DUNG INHABITANTS AND DUNG FEEDERS – A ROGUES’ GALLERY
The following list is not exhaustive; instead it is a guide to the types of insects (and a few other creatures) associated with dung. The world fauna of scarabaeine dung beetles alone is something of the order of 6,000 species, clearly beyond the scope of this book. There would come a point at which this would become a repetitive catalogue. Not very readable.
Identification to species level, for all the families and genera listed below, often requires meticulous examination of specimens under the microscope and the use of complex identification keys and detailed monographs. Instead I offer here a rogues’ gallery of the usual suspects. As well as feeding on or in the dung itself, associates feed on other dung-recyclers, on each other, they shelter under the dung, rest on top of the dung, or use it as bait.
I acknowledge that this list is rather Anglocentric, skewed in its content to north-west Europe, and heavily influenced by my own personal experience of dunging in the grazing meadows of southern England, but some New World and tropical species are also included to give a bit more breadth. Unless otherwise stated most species occur broadly across northern hemisphere Eurasia.
Hornet robber fly, Asilus crabroniformis
Size: Body length 20 mm.
Description: Large dark-brown fly, abdomen nearly black, with bright yellow tail tip; wings smoky, legs bristly yellow, head with prominent pointed snout. Hornet mimic, but harmless to humans, lacks sting.
Life history: Secretive larvae in soil layer, foodstuff uncertain, but possibly preying on other small soil invertebrates, take at least 2 years to reach maturity; adult flies predatory, attacking flies, beetles, grasshoppers or other insects, usually in midair, and killing with powerful skewer mouthparts.
Ecological notes: Adults (summer and autumn flying) sit on dry cow-pats (sometimes bare soil patches or twigs), using them as launch pads from which to attack insects flying past, including dung flies and dung beetles heading in towards the pat. Eggs are thought to be laid in the dung (cattle and rabbit recorded), leading to suggestion that larvae are specialist predators of dung beetle larvae, but no other robber flies (large world-wide family Asilidae) are dung-associated; study is still ongoing.
Snipe fly, Rhagio scolopaceus (and other species)
Size: Length 12 mm, wing length 10 mm.
Description: Grey fly, with long legs and conical abdomen clearly marked with yellow. Wings clear, but with brown and orange mottle pattern.
Life history: Predatory larvae, pale, worm-like, live in soil, leaf litter, wood mould, occasionally invade old dung. Adults are not known to feed.
Ecological notes: Adult flies have distinctive alert stance, resting head-down on tree trunks and other vertical surfaces. Intuitively they look as if they are poised for hunting forays, but these are usually males, and the only hunting they do is after the females which mostly rest on the ground.
Yellow dung fly, Scathophaga stercoraria
Size: Body length 5–6 mm.
Description: Furry or bristly fly, dull greenish grey in female, but densely bright golden furred in male.
Life history: Eggs laid in fresh dung (cow-pats especially preferred); pale whitish-grey, slim, conical maggots feed in the dung for 10–20 days then pupate in the soil; adults emerge 10–80 days later depending on temperature. Adults predatory on flies and other small insects.
Ecological notes: One of the most obvious and important dung insects. Fresh dung, minutes on the ground, soon attracts males which flit about actively and aggressively waiting for females. Mating takes 20–50 minutes, but male remains on female’s back ‘guarding’ her for similar (or longer) time, to prevent other males diluting his sperm through multiple copulations. Female leaves after egg-laying, so active pats often male-dominated, leading to squabbles over incoming receptive females. Wetter or looser dung types preferred: cow or horse in meadows, but cat, dog or fox in urban parks and gardens. Several very similar species.
Black ‘ant’ flies, Sepsis and other genera
Size: Body length 2–6 mm.
Description: Tiny, black, shining flies, with relatively long legs, slim waist and round head, giving an ant-like appearance. Clear, narrow wings tipped with small but distinct black dot near apex.
Life history: Breed in soil, dung, decaying vegetation and other rotten organic matter.
Ecological notes: Large ‘swarms’ of the flies (many thousands) can emerge and crawl/fly about in the grass where the cow-pat or other dropping has long gone. As they crawl over the vegetation, they flick their spotted wings about in territorial or courtship display.
Moth flies, family Psychodidae
Size: Length to 2.5 mm.
Description: Tiny, slightly fluffy, flies with short, broad wings, also covered or patterned with fluffy scales. Wings held flat over back in delta shape, hence moth name. Various species coloured black, through brownish mottles to pure white.
Life history: Tiny pale maggots feed in wet detritus, soil, rotting fungus, leaf litter, manure, compost and dung.
Ecological notes: Often found swarming in clouds close over and around dung. Small size means they can crawl between the pellets or folds of the dropping. They are a major inhabitant of trickle beds in sewage treatment works. Some species breed in drains, including in the algal film growth in domestic sink overflows in kitchen or bathroom, but not implicated in disease spread.
Daddy longlegs, or craneflies, Tipula species
Size: Wing length to 30 mm, body length to 35 mm, leg span to 55 mm.
Description: Narrow-bodied, narrow-winged, long-legged flies, various shades of grey, sometimes with wings smoky or marked with dark mottled patterns. Adults fly with ungainly bobbing flight, often half crawling, through the long grass. Some species attracted to porch lights.
Life history: Stout cylindrical larvae (‘leatherjackets’) are grey wrinkled and short worm-like, feed in soil, leaf litter, root thatch, on living and dead plant material.
Ecological notes: Not really dung specialists, but larvae often found under old dung where soil horizon is blurred.
Biting midges, Forcipomyia species
Size: Wing length to 4 mm.
Description: Small to minute grey or blackish flies. Bodies slightly hunched, wings clouded with dusting of microscopic dark scales. Males have distinctive plumed antennae.
Life history: Pale grey bristly larvae live in soil, dung and leaf litter, eating decaying organic matter, fungi or algae.
Ecological notes: Although part of ‘biting midge’ family, are inoffensive to humans or stock animals, probably biting and sucking body juices from other, larger, insects. A Forcipomyia midge achieved the rare distinction of having the highest recorded wing beat of any insect, at over 2,200 beats per second. However, this was only in the laboratory, and after the researcher had heated up the unfortunate fly, and cut most of its wing membrane off.
Biting midges or ‘punkies’, Culicoides species
Size: Wing length to 6 mm.
Description: Small to minute grey, brown, black or yellowish flies. Body stout and hunched, wings often patterned with dark mottles. Male antennae thickly plumose.
Life history: Tiny pale worm-like larvae feed in dung, or waterlogged soil, eating fungi, algae or decaying organic material.
Ecological notes: Adult flies bite and suck blood from humans and farm animals. These are the midges that make upland areas intolerable for much of the year. In North America, they are blamed for huge financial losses to the tourist industry because their flight season seriously curtails outdoor activities. C. imicola is main vector implicated in spread of bluetongue virus among sheep in Europe.
Non-biting midges, family Chironomidae
Size: Length to 10 mm, wing length 8 mm.
Description: Narrow-winged, narrow-bodied, long-legged flies, various shades of grey, brown or black. Males with broad feathery antennae. Closely resemble mosquitoes, to which they are related, but lack any biting mouthparts.
Life history: Most species are aquatic, larvae living in the mud at the bottom and sides of ponds, ditches and slow streams, feeding on decaying organic matter. Bobbing midge clouds over water are often members of this large family.
Ecological notes: Some species renowned for their ability to live in low-oxygen muds; these larvae contain the strongly red-coloured oxygen-storing chemical haemoglobin (the same as found in mammalian blood), giving them a bright scarlet colour and earning them the name bloodworms. Not surprisingly a few species have been reared from dung.
Window gnat, Sylvicola punctata
Size: Wing length to 6 mm.
Description: Small, greyish brown, mosquito-like fly with tiny spherical head. Wings marked with dark cloud of mottles, sometimes approximating to three dark blotches.
Life history: Slim, worm-like larvae live in soil, dung and leaf litter, eating fungi, algae and decaying organic matter. Adult flies mostly nocturnal.
Ecological notes: Gets its common name from its habit of appearing on the inside of domestic windows after emerging from pupae in the soil or compost brought in with potted house-plants, but does not visit food and is not implicated in any disease spread.
Fungus midges, family Sciaridae
Size: Wing length to 3.5 mm.
Description: Small, dark, delicate flies with long legs, long antennae, long wings and narrow bodies. Wings often darkened. Fly slowly, but scuttle quickly. Tibiae of all legs with long distinct spurs at ends.
Life history: Slow-moving larvae feed on fungi, in soil, in decaying organic matter.
Ecological notes: Often found breeding in old dung when fungal decay has set in. Sometime pest in mushroom farms where they can breed in the rotted manure-based compost, but also potentially attack the hyphae of the crop, or invade any overripe mushrooms on the turn.
Dung midges, family Scatopsidae
Size: Length to 3 mm, wing length 2.5 mm.
Description: Minute dark or black flies with seemingly over-long wings or extra short bodies. Abdomen short oval, legs short, wings whitish with veins unpigmented, antennae short and stout.
Life history: Slow-moving bristly larvae breed in soil, leaf litter and dung. Adults often congregate in large numbers on flowers.
