarily invertebrates (beetles and ants), some vertebrates and plant material¹⁷	Dog food, hard-boiled egg, various insects, mice, chicks, vegetables, leafy greens, browse, Insectivore mixes	-	Dog Pig Rat Mouse	Insect-based diet^d
Kowari (Dasyuroides byrnei) Crest-tailed mulgara (Dasycercus cristicauda)	C	Insects, reptiles, arachnids, small birdsand mammals⁷	Dog food, hard-boiled egg, various insects, mice, chicks, Insectivore mixes	87-150 min (D. byrnei)⁷	Cat Dog	Low risk if fed variety of feed items that include a complete feed

14-Nutrition 239

Table 14.4. (continued) border=0> Spotted-tailed quoll (Dasyurus maculatus) C Greater gliders, rabbits, bandicoots and small macropods year-round. Insects and reptiles increase in summer¹⁸ Dog food, rodents, chicks, hard- boiled eggs, rabbit, various insects, earthworms, fish, yabbies 13-17.1 summer

5.9-7.3 winter (Eastern quoll, D.

Viverrinus)⁷ Cat

Dog Whole prey^e Phascogale spp. O Variety Ofvertebrates, plants and nectars seasonally⁷ Dog food, hard-boiled egg, various insects, rodents, chicks, Insectivore mixes, native blossoms Cat Dog

Pig

Rat Mouse Supplementation may be required Ifcompletefeed is not well accepted

Insect-based diet^d Kultarr

(Antechinomys Ianiger) Cl Arthropods (cockroaches, beetles, arachnids)¹⁹ Properly supplemented mincemeat, insects, cat/dog food²⁰ Cockroaches, mealworms, crickets, rodents, Wombaroo small carnivore mix (Wombaroo Food Products, Glen Osmond, SA)²¹ 3.9²¹ Cat

Dog Insect-based diet^d Ningaui

(Ningaui spp.) Cl Cockroaches, crickets, centipedes, spiders, butterflies and moths²² Various insects, dog/cat food, hard-boiled egg, Insectivore mix - Cat

Dog Insect-based diet^d Planigalespp. Cl Variety of insects (beetles, spiders, isopods, grasshoppers)⁷ Variety of insects, cat/dog food, hard-boiled egg, mice, chicks, Insectivore mixes, native blossoms - Cat

Dog Insect-based diet^d Dunnart

(Sminthopsis spp.) Cl Variety of insects (beetles, spiders, isopods, grasshoppers)⁷ Variety of insects, cat food, chicks, earthworms, rodents, Insectivore mix 53 min (S.

Crassicaudata)²⁰ Cat

Dog Insect-based diet^d Tasmanian devil (Sarcophilus harrisii) C Scavenge on medium- to large-sized mammals. May also consume birds, fish and insects²³ Variety of whole prey; kangaroo, chickens, rabbits, rodents — Cat

Dog Whole prey^e Order Peramelemorphia Greater bilby (Macrotis Iagotis) O Predominantly seed-based in winter. Invertebrates, vertebrates and plant material (leaves, roots, grasses, stem/bark, seeds, flowers, berries) and fungi^24,25,26 Variety of vegetables, seeds, insects and dog food 17.9²⁷ (fluid)

23.5²⁷ (particle) Dog Pig

Rat Mouse Seed-based diet^f

^aFeeding strategy: C (carnivore), Cl (insectivorous carnivore), CP (piscivorous carnivore), O (omnivore), B (browsing herbivore), G (grazing herbivore), I (intermediate herbivore), N (nectarivore). Fibre fermentation: The herbivores (H, I, B) can be further divided by their site of fibre fermentation; FG (foregut), HG (hindgut - caecum and/or colon)

^bForm ofmanaged care diet is not outlined. Complete feeds should meet nutrient requirements for domestic animals (i.e. AAFCO 2014). Herbivore pellets may be formulated for cow, horse goat, guinea-pig, rabbit or kangaroo. Dog/cat foods (dry kibble or wet) may need to be soaked Orcrushed for smaller species. Whole prey must be Ofappropriate size or Cuttoappropriate size. Supplements may be required for total diet to meet minimum requirements of animal model suggested.