Ecological notes: A poorly studied group of minute flies, little is known about their habits or life histories as demonstrated by their other common name ‘minute black scavenger flies’. In marshy areas of grazing meadows they sometimes form clouds of many thousands skittering about in the long grass, or over scrub.
St Mark’s flies, Bibio marci (and other species)
Size: Length to 15 mm, wing length 12 mm.
Description: Large, black, sooty-looking fly, with robust body. Broad wings pale but dark-edged in male, heavily blackened in female. Flies heavily, with long black legs hanging down.
Life history: Pale larva, live in the soil, feeding on decaying (possibly also living) plant matter, so regularly invade old dung where humus horizon is blurred.
Ecological notes: Named for appearance, often in large clouds, around St Mark’s day, 25 April, although this is very approximate. Many similar species, some with bright red front legs. Smaller soil-inhabiting bibionids often called fever flies, for no very sound reason, can occur in huge numbers; larval densities of 37,000 per square metre are recorded.
Broad centurion, Chloromyia formosa
Size: Length 9 mm, wing length 7 mm.
Description: Broad, flat, parallel-sided, metallic green to violet soldier-fly, abdomen of males golden or bronze, wings broad and delicate, veins hardly visible, legs and antennae short.
Life history: Short, stout grey larva lives in dung, damp soil and wet leaf litter, feeding on decaying organic matter.
Ecological notes: Really a denizen of damp soil, but regularly occurring in the blurred boundary between humus and old dung. Common in fields, woods and gardens. Gets soldier name because related species sometimes brightly coloured like military dress uniforms.
Black-horned gem, Microchrysa polita
Size: Length 5 mm, wing length 4 mm.
Description: Tiny metallic green, or bluish, soldier fly, abdomen short, squat, parallel sided, almost hexagonal. Legs and antennae black (yellow in other species). Wings delicate, clear, veins hardly visible.
Life history: Tiny grey larva short and stout, breeds in and under fresh dung, in manure heaps and compost bins.
Ecological notes: Needs moist, putrescent organic matter. Many other species of soldier fly are aquatic, breeding in muddy pond edges, ditches and slow streams.
Twin-spot centurion, Sargus bipunctatus
Size: Length to 10 mm, wing length to 9 mm.
Description: Narrow, parallel-sided body, metallic green thorax but abdomen golden bronze in male, dark metallic blue with pinkish-orange girdle in female. Legs yellow.
Life history: Several similar species breed in wet meadows, marshy woods, in the damp soil, larvae feed on decaying organic matter.
Ecological notes: The most closely dung-associated of this large genus in Europe; often the only species found in towns and cities where it regularly breeds in cat, dog and fox dung.
Black soldier fly, Hermetia illucens
Size: Length to 16 mm, wing length to 14 mm.
Description: Dusky black fly, wings clouded, legs marked with white or yellow.
Life history: Native to south-eastern USA, breeding in compost and manure, animal dung and carrion. Larvae (‘phoenix worms’) are dung/decay feeders.
Ecological notes: Introduced throughout much of the northern hemisphere, either because maggots are supplied by pet shops as lizard or fish food, or as part of manure control in farms. The larvae are amenable to captive rearing on an industrial scale to process chicken or pig manure. The larvae are then edible to the livestock – a particularly neat recycling of nutrients. Also potentially edible by humans, but in processed ground meal form.
Long-footed flies, Dolichopus species (and many similar genera)
Size: Length to 6 mm, wing length 5 mm.
Description: Small greyish, or often metallic greenish, flies with slim legs on which the short body seems to perch, head up, tail down. Active and fast, scuttling and skipping about.
Life history: Predatory as adults, hunting and killing other small flies. Larvae in soil, mud, rotten wood, standing water; most are predatory eating small invertebrates.
Ecological notes: Not really dung-feeders, but will visit dung to prey on the other small flies it attracts.
Metallic scavenger fly, Physiphora alceae (formerly Chrysomyza demanata)
Size: Length 5 mm, wingspan 9 mm.
Description: Small, squat, short-bodied black fly with distinct metallic green reflections. Head red, eyes prettily striped red and orange. Wings pale, clear. Female abdomen pointed with telescopic egg-laying tube.
Life history: Breeds in manure heaps, compost and other mainly decaying plant matter, but also sometimes recorded on dung.
Ecological notes: An elaborate courtship dance has been reported, involving male leg-waving and wing-flicking.
Lesser dung flies, family Sphaeroceridae (formerly Borboridae)
Size: Length to 4.5 mm, wing length 4 mm.
Description: Tiny to very small black or dark grey flies, rather hunched, scuttle readily on their stout legs. Very many extremely similar species.
Life history: Larvae of some species are specialist dung-feeders.
Ecological notes: Adult flies will enter the dung to lay their eggs, using tunnels dug by other dung-users, mainly dung beetles. Their relatively short wings and preference for running, rather than flying, may aid them in their exploration of the dung tunnels and crevices.
Snouted hover fly, Rhingia campestris
Size: Length 12 mm, wing length 10 mm.
Description: Highly distinctive hover fly, thorax grey, streaked with darker markings, abdomen pinkish-orange with dark central stripe and dusky tail tip. Large eyes red, head greyish, but face hugely extended into a narrow orange ‘beak’ under which the long elbowed tongue is stored.
Life history: Stout, flattish larvae feed in dung, cow especially, and also sometimes in manure or compost heaps, and probably in other decaying organic matter.
Ecological notes: Adults rarely (if ever) found in the immediate vicinity of dung, and seemingly visit only to lay eggs, then depart. They are mostly found resting on herbage or visiting flowers and can use their long tongue to reach deep into plants with long corolla, which other flower visitors cannot reach. For a time this insect was called the Heineken fly, echoing the catchphrase of a long-running and highly successful 1970s advertising campaign, where the beer was able to ‘refresh the parts other beers cannot reach’.
Thick-legged hover fly, Syritta pipiens
Size: Length 6 mm, wing length to 6 mm.
Description: Slim, narrow hover fly; body dark, but abdomen clearly patterned with four yellow triangles in male, or two yellow triangles and four white tick marks in female. Face white, head dominated by large dark eyes. Legs dark brown, but marked with orange. Femur of hind leg grossly enlarged into bulbous swollen segment, with ridge of small sharp spines running beneath it.
Life history: Larvae in wet organic matter, including compost and manure heaps, silage and dung.
Ecological notes: Adults never seen at dung, but ubiquitous visiting flowers, where large numbers can congregate, making it one of the commonest of insects. Superb hoverers, males engage in head-to-head hovering jousts, making sudden darting movements (to other insects too), retreating and dodging, before eventually one retires and the other continues patrolling. Reason for thickened legs unknown.
Drone fly, Eristalis tenax
Size: Length 15 mm, wing length 13 mm.
Description: Glossy black or dark brown hover fly, stout body marked with orange-brown triangles or dashes on sides of abdomen. Eyes very large, dominating head. Closely resembles a honeybee (especially the large-eyed males – drones), for which it is frequently mistaken, especially by journalists and subeditors.
Life history: Semi-aquatic larva lives in watery detritus, has long rat-tail telescoping breathing tube to reach surface for air. Adult flies active all year, visiting flowers.
Ecological notes: Not strictly a feeder in dung, but usually in ditches where nutrient-rich run-off has entered from sewers, manure heaps, farmyards, slurry pits. This is the bugonia or oxen-born bee of the ancients who thought honeybees could spontaneously generate from the carcass of a cow, not realising that Eristalis was quite at home breeding in the putrescent semi-liquid decay of the corpse. The wasp-mimicking black and yellow Myathropa florea occurs in similar places.
Face fly, Musca autumnalis
Size: Length 7–8 mm, wingspan 13–18 mm.
Description: Superficially similar to house fly in shape, though slightly larger and sturdier. Grey, thorax dark-streaked, abdomen tessellated with chequer pattern of light and dark greys in female, male abdomen orange with dark central streak and ticks down back. Wings clear, but stained orange near bases. Eyes dark red.
Life history: Yellowish-white maggots feed in manure, slurry and dung. Adults visit flowers, but also torment stock by trying to sip liquid around the eyes and mouth.
Ecological notes: Will enter houses in autumn (hence scientific name), but only to hibernate, so not implicated in disease spread onto human foods. Does, however, spread eyeworm (nematode) and pinkeye (infectious conjunctivitis) in cattle. Walkers and ramblers through cattle pasture can be pestered by clouds of them flying around the head.
Bush fly, Musca vetustissima
Size: Length 6–7 mm, wingspan 13–15 mm.
Description: superficially very similar to house fly. Grey mottled fly, thorax with two broad darker streaks, abdomen greyish.
Life history: Australian. Breeds in moist decaying organic matter, and although not overly fond of native marsupial droppings because they were too dry, it thrived in the semi-liquid droppings of cows when these were introduced by European settlers.
Ecological notes: Particularly a problem in summer (October–March) when the air is thick with clouds of them. There are regular tales of travellers in the bush being unable to speak because immediately they opened their mouths in the swirling maelstrom, flies flew in. This gave rise to the stereotypical Aussie bush-hat with corks dangling on strings, and the ‘Australian salute’ flick of the hand across the face. It was partly to control bush fly numbers that African and Eurasian dung beetles were introduced into Australia in the mid to late 20th century.
House fly, Musca domestica
Size: Length 6–7 mm, wingspan 13–15 mm.
Description: Short and stout fly, grey, thorax with four darker streaks, abdomen with sides pale translucent yellow, legs dark grey to black.
Life history: Pale grey maggots feed in semi-liquid decaying organic matter – compost, manure, dung, carrion. Adult flies feed in the same breeding material, but then come into human houses and visit food left out. They eject enzyme juices, including contaminants from previous faecal encounters, from internal glands, then suck up digesting food.
Ecological notes: Long blamed for disease spread, directly by the method of eating by regurgitating crop contents from previous meal, but also by traipsing filth over human food. Reviled as ‘the insect menace’, houseflies were found to harbour more than 100 types of bacteria, virus and protozoa, and in one report over 6 million bacteria on a single fly. Advances in sanitation and food storage have improved the situation. Now much less common in towns and cities, still abundant in rural areas and really might be called the farmhouse fly.
Lesser house fly, Fannia canicularis
Size: Length 3–4 mm, wingspan 8–9 mm.
Description: Mottled grey fly with vaguely pale-streaked thorax, and yellow side markings on abdomen. Legs dark, wings clear.
Life history: Spiny wrinkled larvae live in almost any decaying organic matter, including compost, fungi, carrion and leaf litter in the soil. Adults come indoors and fly mad zigzags under pendent light fittings, but are not attracted to food and are not implicated in disease spread.
Ecological notes: Although this species is most often associated with old dung, where it may be feeding on fungal hyphae, the closely related F. scalaris has earned the name latrine fly for its very close association with human and other fresh dung. There are very many similar species with varying degrees of association with dung and manure.
Greenbottles, Lucilia species
Size: Length to 10 mm, wingspan 15–20 mm.
Description: Broad, stout, shining metallic green flies, sometimes with red, copper or gold tints, or appearing slightly bluish. Short legs dark. Large eyes dark red. Wings clear. Many similar species. The larger ‘dead dog fly’, Cynomya mortuorum, to 15 mm long, darker, bluer, bristlier.
Life history: Fat, pale-grey maggots feed in carrion or other decaying animal matter such as found in food leftovers dumped into compost heaps of domestic rubbish bins, or in commercial rubbish tips.
Ecological notes: Although not known to breed in dung, these attractive flies visit fresh dung to feed and can arrive within minutes of deposition. Large numbers can occur, creating a loud buzz when they are disturbed. Are attracted to meat-based human foods (fresh or cooked) so potential for disease transmission on the barbecue, picnic or open-air market stall, although not regular household visitor indoors.
Bluebottle, Calliphora vomitora
Size: Length 12 mm, wingspan, 20 mm.
Description: Large stout shining blue fly, verging on metallic, bristly.
Life history: Like its greenbottle cousins, larvae in decaying animal material.
Ecological notes: Likewise does not breed in dung, but visits fresh droppings to feed, often in loud buzzing hordes. Will come indoors, buzzing lazily about, and often called blow fly for its habit (before fridges invented) of laying eggs on stored meat, then said to be ‘fly-blown’. The blowing of its name may be derived from bloated (‘blowted’) carrion and decaying meat. External feeding by spitting out digestive juices onto food, then sucking this back up, echoed in its scientific name; visiting human food after dung-feeding thus not a pleasant thought.
Horn fly, Haematobia irritans
Size: Length 4 mm, wingspan 7 mm.
Description: Small grey compact fly reminiscent of house fly. Dark grey, with vague pattern of lighter mottles. Wings clear.
Life history: Eggs laid in very fresh dung. Requirement for blood-heat freshness of the dung has been invoked; sometimes flies said to lay eggs before defecation has finished. Pale larvae hatch almost immediately, feed-up quickly and take about 7 days to mature enough to pupate. Continually brooded until late autumn in temperate areas, when some pupae remain dormant until adult emergence in spring.
Ecological notes: Adult flies suck animal (but not human) blood and can reach nuisance proportions around stock animals, many thousands per cow, causing distress and lowered milk yields. The similar stable fly, Stomoxys calcitrans, will (painfully) bite humans; it breeds in stable litter and manure in temperate zones (overwintering as slow-developing larva), but directly in dung in open grassland in warmer regions.
Flesh flies, Sarcophaga species
Size: Length to 20 mm, wingspan to 22 mm.
Description: Large, stout, mottled grey and black flies, chequered pattern shifting with the light. Eyes bright red
Life history: Mostly carrion breeders (even small items such as dead snails and insects), requiring decaying animal matter for the maggots to feed, but some dung-breeding species are known throughout the world.
Ecological notes: Even if not breeding in the dung the adult flies often occur as casuals, resting on it, and presumably attracted by the same aromatic decay chemicals.
Horse bot fly, Gasterophilus intestinalis
Size: Adult fly length 18 mm, wing length 15 mm; larva to 20 mm.
Description: Adult flies are all-over orange-brown and grey-brown, furry, sometimes said to resemble bees. Wings clear with dark blotch in middle. Female fly with narrow tail, egg-laying tube, tucked underneath. Larva stout, cylindrical, pale orange-brown, with each segment edged with ring of spines.
Life history: In early summer flies lay slim white eggs (up to 1,000) on horse flanks, each glued to one of the host’s hairs. These are inadvertently swallowed when the horse licks itself. Larva attaches to the inside of the horse’s mouth, releasing and attaching to the stomach lining after about 4 weeks. The maggot eats the mucosal layer of the intestinal tract. When fully grown, during late winter or early spring, the larva releases its hold and is passed with the dung, in which it pupates.
Ecological notes: Not a dung feeder, but larvae (also pupae) are found in the dung. Adult flies have no functional mouthparts and do not (cannot) feed. Now a very rare insect in many areas where the valuable livestock are treated with systemic chemical drugs to rid them of bots, worms, warbles and other disease-status organisms.
Noon fly, Mesembrina meridiana
Size: Length to 12 mm, wingspan 23 mm.
Description: Large, shining, jet-black, very bristly fly. Wings clear, but brightly marked with orange at base and along fore-edge. Face and feet also bright orange.
Life history: Breeds in dung (mainly cow). Female lays only about five eggs in its lifetime, singly, often a day or two apart. Egg hatches quickly (1 hour), but if suitable pat is not found it hatches in the fly’s egg-laying tube and live larva is laid instead. Deep yellow maggot (largest fly larva in north European dung pats) has large jaws and will eat other fly grubs if they are available.
Ecological notes: A spectacular and handsome insect, often found sunning itself on tree trunks, leaves or fence posts, at the height of the day, hence common name. Also visits flowers. Given the chance the larva is fiercely predatory, but can survive on a dung-only diet.
Mottled dung fly, Polietes lardarius
Size: Length 10 mm, wingspan 18 mm.
Description: Medium-sized, stout, broad, grey fly, prettily marked with dark stripes down thorax and black bars across abdomen, creating a harlequin-like pattern that seems to shift as the light angle changes. Legs dark grey. Large eyes dark red.
Life history: Pale larvae live in dung, and are probably predatory on other fly maggots, at least when they are near fully grown. Adults rest of leaves and tree trunks as well as visiting dung.
Ecological notes: Several similar species. P. hirticrura ferociously predatory and recorded taking on much larger predatory larvae of Mesembrina (see above). These two are cow dung specialists, but other species apparently live only in horse dung.
Ground beetles, family Carabidae
Size: 3–30 mm.
Description: Very large group of diverse beetles, generally long oval, long legs, long antennae, flattened or cylindrical bodies, shining black or dark metallic.
Life history: Predatory as dark armoured larvae, and adults. Very active, fast running. Sometimes found under dung, and would not doubt take advantage of dung insect prey if they found it, but not really part of the dung fauna.
Ecological notes: Denizens of the grass thatch and have specially strengthened hind legs to wedge-push their way through the tight stems and roots, in pursuit of prey and to escape enemies. Often found under logs and stones, and treat dung as similar shelter.
Swimming dung beetle, Sphaeridium scarabaeoides
Size: Body length 5.5–7.5 mm.
Description: Shining, round, slightly domed beetle, black with reddish patches near shoulders and across tips of wing cases; legs broad and spined. Several very similar species.
Life history: Wrinkled, grey/brown, maggot-like larvae are predatory, attacking and eating other small dung- and soil-dwelling insects and their larvae.
Ecological notes: Flattened, rounded and smooth outline and broad paddle-like legs allow it to ‘swim’ through fresh semi-liquid cow dung. Flying new arrivals to the runny pat land and break straight through the thin skin-like rind forming in the sunshine. Related beetles in this family (Hydrophilidae) live in water, or mud.
Scavenger dung beetles, Cercyon and other genera
Size: 1.2–4.5 mm.
Description: Minute to very small, highly convex, domed, shiny beetles. Usually black, dark brown or dirty red, but sometimes marked with brighter red or yellow. Legs and antennae short. Several similar genera include Cryptopleurum, which is slightly hairy under the microscope, and Megasternum which has very broad, deeply notched front legs.
Life history: Various species breed in decaying organic matter of all types, including compost, carrion, rotting fungi, pond edges and seaweed strandlines; a few are more or less confined to dung and manure.