^cMeat-based diets are imbalanced if not supplemented. Common deficiencies include calcium, copper, zinc, and vitamin E. ^dlnsect-based diets are Susceptibleto imbalances if no complete diet is fed. Common deficiencies include calcium, vitamins A, D, E, thiamine, methionine, arginine, carotenoids).^9,10,11'¹²'^{13 e}WhoIe prey may require vitamin E supplementation.

^fVitamin EandVitamin B deficiencies are common in diets that Iackvarietyand are predominantly seed-based.

?Forage-based diets are Susceptibleto deficiencies from hay and grasses. These will vary geographically and require nutrient testing. In NSW common deficiencies include calcium, copper, zinc and vitamin E. A complete herbivore feed should comprise 25-30% of the total diet. Acidosis if fed excessive concentrate or high sugar items.

^hln-house honey mixes are often deficient in calcium, copper, zinc, and vitamin E. ^lFish-based diets should be supplemented with vitamin B₁ (30 mg/kg diet) and vitamin E (100 IU∕kg diet) to prevent deficiency.

¹Spencer and Richards (2009); ²Sprent and Nicol (2016); ³HoIz (2015); ⁴Bethge etal. (2003); ⁵McLachIan-Troup etal. (2010); ⁶KIamt etal. (2016); ⁷Hume (1999); ⁸AAFCO (2014); ⁹Attard (2013); ¹⁰Finke (2002); ¹¹Ferrie etal. (2014); ¹²Gietzen etal. (2007); ¹³Rodriguezand Pessier (2014); ¹⁴Cooperand Withers (2004); ¹⁵Hogan etal. (2013); ¹⁶Vernes and Dunn (2009); ¹⁷Paveyefa/. (2016); ¹⁸GIen and Dickman (2006); ¹⁹Stannard and Old (2014); ²⁰Geiser (1986); ²¹Stannard and Old (2011); ²²Bos and Carthew (2007); ²³Pemberton etal. (2008); ²⁴O'Hara etal. (2012); ²⁵Vernes (2014); ²⁶Vernesefa/. (2015); ²⁷Gibson and Hume (2000); ²⁸PhiIIipsand Pratt (2008); ²⁹DierenfeId (2009); ³⁰Vernesefa/. (2001); ³¹Smith (2009); ³²Hayward (2005); ³³MiIneef al. (2016); ³⁴Roswag efα∕. (2012); ³⁵SchmeIitschekefa/. (2009); ³⁶Jackson (2003); ³⁷Goto etal. (2008); ³⁸Lanyon and Marsh (1995); ³⁹Brookand Kutt (2011); ⁴⁰Behrendorffefa/. (2016); ⁴¹Berryefa/. (2017); ⁴²Hardyefa/. (2017); ⁴³CIaussand Paglia (2012)

(Macropus giganteus), red-necked wallabies (N. rufogri- seus), yellow-footed rock-wallabies (Petrogale xanthopus) and tammar wallabies, with castrated males at a higher risk of developing urolithiasis (Bryant and Rose 2003). Excessive calcium, either by unintentionally increasing the calcium to phosphorus ratio in the diet (e.g. high proportion of lucerne) or consumption of plants grown in high calcium soil, high levels of vitamin D and possibly hypovitaminosis A may be contributing factors in the development of calcium based uroliths (Bryant and Rose 2003; Lindemann et al. 2013).

Golden bandicoot (Isoodon auratus)	O	Predominantly invertebrates (particularly arthropods in breeding season), vertebrates and plant material (leaves, roots, grasses, stem/bark, seeds, flowers, berries) and fungi^24,25,26	Variety of vegetables, seeds, insects and dog food	24—33⁷ (fluid) 11-27⁷ (particle)	Dog Pig Rat Mouse	Vitamin B deficiency reported when fed all-vegetable diet^7,33
Order Diprotodontia
Suborder Vombatiformes
Koala (Phascolarctos cinereus)	B, HG	Eucalyptus spp. (primarily tip and young leaves).	A variety of eucalyptus spp. (5 branches 2m in length daily).	213⁷ (fluid) 99⁷ (particle)	Horse	Low risk if fed a variety of Eucalyptus spp.
Southern hairy-nosed wombat (Lasiorhinus Iatifrons)	G,HG	Grasses²⁸	Herbivore cube (horse), grass hay, fresh grass, small amount of root vegetables	36⁷ (fluid) 62⁷ (particle)	Horse	Forage-based diet^g
Suborder Phalanderiformes
Southern common cuscus (Phalanger mimicus)	F, HG	Fruit and tree leaves⁷	Native browse, herbivore pellet, fruit, glider mix, blossoms	8⁷ (fluid) 51⁷ (particle)	Rabbit Guinea-pig	Prone to obesity Honey mixes^h
Eastern ring-tailed possum (Pseudocheirus peregrin us)	B, HG	Eucalypt spp. (leaves), native fruits and blossoms	Native browse, herbivore pellet, variety of low sugar and low starch vegetables, leafy greens	63⁷ (fluid) 37⁷ (particle)	Rabbit Guinea-pig	Pronetoobesity Limit brassicas to avoid bloat Honey mixes^h
Yellow-bellied glider (Petaurus australis)	I, HG	Plant and insect exudates, blossoms, insects, spiders and pollen⁷	Glider mix, fruit, acacia gum, insects, native blossoms	-	Rabbit Guinea-pig	Pronetoobesity Honey mixes^h
Sugar glider (Petaurus breviceps), Leadbeater's possum (Gymnobelideus Ieadbeateri).	N, HG	90% Acacia gum, nectar and pollen²⁹	Glider mix, vegetables, leafy greens, acacia gum, insects, native blossoms	29 (P. breviceps)⁷	Rabbit Guinea-pig	Prone to obesity May be Susceptibleto haemosiderosis⁴³ Honey mixes^h
Common brush-tailed possum (Trichosurus vulpecula)	B, HG	Eucalyptus spp. (leaves)⁷	Native browse, herbivore pellet, variety of low sugar and low starch vegetables, leafy greens, insects	64 (fluid -high fibre)⁷ 71 (particle - high fibre)⁷ 51 (fluid - natural foliage) ⁷ 49 (particle -natural foliage)⁷	Rabbit Guinea-pig	Limit brassicas to avoid bloat Pronetoobesityand May be susceptible to haemosiderosis ⁴³ Honey mixes^h

Feather-tailed glider (Acrobates spp.)	N, HG	Nectar, pollen and insects⁷	Glider mix, fruit, pollen, insects	7 (32% pollen in diet)⁷ 24 (4% pollen in diet)⁷	Rabbit Guinea-pig	Pronetoobesity, iron storage disease, lipid keratopathy Honey mixes^h
Greater glider (Petauroides volans)	B, HG	Eucalypt spp. (leaves)	Native browse, blossoms, glider mix, acacia gum, pectin	51⁷ (fluid) 23⁷ (particle)	Rabbit Guinea-pig	Prone to obesity, iron storage disease Honey mixes^h
Suborder Macropodiformes
Eastern grey kangaroo (Macropus giganteus)	G, FG	Grasses⁷	Grass hays, vegetables, leafy greens, herbivore pellet	14⁷ (fluid) 30⁷ (particle)	Goat sheep cattle	Forage-based diet^g Obesity Dental lesions Calcium uroliths
Tammarwallaby (Notamacropus eugenii)	I, FG	Grasses and forbs⁷	Herbivore pellets, lucerne and grass hay, low sugar vegetables, grasses, native browse	16.7⁷ (fluid) 25.8⁷ (particle)	Goat sheep cattle	Forage-based diet^g Prone to calcium uroliths
Long-nosed potoroo (Potorus tridactylus) Rufous bettong (Aepyprymnus rufescens)	G, FG	Hypogeal fungi, roots, tubers, bulbs, fruit, seeds and invertebrates³⁰	Herbivore pellets, insects, vegetables, seeds, nuts	20-31⁷ (fluid) 20-37⁷ (particle)	Goat sheep cattle	Forage-based diet^g Prone to obesity
Tree-kangaroo (Dendrolagus sp.)	B, FG	Leaves, fruit, blossoms, seeds⁷	Variety Ofvegetables, leafy greens, lucerne, herbivore pellets, browse	-	Goat sheep cattle	Limit brassicas to avoid bloat Acidosis (Eucalyptus spp. are toxic to the forestomach bacteria and should not be fed)⁷
Swampwallaby (Wallabia bicolor)	I, FG	Leaves from shrubs and vines, grasses, young seedlings, ferns, fungi and reeds⁷	Herbivore pellet, lucerne and grass hay, vegetables, leafy greens, native browse	-	Goat sheep cattle	Forage-based diet^g
Nail-tailed wallabies (Onychogalea spp.)	G, FG	Grass and forbs³¹	Herbivore pellet, grass hay, low Sugarvegetables	-	Goat sheep cattle	Forage-based diet^g
Hare-wallabies (Lagorchestes spp.)	I, FG	Grass and browse³¹	Herbivore pellet, grass hay, low sugar vegetables, native browse	23⁷ (fluid) 39⁷ (particle)	Goat sheep cattle	Forage-based diet^g
Quokka (Setonix brachyurus)	B, FG	Primarily leaves and stems, insect elytra, seeds, fungi, pollen and blossoms³²	Herbivore pellet, native browse, IowsugarvegetabIes_zVitamin E supplementation		Goat sheep cattle	Vitamin E should be supplemented Prone to obesity, mesenteric fat necrosis, dental disease Limit lucerne to avoid calcium build up and uroliths