Ecological notes: Large numbers of many different species can inhabit the same pat. Many species in the same family (Hydrophilidae) are aquatic and the smooth lines and broad short legs of Cercyon specimens allows them to push right into soft dung.
Minute dung beetle, Ootypus globosus
Size: 1.0–1.5 mm.
Description: Minute, convex, domed, almost hemispherical, very shiny beetle; black to dark reddish chestnut brown. Legs short and slim.
Life history: Little is known other than it breeds in dung and other rancid decaying organic matter. Several extremely similar species, including Atomaria, also named for their small size.
Ecological notes: Just about the smallest dung beetle to be found. Probably a fungus feeder, eating the fungal hyphae and grazing mould and spores.
Clown beetle, Hister unicolor (and other species)
Size: Length 8 mm.
Description: Broad, round, domed, smooth, shining black, heavily armoured beetle. Antennae short, clubbed. Legs broad, armed with teeth, flattened, and which can be pulled in and recessed into grooves on the underside of the body. Many similar species, some marked with vague red blotches.
Life history: Adults and larvae are predatory on other dung-inhabiting invertebrate larvae, notably fly maggots.
Ecological notes: Not common, so play only a minor role in dung ecology, but distinctive and attractive beetles. Many similar species, some marked with reddish blotches, some metallic greenish or bronze. Also found in carrion, rotting fungi, decaying plant material. Sadly the obviously dung-named Margarinotus merdarius is mostly a species of compost and manure heaps, or bird nests in tree hollows. Smaller and shinier Hypocaccus species usually confined to sand dunes, where dog dung is only available food. Common name is of dubious etymology, either Latin hister meaning dirty lowly being, or histrio (from which also histrionics) meaning actor.
Ridged clown beetle, Onthophilus striatus
Size: Length 1.8–2.4 mm.
Description: Tiny, globular, dull black beetle. Wing cases each with three strong and three smaller raised ridges. Thorax with six raised ridges. Legs and antennae slim.
Life history: Predator of other even smaller dung invertebrates, particularly fly larvae.
Ecological notes: Seemingly only ever in horse dung, but closely related, larger (2.5–3.5 mm) O. punctatus, with five thorax ridges, in mole nests, fox dens or badger setts.
Feather-winged beetles, family Ptiliidae
Size: 0.5–1.2 mm.
Description: Microscopic black or brown beetles, short oval, some with wing-cases shortened, exposing the tail of the abdomen. Antennae long and very thin with rings of long hairs.
Life history: Feeders on fungal hyphae and spores.
Ecological notes: Found in old dung where powdery mildew fungi and moulds have started to grow. Large number of extremely similar species. Also found in leaf litter, rotten wood, fungi, animal nests, compost, mouldy hay, ant nests, decayed seaweed. Ironically named Nephanes titan (0.55–0.65 mm) one of many frequently described by entomologists in terms of ‘the size of the full-stop at the end of this sentence’. Dung associations broad, but sometimes blurred; Euryptilium gillmeisteri known (in UK) only from three specimens found in leaf litter infected with bird droppings at base of large oak tree.
Ridged rove beetle, Micropeplus porcatus
Size: Length 2.5–3.0 mm.
Description: Short, broad, flat, dull brown rove beetle. Wing cases each bearing three sharp ridges, short, exposing four or five abdominal segments. Thorax strongly sculptured. Legs and antennae very slim. Slow moving. Several very similar species.
Life history: Feeds on mould and fungal hyphae in decaying organic material.
Ecological notes: In old dung, also in manure heaps, rotten straw, mouldy hay, compost, leaf litter and other dry, decaying organic matter. Also occurs in mud at the side of lakes and streams, suggesting it feeds on liquid decay as well as dry fungoid rot.
Impossible rove beetles, Atheta species
Size: Length 1.1–5.0 mm.
Description: Minute to very small, slim, black or brown rove beetles. Legs and antennae slim.
Life history: Very little is known about their habits. Some species are known to eat free-living nematode worms, so predation of micro-organisms and other soft-bodied invertebrates seems likely.
Ecological notes: Occur in a wide variety of decaying organic matter with several specifically in dung. Huge number of extremely similar species, which require expert knowledge and detailed examination to identify. Most identifications rely on dissection and special mounting of internal genitalia, notably the female’s sperm-storage organ (spermathecal). Males of some species unknown or not possible to identify. That’s not really their English name, just one I have coined in my own frustration at finding them so impossible to identify correctly.
Flat rove beetles, Megarthrus species
Size: Length 2.5–3.5 mm.
Description: Short, very broad, flat, brown or black rove beetles. Hind corners of thorax incised with small notch. Wing-cases long (for a rove beetle), about as long as, or slightly longer than hind body.
Life history: Both adults and larvae are predators of smaller invertebrates.
Ecological notes: In dung, manure or other decaying organic matter. Many similar genera.
Sculptured rove beetles, Platystethus, Anotylus, Oxytelus and other genera
Size: 1.2–5.5 mm.
Description: Numerous group of slightly flattened, black or dark brown rove beetles, sometimes smudged with pitchy-reddish blotches. Very parallel-sided with broad heads, broad rectangular thorax usually ridged, dimpled or grooved to give sculptured appearance.
Life history: All feed as larvae in decaying organic matter, including dung, manure, compost, rotting seaweed, putrescent fungi, haystack refuse, leaf litter, and moss.
Ecological notes: Slow-moving under dung. Spiny legs imply they burrow in the dung or soil beneath. Other closely related groups with similar broad, shovel-shaped front legs are confirmed soil-diggers.
Pride of Kent, Emus hirtus
Size: Length 18–28 mm.
Description: Large, broad, powerful, spectacular rove beetle. Black, but head, part of thorax and tail tip covered with bright yellow hairs. Rear half of wing-cases thick with grey fur. Legs short and stout. Jaws large.
Life history: Predator of other invertebrate inhabitants of cow dung.
Ecological notes: Extremely rare in the UK, with most records from the grazing marshes of north Kent. There was much excitement when it was rediscovered, after many years, running on the tarmac near a public lavatory on a Kent nature reserve. Usually associated with very fresh cow pats, on which it scurries in frantic activity, attacking the flies which are attracted. Possibly a bumblebee mimic when flying. Note: Creophilus maxillosus is another large, broad, furry rove beetle, but is patterned with pale grey hairs, not yellow; it occurs in carrion rather than dung, and is sometimes misidentified as Emus by excited, but inexperienced, observers.
Mottled rove beetle, Ontholestes tessellatus
Size: Length 14–19 mm.
Description: Large, stout, rove beetle. Black, but forebody (head, thorax, wing-cases and first segment or two of abdomen) patterned with golden-green hairs to give a shifting chequered appearance. Small patch of matt black hairs at join of thorax and wing-cases. Tail black, marked with grey patches. Legs reddish golden. Smaller (10–15 mm) and narrower, O. murinus (right) is less golden, with legs black.
Life history: Predator of flies and their larvae. Also occurs on carrion.
Ecological notes: Very fast and active insect. Despite its silky appearance it copes with sticky dung very well, and grooms itself using front legs on head and antennae, middle legs for thorax and wing-cases, and rear legs for tail segments.
Red-girdled rove beetle, Platydracus stercorarius
Size: Length 12–13 mm.
Description: Black, not very shining, but wing-cases strongly contrasting bright orange-red; abdominal segments decorated with bands of silver hair, which catch the light as the animal runs; legs and antenna red. Several similar species, some with wing-cases black, or all-over mottled with vague silvery grey pubescence.
Life history: Predatory as both larva and adult.
Ecological notes: Occurs under dung, but also under carrion, in manure and compost heaps. Very agile and active.
Parasitoid rove beetles, Aleochara species
Size: Length 1.5–10 mm.
Description: Relatively broad, stout, black, brown or pitchy-red rove beetles, usually shining; sometimes wing-cases marked with reddish blotches. Wing-cases seeming very short compared to long hind body.
Life history: Very large and diverse group, which are specialist predators of fly puparia; feeding completely inside the pupa, they may really be described as parasitoids.
Ecological notes: In all types of dung, but also in carrion and other putrescent decay. When fully grown, larva pupates inside the almost empty fly puparium. Adult beetle emerges through a jagged hole it chews in the puparium shell. This contrasts with the usually perfectly round hole through which parasitic wasps emerge.
Japanese rove beetle, Philonthus spinipes
Size: Length 13–17 mm.
Description: Head and thorax shining black; wing-cases orange-red, covered with red pubescence; hind body iridescent black. Antennae black. Bristly legs black at base, and orange from knees.
Life history: Predator as both larva and adult. Very fast and active.
Ecological notes: Particularly in horse dung. First described (1874) from Japan, but spreading through Asia and Europe in late 20th century. Arrived in UK in 1997. Reason for range expansion unknown.
Splendid rove beetle, Philonthus splendens
Size: Length 12–14 mm.
Description: Shining black parallel-sided rove beetle; head and thorax with slight metallic bronze sheen, wing-cases more strongly brassy green metallic.
Life history: Predator as both larva and adult, mostly on flies and their maggots.
Ecological notes: Very fast and active predator of other small invertebrates in dung, also manure heaps, carrion, putrid fungi. Very many similar species requiring expert knowledge to identify. Flies readily and can land, fold away wings in an instant and vanish under fresh dung in a blur.
Devil’s coach-horse, Ocypus olens
Size: Length 20–28 mm.
Description: Huge, broad, dull black rove beetle. Entire body surface covered with dense array of minute pinprick dints and short black pubescence, giving a matt appearance.
Life history: Predator as both larva and adult. Commonly under stones, logs, clods of earth, in compost and manure heaps. Not a specialist dung species, but regularly found treating old dung as suitable shelter.
Ecological notes: Often found running on paths, and familiar to gardeners and many non-entomologists. If threatened will rear up head, displaying its large jaws (can nip), and tail, exuding droplet of smelly liquid from its tip.
Smooth rove beetles, Tachinus species
Size: Length 3–10 mm.
Description: Elegant, smooth, shining black, brown or reddish rove beetles. Hind body strongly narrowed behind, last segment deeply notched, or extended at sides into long teeth. Many similar species. Note: closely related and hugely abundant genus Tachyporus specimens are smaller (2–4 mm), and occur in grass thatch.
Life history: Predator as larva and adult.
Ecological notes: In rancid decaying matter such as carrion, manure, putrid fungi, compost heaps and dung.
Burying beetles, Nicrophorus (formerly Necrophorus) species
Size: Length 10–30 mm.
Description: Large, stout, heavily built, parallel-sided beetles, generally black with bright orange bars (broken or solid) across wing-cases; some species all black, others orange on thorax.
Life history: Carrion feeders, mostly of small carcasses such as voles or birds. Dig out earth from underneath creating a void into which the body subsides, and spoil from which eventually engulfs it. Then feeds on putrescent decay, and lays eggs. Male and female generally work together.
Ecological notes: Not really a dung feeder, but often found under carnivore dung if stool still contains enough decaying meat material to give off the right scents. Unlikely to result in interment, egg-laying or breeding though.
Hide beetles, Trox scaber (and other species)
Size: Length 5–7 mm.
Description: Stout, blunt, oval, domed beetle, body roughly sculpted with raised tubercles and covered with short upstanding scales or broad stiff hairs, giving a slight bristly appearance, and sometimes obscuring the beetle by accumulating a covering of dust and debris.
Life history: Mostly carrion feeders, appearing at the dry tendon-and-fur stage after putrescence has passed.
Ecological notes: Not really a dung feeder, but sometimes found under carnivore scats, especially if they are rich in fur, feathers or hair.
Dumbledor or dor beetle, Geotrupes stercorarius (and other species)
Size: Body length 16–26 mm.
Description: Massive, convex, broad domed, shining black beetle, with purplish or bluish tints, stout powerful toothed legs, antennae clubbed.
Life history: Adults dig deep burrows into the soil, provisioning side tunnels with dung balls in which eggs are laid. Fat, pale whitish C-shaped grubs feed in these dung cells and emerge as adults, pushing up through soil, weeks or months later (depending on temperature). Some species also use decaying leaf litter instead of dung.
Ecological notes: Important dung recyclers because of their size. Despite heavy build, they are good fliers across meadows on warm evenings, giving off loud buzzing whirr, leading to the onomatopoeic English names; also known as lousy watchman (often found carrying louse-like mites, watchman is reference to ‘clock’, an old name for any large beetle). Several different species distinguished by size, colour hints, leg tooth shape and smoothness or ridge numbers on wing cases.
Minotaur beetle, Typhaeus typhoeus
Size: Body length 14–22 mm.
Description: Stout, convex, broad domed, shining black beetle with broad, toothed legs and clubbed antennae. Male with three sharp spines projecting over head from front of thorax, usually side ones longer, but varying greatly; female with slight bumps instead.
Life history: Like dor beetle, buries dung balls in a tunnel dug down in the soil, on which eggs are laid.
Ecological notes: Prefers sandy soil where burrows can be up to 2 m long; mostly on rabbit or sheep dung. Drags, rather than rolls, the small pellets. Overwinters as an adult in the burrow so is sometimes active on mild winter days. Attracted to lighted windows and porches, and regularly found in moth light traps, even in early February.
Lesser scarab, Odonteus armiger (formerly Odontaeus mobilicornis)
Size: Body length 6–8 mm.
Description: Short, round, almost globular, smooth, shining, dark brownish black, legs reddish. Male with long backwards curving moveable spine on its head, female with small prominences.
Life history: Usually found flying on warm summer evenings. Thought to feed on subterranean fungi, and excavates burrows to 40 cm deep.
Ecological notes: Reported from dung in some of the old books, and possibly associated with buried dung on which fungi develop. Closer association with rabbit warrens has been suggested, but evidence is flimsy.
Common dwelling dung beetles, Aphodius species
Size: Length 2.5–15 mm.
Description: Stout, semi-cylindrical long oval beetles, with wing-cases distinctively rounded at tail end, broad domed thorax and blunt shovel-shaped head. Without horns or prominences on head or thorax. Very many different species. Typical common European species include: A. fimetarius (6–8 mm), jet black but wing-cases and front corners of thorax bright red; A. contaminatus (5–6 mm) wing-cases straw coloured, flecked with black marks in vague chevron shape; A. sticticus similar but shinier, markings crisper; A. rufipes (11–13 mm) long, parallel-sided, deep chestnut red-brown; A. fossor (9–12 mm) thickset, black, shining; A. prodromus (4–7 mm) wing-cases dirty yellow, with large darker smudge at sides and end; A. erraticus (6–9 mm) wing-cases dull beige, very broad; A. haemorrhoidalis (3–4 mm) all black, wing-cases with reddish tip.
Life history: Fly readily to colonise dung, burrow into and beneath the dropping, where they lay their eggs. Huge numbers and/or multiple species can occur in a single pat.
Ecological notes: In temperate regions this is the most important and numerous dung beetle genus. Despite their ubiquity they have no widely accepted common name, so I’ve coined this one since they are the dominant ‘dweller’ beetles (as opposed to tunnellers or rollers). Some species supposedly more associated with dung of particular animals. There is some evidence that moisture content is important so some species cope better with wet cow dung, others are more at home in drier sheep or rabbit pellets, but most are fairly broad in tastes; thus deer dung species more likely to be just adapted to woodland shade rather than stool content.
A. fossor
A. prodromus
A. nitidulus
A. varians
A. haemorrhoidalis
A. sticticus
A. paykulli
Common tunnelling dung beetles, Onthophagus species
Size: Length 3–14 mm.
Description: Short, compact, oval convex beetles, head and thorax about equal in size to abdomen covered with wing-cases. Legs flat and broad, armed with large teeth. Variously coloured black, yellowish, brown, reddish; wing-cases sometimes mottled or blotched, matt to polished shining black, metallic green or bronze. Males often with pronounced, sometimes bizarre, horns and spines from head or front of thorax. Widespread (but never common) European species include: O. joannae (4–6 mm) all dull black; O. taurus above right (8–12 mm) all shining black or dark brown, male head with two long curved horns reaching back over sides of thorax; O. coenobita (6–10 mm) head and thorax dull dark metallic green, wing-cases pale brown, usually flecked with vague darker blotches; O. similis bottom (4–7 mm) thorax black, wing-cases dark beige with chequer-like pattern of black speckles.
Life history: Digs burrow under the dung and removes pellets or boluses down into the blind end, on which to lay eggs.
Ecological notes: Hugely varied and important genus of beetles, and the dominant tunnelling genus through most of the world. Some of the horns are truly spectacular and the basis of studies trying to understand how and why these decorations have evolved and developed. Head-to-head pushing and shoving contests between males may take place in the tunnels. Three Australian species have prehensile claws to grip fur around wallaby anus until dung is dropped, then release hold and bury pellet whole.
English scarab, Copris lunaris
Size: Body length 17–23 mm.
Description: Stout, domed, shining black beetle, but with sharp ridge along cut-in front edge of thorax, and head with long (male) or short (female) spine projecting up from broad, rounded head.
Life history: Like dor beetle, buries dung balls in a tunnel dug in the soil, on which eggs are laid.
Ecological notes: Usually on cow dung and said to prefer sandy or chalky places. Extremely rare in Britain, known in UK only from a handful of sites in south-east England and likely to be extinct; not reliably recorded since 1950s. ‘English’ scarab after conservation exercise in 1990s meant priority species were all given common names. Widespread and sometimes common in Europe.
Tumblebugs, Canthon pilularius (formerly C. laevis) and other species
Size: Length 10–20 mm.
Description: North America. Broad, smooth, dull black beetle, with coppery tinge or (south-western colonies) brighter blue or green sheen. Head broad, strongly rounded and flattened. Front legs broadened with three deep teeth; middle and back legs longer and slimmer.
Life history: Flies in low zigzags upwind to fresh dung. Landing, it immediately starts to cut away a lump of dung to shape it into a ball. Balls are roughly 30 mm across and take about 20 minutes to sculpt. Sometimes the ball is rotated in the dry ground to give it a coating of sand. The ball is then rolled away from the pat, to be buried.
Ecological notes: Rolling is by the beetle pushing, head down, standing on its front legs and using its middle and back legs to manoeuvre the ball away. Quite an ungainly animal, by all accounts, frequently tumbling and losing its grip on the ball, which rolls away and has to be searched for, antennae outstretched. Route away from the pat not very straight, often zigzag.
Perching dung beetle, Canthon viridis
Size: Length 2–5 mm.
Description: North and Central America. Small, round, stout, globular, dung beetle; shining metallic purple, bronze (northern) or bright glossy green (southern). Head broad, flat, rounded. Legs slim. Several similar species.
Life history: Breeds in dung of woodland animals such as deer, monkeys, peccaries; also in rotting fungi and carrion.
Ecological notes: A woodland or forest species. Noted for its perching behaviour, on leaves and stems in the dappled light of often dense treescapes, its round little body upright on its slim legs and with its antennae outstretched, three flat terminal club segments fanned out. Possibly waiting to catch scent of freshly dropped dung, or waiting for dangerous predatory rove beetles to leave the dung.
Carolina scarab, Dichotomius carolinus
Size: 20–30 mm.
Description: North America. Large, heavy, broad, stout dung beetle; shining black, but underside fringed with brown or orange hairs. Legs black, antennae orange-brown. Wing-cases deeply grooved. Thorax with heavy jutting shoulder across front. Head large, broad, rounded, almost flanged.
Life history: Breeds in dung of horses, cows, deer, digging a tunnel 30–40 cm deep into the soil and packing its end with dung fragments to make a food store or brood ball. Single white egg laid on brood mass. Generation time about 2 months. Large mounds of excavated soil mark the burrow entrances.
Ecological notes: Nocturnal, and often attracted to lighted porches or verandas during the summer. Thought originally a forest species, of open clearings, but regularly found in grazing pastures abutting woodland. One of the largest and heaviest dung beetles in North America. So powerful it is difficult to hold in a clenched fist.
Rainbow scarab, Phanaeus vindex
Size: Length 11–22 mm.
Description: North America. Beautiful and striking brilliant metallic green and gold, sometimes blue or black, dung beetle; broad, almost square. Thorax flattened and produced into two broad backward sloping, tooth-like lobes. Head of male armed with huge, backwardly curved horn. Whole body covered in wrinkled, dimpled sculpture. Legs broad and stout. Several similar species.
Life history: Tunneller, digging burrow under the dung and throwing up spoil heap beside the pat. Roughly pear-shaped dung ball is placed at the end of the burrow. Side tunnels may be created to accommodate more brood masses. Single egg on each mass.
Ecological notes: Widespread and often common; in wide variety of dung including pig, opossum, dog, cow, horse and human.
Giant African dung beetle, Heliocopris gigas
Size: Length 37–60 mm.
Description: Africa. Very large and stout, domed, broad, almost square black or very dark brown dung beetle. Black, slightly shining, underside with fringing of brown hairs. Legs broad, especially front pair, flattened and armed with large teeth. Head very large, broad, round, spade-shaped in female, armed with two broad twisted back-swept horns in male. Thorax with nearly vertical cliff-like front, produced, in males, into sharp spines at sides and a large prong jutting over the head. All over dimpled and wrinkled to give tough leather-like appearance
Life history: Breeds in elephant and other dungs, digging a tunnel beneath the pat and stocking the ends of the burrows with brood balls, each about 5 cm in diameter, on which single eggs are laid.
Ecological notes: Although once thought to breed only in elephant dung, it occurs widely in the elephant-free Arabian Peninsula, and will use droppings from wild animals, also camel, cow and horse.
Sacred scarab, Scarabaeus sacer
Size: Length 26–40 mm.
Description: Southern Mediterranean Europe and North Africa, Indus Valley. Very large and stout, broad, flat, short oval dung beetle. Black, moderately shining. Head flattened, armed with six tooth-like projections. Legs relatively long; middle and hind pairs slim, front pair with each leg armed with four broad teeth. Several similar species.
Life history: Scoops balls of dung using its broad head and rake-like front legs, then rolls these balls backwards, head down, away from the pat some distance (often several metres) before burying them in the soil. Each brood ball with a single egg laid in it.
Ecological notes: Uses the sun (or moon, or indeed Milky Way) to orient its journey so that even though it does not look where it is going, it can successfully roll away the dung ball in a straight line, despite uneven ground, ridges, boulders or log barriers which would otherwise interrupt its path.
Egyptian scarab, Kheper aegyptiorum (and other species)
Size: 25–40 mm.
Description: North-east Africa. Very large, broad, flattened, short oval dung beetle. Black, moderately shining, often with vague metallic bronze tinge. Flat head armed with six projections. Front legs similarly armed with flat blade-like teeth.
Life history: Like Scarabaeus (it is sometimes also included in this genus) rolls large ball of dung away from the main pat, and buries these in the soil some distance away.
Ecological notes: Active during the day, where many African dung beetles are semi-nocturnal. Once a male has shaped a dung ball and moved it a short distance from the main pat, it adopts a head-stand pose, angled at about 45°, on top. It is releasing a nuptial pheromone from the underside of its abdomen, pumped into the air by simultaneous contractions of its back legs. This attracts a female and the two beetles work together to bury the dung ball, before the female lays an egg in it.
Gracile dung beetle, Sisyphus (now sometimes Neosisyphus) mirabilis
Size: Length to 12 mm.
Description: South Africa. Body small oval, dull black; head broad, rounded, paddle-shaped. Legs very long, especially hind pair which are armed with various spines and pegs. Middle legs each armed with strong curved spine. Front legs short, stout, armed with small teeth. Several similar species worldwide, though not all as long-legged.
Life history: A dung roller, digging out a marble-sized ball of dung and rolling it off at high speed away from the pat, to a safe place where it can be buried in the soil and an egg laid.
Ecological notes: It rolls by running on its front legs, head down, tail in the air, using its hooked middle legs and long back legs to keep control of the ball it is pushing. In Greek mythology, Sisyphus was the conceited king of Ephyra (Corinth), condemned to an eternity of rolling a large boulder up a hill, only to have it roll away from him near the top so he would have to start all over again.
Sloth dung beetle, Pedaridium (formerly Trichillum) bradyporum
Size: Length 3 mm.
Description: South and Central America. Small short oval, domed, very convex dung beetle. Black or pitchy with reddish brown tints, verging on metallic. Upper surface covered all over with short, widely spaced, but distinct upstanding hairs. Legs short.
Life history: Associated with three-toed sloths (Bradypus species), in the fur of which they lodge (nearly 1,000 recorded from one individual). When the host descends to the forest floor to defecate (they bury their dung to hide their presence from predators), the beetles drop off and lay their eggs.
Ecological notes: Several similar species are recorded from cow and human dung, but close association with any particular hosts is unknown or unrecorded.
Three-striped chafer, Macroma (sometimes Campsiura) trivittata
Size: 20–23 mm.
Description: Africa. Large broad, smooth straw-yellow and black chafer beetle. Varyingly coloured from yellow to reddish, and variously marked, but usually with three black longitudinal stripes on thorax, small black tick marks on shoulders and jagged black bar across near tip of wing-cases. Legs and antennae reddish.
Life history: Fat C-shaped grub lives in leaf-litter and soil, eating decaying plant material. Adults fly actively.
Ecological notes: Has been recorded in Ivory Coast breeding in elephant dung. Unlike in the open savannahs, there appears to be less competition for dung in the tropical rainforest hereabouts and elephant dung remains in large quantities for secondary users.
Cave larder beetle, Dermestes carnivorus
Size: length 6.5–7.5 mm.
Description: Small, long-oval beetle. Black, upperside with scattering of brownish scales, underside with dense blanket of silver-white scale-like hairs. Legs medium length. Antennae brown, clubbed.
Life history: Active bristly larvae are scavengers. Original habitat probably carrion, at its dried stage where only bone, sinew, fur and feather remain. Thought to have originated in neotropics, but after invading human homes as larder pest (in stored meat products) has been transported across the globe.
Ecological notes: Major decomposer of bat guano in caves. Will probably also feed on dead bats. As well as making jump to human habitations to feed on spilled food, has been recorded in bat droppings in abandoned buildings.
Shining spider beetle, Gibbium aequinoctiale
Size: Length 1.5–3.5 mm.
Description: Small, shining globular beetle. Brown to beige, wing-cases usually glossy chestnut. Abdomen nearly spherical, humped, head hidden under narrow thorax. Legs relatively long, giving spider-like appearance.
Life history: A scavenger species, part of a large group of similar species that feed on spilled food in houses and warehouses, or in seeds and nuts cached by wild animals or dropped inside their nests.
Ecological notes: Completely bizarrely found 800 m underground in Yorkshire coal-mines, feeding on human dung. No toilet facilities being available in the mine workings, miners allocate disused tunnels as unofficial latrines. With no ‘natural’ dung fauna to bury or recycle it, the excrement remains, but dries out into a cake-like consistency. The beetle is flightless, lacking wings, and with its wing-cases fused together; how it got down the mine (and others in Durham and Staffordshire) is still a mystery.
Click beetles, Agriotes species
Size: Length 4–9 mm.
Description: Small, smooth-outlined, long-oval parallel-sided, almost cylindrical beetles; brown, greyish black to dull chestnut. Legs slim, antennae long. Many similar species.
Life history: Soil-dwelling larvae (called wireworms by gardeners) are long, cylindrical, pale yellowish-orange with brighter orange, brown or black head capsule. They feed in the soil, partially predatory, but also eat plant roots, and are sometimes regarded as horticultural or agricultural pests.
Ecological notes: Not really dung feeders, but larvae regularly invade the blurred soil horizon under old dung; adults less so. Adult beetle can make audible click by sudden jackknifing of body, which propels it into the air, or out of harm’s way, if attacked or disturbed.
Lesser mealworm, Alphitobius diaperinus
Size: Length 5–6 mm.
Description: Long oval, smooth, glossy, black or dark brown beetle, sometimes with reddish tinge. Legs and antennae same colour as body.
Life history: A scavenger of dry material, including fungoid wood, stored food products such as wheat, barley, beans, tobacco and dried meat. Has been transported around the globe and can be serious domestic or commercial pest.
Ecological notes: Often found in caves, feeding primarily on dry bat guano. It is also found in chicken houses, where it may be feeding on droppings.
Dung weevil, Tentegia ingrata
Size: Length 8–12 mm.
Description: Australia. Dumpy, hunched weevil with bulbous thorax, wing-cases ridged or knobbled, dark, sometimes black, sparsely flecked with small paler scales. Snout long and narrow. Legs relatively long.
Life history: Adult beetle awkwardly manoeuvres pellets of wallaby or kangaroo dung from dry grassland several metres to small log, under which it caches them. It makes a small hole in the soil, or uses already existing cavity, to store the dropping, then lays an egg in each pellet. Very similar T. bisignata has been reared from pellets of possum dung found hidden in similar caches. It is unclear why the unfortunately named T. stupida is so called.
Ecological notes: The only known dung-feeding genus of weevil. Australian marsupial droppings are dry, so the weevils treat it as merely preprocessed plant material. Ordinarily female weevils use their long snout, with small biting jaws right at the end, to chew a deep drill hole into leaf, stem, seed, then turn round and use a telescopic ovipositor to lay egg deep inside. In this case it seems that they chew into the dung nugget to make an egg tunnel.
LEPIDOPTERA – BUTTERFLIES AND MOTHS
Butterflies, various species
Size: Wingspan 25–105 mm.
Description: Unmistakable. Distinctive colourful day-flying insects. Huge variety of different sizes, shapes, colours and patterns.
Life history: Mostly plant-feeders as caterpillars (with chewing mouthparts), taking weeks or months to achieve bulk enough to pupate and transform into adult. Adults have long tongue for sipping nectar from flowers.
Ecological notes: Not really dung feeders, but regular visitors (as adults) to fresh droppings where they sit and drink from the moist surface. In tropical countries muddy banks of streams and ponds, where animals drink and defecate, can be ablaze with scores, even hundreds, of butterflies, often a mixture of different species. This ‘puddling’ behaviour likely supplies minerals, salts and other micronutrients.
Sloth moth, Cryptoses choloepi (and other species)
Size: Wing length 8–11 mm.
Description: South and Central Americas. Small grey-brown moth, wings narrow, held tight furled to body when at rest. Forewings brownish or purplish grey, vaguely streaked with dirty cream marks. Hind wings uniform pale grey.
Life history: Lives in the fur of sloths (mainly three-toed, Bradypus species). Once a week sloth descends tree, scrapes small hollow with vestigial tail, defecates and covers dropping with leaf litter. Female moths fly down and lay eggs in the dung. Larvae feed in the dropping. Emerge some weeks later and fly up into canopy to find new sloth host.
Ecological notes: Not a commensal (minor non-damaging parasite) – sloth and moth are mutualists; both derive advantage. Moths get readily available food supply for their larvae. Sloths also benefit from moths, which bring biomass into the fur where green algae grow. This camouflages the slow-moving sloths in the canopy. They groom themselves and derive significant nutrients from eating the algal bloom in their fur.
Owl pellet moth (tapestry moth), Trichophaga tapetzella
Size: Length 5–9 mm, wingspan 13–22 mm.
Description: Small grey and white moth; generally mottled, but basal third of wings darker, brownish or purplish grey. Wings held furled tight against body when at rest. Bird-dropping mimic.
Life history: Small pale caterpillar feeds in a sock-like case woven of its own silk, which it carries about as it crawls. Lives in bird and animal nests, feeding on moulted fur and feathers.
Ecological notes: Sometime minor household pest, eating woollen fabrics (including tapestries), rugs, horse hair furniture stuffing, pillow feathers. Regularly reared from owl pellets, where the larvae feed on the undigested feather and fur remains coughed up. Strictly speaking these are not dung, but droppings in a much broader sense.
DICTYOPTERA – TERMITES AND COCKROACHES
Termites, Microtermes, Odontotermes, Macrotermes, Synacanthotermes and others
Size: Length 1–15 mm.
Description: Small pale creamy or yellowish-brown insects. Head relatively large on soft, narrow, short-legged body. Sometimes called white ants for their vague resemblance, large numbers and nest formation, but are unrelated.
Life history: Complex colony of fertile male and female and many thousands of sterile females (workers), of different shapes and sizes adapted to tasks of foraging, tending young, nest building, fighting. Build large nests of soil mixed with saliva to make tough concrete, elaborate structures to allow ventilation and food storage, collect plant material. Can digest cellulose because of gut micro-organisms. Some species culture fungus on stored leaf fragments in the nest and eat fruiting bodies.
Ecological notes: Only considered a dung-feeder since 1970s when they were discovered to be major recyclers of dung during African dry season when dung beetles less active. Removes remains of elephant, cow, camel and many other types of dung, treating the quickly drying droppings as merely part-processed plant particles.
Cave cockroaches, guanobies, Trogoblatella, Spelaeoblatta, Eublaberus and other species
Size: Length up to 50 mm.
Description: Broad, flat, long oval insects, with long legs and very long, many-segmented antennae. Some species are winged, with leathery overlapping wing-cases, although these are sometimes shortened, and many flightless species lack them completely. Tip of abdomen with cerci – short antenna-like feelers.
Life history: General scavengers, eating whatever comes their way. Eggs laid in ootheca, a tough case made of hardened protein material secreted around the eggs inside the female abdomen.
Ecological notes: Cockroaches are a major component of cave faunas around the world, where bat droppings accumulate on the cave floors. They feed on the guano, on moulds, dead insects, dead bats and anything else. Some cockroaches have become domestic pests across the world. These started off as generalist detritivores in tropical leaf-litter rather than as cave species, but their sometime attraction to sewers and human latrines makes their kitchen-visiting habits unhygienic and unhealthy.
Social wasp, yellowjacket, Vespula vulgaris and other species
Size: Length 20 mm, wingspan 40 mm.
Description: Distinctive long, narrow, strikingly marked, black and yellow insects, with notable narrow ‘waist’ between thorax and abdomen. Head with large kidney-shaped eyes, stout antennae, yellow face and sharp jaws. Wings (large pair plus smaller pair) clear, membranous, folded into pleats when at rest.
Life history: Complex colony formation with single fertile female (queen) laying eggs and up to several thousands of sterile females (workers) foraging and nest building. Late in season new males and queens are produced which mate. Mostly predators of small insects, chewed bits of which are fed to the grubs in the nest combs, but some flower visiting.
Ecological notes: Not strictly dung feeders, but will visit carrion and other putrescent decay to feed. Sometimes sits in wait by a dropping to attack the dung flies and dung beetles that are attracted.
Scarab wasps, Tiphia femorata, T. minuta and others
Size: Length to 12 mm, wing length to 9 mm.
Description: Narrow, black, shining wasp-like, with narrow waist. Legs red or black.
Life history: Burrows into the soil (mostly sandy places) to find find dung beetle larvae or pupae, on which it lays its eggs. The maggots then devour the host grubs.
Ecological notes: Not strictly a dung-feeder, but definitely part of the dung community. Not generally associated with fresh dung, but a latecomer, after the dung has been partly or wholly removed.
Digger wasp, Mellinus arvensis (and other species)
Size: Length 15 mm.
Description: Shining, narrow, black wasp, prettily marked with bright yellow flashes on head and thorax, and bars across slightly bulbous abdomen. Legs yellow. Wings, slightly brownish.
Life history: Captures and kills insect prey to stock a small burrow nest made usually in sandy ground. Egg laid in each prey-stocked cell, maggot eats store and emerges the following year. Each female works alone, but many gather in loose aggregations in suitable bare ground to form a ‘wasp village’.
Ecological notes: Not a dung-feeder, but regularly observed sitting on fresh dung waiting to pounce on the blowflies, greenbottles and dung flies, often larger than itself, which are attracted. Mad scramble at the pounce, uses formidable jaws and sting to subdue its prey.
Parasitoid wasps, families Pteromalidae, Braconidae, Ichneumonidae, etc.
Size: Length 0.5–35 mm.
Description: Vast array of minute to large, slim, usually dark-bodied, sometimes metallic, wasp-like insects. Wings, one large pair, one smaller pair behind, usually clear membranous. Smaller species run-hop, larger species hawk low, quartering the ground looking for host victims.
Life history: Internal parasitoids of other insects. Eggs are laid direct into insect eggs, larvae or pupae and the hatching maggots eat the host alive, from the inside, finally killing it. Very often host-specific, each parasitoid species only attacking one particular host species, genus or close-knit guild of organisms.
Ecological notes: A ubiquitous, but poorly studied group of insects, despite many being large and brightly coloured. Virtually every insect in the world will have its own particular parasitoids. Many dung-feeding beetles, flies and other insects have their own parasitoid species attacking them, but rearing records are pitifully few.
Ants, family Formicidae
Size: 2.5–25 mm.
Description: Unmistakable ant-shaped insects with large broad head, narrow thorax and waisted segment (petiole), bulbous abdomen. Legs relatively long. Antennae long, elbowed in middle. Black, brown, red or yellow, or blotched combinations.
Life history: Complex colonies with fertile female (queen) laying eggs and large numbers, sometimes many thousands, of infertile females (workers), foraging, nest-building and brood-caring. Usually nest in soil, occasionally making small or large mounds. Most temperate ants feed on aphid honeydew (excess, barely processed plant sap, passing through the body), on plant leaves and stems, but also on roots.
Ecological notes: Not dung-feeders, but regularly found sheltering, or even nesting under old dung. A few predatory species may be after small dung organisms. May also be attracted by moisture in fresher droppings.
Shorebug, Saldula orthochila, and other species
Size: Length 3–5 mm.
Description: Small, oval, flattish bug. Black or very dark brown, mottled with pale speckles. Prominent whitish spot on margin of forewing. Legs pale. Many very similar species.
Life history: Fast, active, hop-flying predatory bug. Most species hunt through the herbage, or across bare mud at pond and stream edges, or salt-marshes, hence common name.
Ecological notes: In northern Europe, this is about the only shorebug species known to inhabit dry places, such as wastelands, dunes and fields, and turns up surprisingly often under fairly fresh horse, sheep and cow dung, perhaps relieved by the moisture.
Lesser earwig, Labia minor
Size: Length 4–7 mm.
Description: Small, parallel-sided, vaguely flat insect. More or less uniform pale brown, head and antennae darker. Similar to the very characteristic and well-known common earwig (Forficula auricularia), but smaller, neater, more compact, less shining, with tail-tip forceps shorter, and less curved.
Life history: Scavenger, feeding on decaying plant material in leaf litter, root thatch, compost heaps. Shelters under stones and logs. Female remains to care for egg batch (cleaning away moulds), and regurgitates food for young hatchlings.
Ecological notes: Not a true dung-feeder, but commonly found in manure heaps. Once a very common urban species, in the days of horse transport, living in the frequent heaps of dung cleared from roads. Flies readily. Forficula is sometimes found sheltering under old dung.
Cave crickets, Ceuthophilus, Caconemobius, Hadenoecus and other genera
Size: Body length 15–20 mm.
Description: Characteristic cricket form, with short stout body and long legs, especially hind pair. Antennae very long. Cave species typically lack wings. Colour pale brown to near white.
Life history: Crickets are generally scavengers, eating a mixture of decaying plant material.
Ecological notes: Many crickets have become adapted to cave life, feeding on bat guano, but also scavenging dead insects and whatever other organic matter they can find. In North America, bats are less common in caves, so cave cricket frass has become a specialist microhabitat for yet another layer of organisms. Surface species ‘sing’ by rubbing wings together in courtship displays, but flightless subterranean species have lost this ability. Many have also lost ability to hop, and although retaining long hind legs, the musculature appears to be atrophied.
Centipedes, Lithobius, Haplophilus and other genera
Size: Length 18–80 mm.
Description: Long, narrow, usually pale yellowish, orange or pinkish-brown worm-like animals with many legs, and long antennae. Leg numbers vary from 30 to 200, but only one pair of legs per body segment.
Life history: Predators after small invertebrates in the soil, leaf-litter and root thatch. Shorter, broader species with fewer but longer legs run fast over the ground. Sharp jaws are modified limbs reaching around the head capsule. Longer, narrower species with more but shorter legs adapted to burrowing into the soil or litter layer.
Ecological notes: Regularly invade old dung after prey, or use it as shelter.
Millipedes, Polydesmus, Tachypodoiulus and other genera
Size: 15–60 mm.
Description: Snake millipedes (Tachypodoiulus, etc.) cylindrical, shining, black (sometimes marked with red or yellow patches), up to 250 legs. Flat-backed millipedes (Polydesmus etc) gnarled, knobbled, appearing flat because of flange-like edges, about 80 legs. All millipedes have two pairs of legs per body segment.
Life history: Feed on decaying plant material in leaf-litter, root thatch, under logs, etc.
Ecological notes: Sometimes found under old dung and probably genuinely eating the decaying remains of the plant material within it. In a curious twist of ecological fate, some millipedes produce a secretion to which a few species of dung beetles are attracted. Some beetle species have evolved to become specialist dead millipede scavengers, and one, Deltocheilum valgum, is now an obligate millipede predator.
Mites, Order Acari
Size: Length 0.1–5.0 mm.
Description: Microscopic to minute, round or broad oval, dull yellowish through brown to almost jet black. Body seemingly only one segment. Eight legs.
Life history: Huge diversity but those encountered in dung are either feeding on the decaying plant material, or sucking the body fluids of other dung inhabitants.
Ecological notes: Tiny (0.5 mm) shining black convex oribatid mites are very common in the soil and leaf layer or under rotten logs, and feed on decaying plant material. Large dung beetles are often infested with many larger (1–2 mm) pale mites, which use the adult beetles as transport onwards to the next pat. Sometimes these attached mites are sucking body fluids through chinks in the beetles’ armour, although many disperse on landing to attack much softer fly larvae.
Woodlice, Oniscus asellus, Porcellio scaber and other species
Size: Length 2–20 mm.
Description: Familiar broad, domed, multi-segmented creatures, with relatively long antennae and 14 short legs. Mostly grey, but some flecked with lighter, brighter colours including pink, yellow or orange.
Life history: Scavengers feeding on rotten plant material. Common in leaf-litter, under rocks and stones, logs and loose bark of dead trees.
Ecological notes: Often found under old dung, and probably feeding on the rotting detritus. They may just be sheltering in a dark, damp place, since they are just about our only terrestrial crustaceans and liable to desiccation in hot weather.
Slugs, various species
Size: Length up to 20 cm.
Description: Soft-bodied, elongate, slimy creatures with two short telescopic feelers at front and two longer telescopic eye stalks above. Covered with mucus, they glide along on silvery lubricated trails.
Life history: Mostly scavengers in soil and leaf-litter, eating any decaying organic matter, but many are predators on other slugs, and snails. Lacking the protective but bulky shell possessed by snails, slugs can squeeze into the tightest of spaces and are mainly subterranean, burrowing into the soil, but emerging to feed at night or in moist weather.
Ecological notes: Often found under old dung and may be feeding or sheltering, but also attracted to fresh dung, especially that of carnivores such as cat, fox and badger.
Nematode worms, various species
Size: Length 0.1–2.0 mm.
Description: Microscopic to minute, apparently unsegmented hair-like worms, circular in cross-section. White, black, brown or other colour.
Life history: Mostly parasitic inside a vast diversity of other animal species, including insects.
Ecological notes: Those species found in dung are usually gut or other parasites in the animals that dropped the dung. Often these are the larvae (or eggs), which only develop further if they are reingested by new hosts eating grass contaminated by dung, or on which the infectious stage has lodged after the dropping has been weathered, removed or recycled. Sometimes there is an intermediate host, for example Gongylonema eggs are eaten by dung beetles, which must then get accidentally eaten by cattle, where they infest the digestive tract.
Earthworms, Lumbricus terrestris (lobworm) and other species
Size: Length to 30 cm.
Description: Large, soft, many-segmented worm, bruised pinkish with purple or blue hints. Cylindrical, with swollen ‘saddle’ (containing the reproductive organs) about one-quarter from front end. Tail blunt and flattened at tip.
Life history: Creates temporary mucus-lined burrow in the soil, from which it half ventures out onto the surface at night to feed. It grips leaf fragments in its mouth, and draws them down into the burrow to feed. Two hermaphrodite worms mate, above ground, at night, by aligning saddles and exchanging sperm.
Ecological notes: Soil-dweller (sometimes called the night crawler) and plant-feeder, but often found sheltering under dung, on particles of which it will feed. In temperate zones worms are late arrivers at the pat, but major consumers of the leftovers.
Brandling or tiger worms, Eisenia fetida
Size: Length to 20 cm.
Description: Long, narrow, multi-segmented worm, bright red, often appearing banded red and pink, or orange. Slightly swollen saddle area much less pronounced than in lobworm.
Life history: Not a soil-burrowing worm, but a denizen of the leaf-litter layer. Eats decaying plant material.
Ecological notes: Not strictly a dung-feeder, but often found in manure heaps, as well as compost bins and piles of grass-cuttings. This is the worm which is commercially available for wormeries, free-standing layered composting systems for the garden.
Dog, Canis familiaris
Size: Hugely variable from toys 6 cm at the shoulder to giant mastiffs over 1 m high.
Description: Varying from stunted squat to slim and muscular or nearly skeletal. Colours any combination of black, brown and white. Short or shaggy fur. Pert or flop-eared.
Life history: Familiar