Red-necked pademelon (Thylogale thetis)	B, FG	Ferns, leaves, short grass, fruit³¹	Native browse, lucerne hay, herbivore pellets	12.5⁷ (fluid) 25.1⁷ (particle)	Goat sheep cattle	Vitamin E should be supplemented
Order Chiroptera
Ghost bat (Macroderma gigas)	C	Small vertebrates such as lizards, frogs, birds and other bats³³	Variety of whole prey (chicks and rodents)	-	Cat	Meat based^c Whole prey^e
Microbats	Cl	Wide variety of insects, predominantly Lepidoptera but also Coleptera, Hemiptera, and Hymenoptera³³	Variety of insects (mealworms, crickets, cockroaches, silkworms, fly pupae)	86-332 min (Myotis spp.) 180-280 min (Pipistrellus spp.)³⁴	Cat	Insect-based diet^d
Grey-headed flying-fox (Pteropus poliocephalus)	F, HG	Rainforest fruits to pollen and nectar of Myrtaceae and Proteaceae species from the beginning of December to the end of March³⁵	Fruit, native blossoms, insects, protein and vitamin/mineral supplementation		Rabbit Guinea-pig	Multiple deficiencies if fed only fruit without supplementation
Order Rodentia
Greater stick-nest rat (Leporillus conditor) Black-footed tree-rat (Mesembriomys gouldii)	O	Fruit, large seeds, grass, termites and molluscs (M. gouldii)³⁶	Variety Ofvegetables, leafy greens, browse, lucerne hay, seeds and nuts	-	Rat Mouse Rabbit Guinea-pig	Pronetoobesity, diabetes, cataracts. Should be fed a more herbivorous diet
Order Sirenia
Dugong (Dugong dugon)	B, HG	Seagrasses (including roots and rhizomes)³⁷	Variety of greens (cos, kale, spinach, endive, iceberg lettuce), small amounts of roots or tubers	146-166³⁸	Horse	Prone to anaemia when fed cos lettuce alone (see Chapter 47)
Order Carnivora
Surborder Caniformia
Dingo (Canis fam Hiar is)	C	Medium-sized macropods, small-sized marsupials, European rabbit (Oryctolagus cuniculus), domestic livestock and poultry³⁹ Stranded cetaceans and turtles in coastal populations⁴⁰	Variety of whole prey, carcasses (rabbit, chicken, deer, kangaroo, rodents, bones) and limited dog food		Cat Dog	Meat based^c Whole prey^e

Long-nosed fur seal (Arctocephalus forsteri), Australianfurseal (Arctocephalus pusillus) Australian sea lion (Neophoca cinerea}	CP	Dominated by cephalopods in summer and fish in winter^41,42	Fish, squid, vitamin E and vitamin B₁ supplement		Cat	Fish diet'
Order Cetacea
Suborder Odontoceti
Dolphins	CP	Fish	Fish, squid, vitamin E and vitamin B₁ supplement	-	Cat	Fish diet' Prone to iron overload⁴³

MACROPODS

More on the topic MACROPODS: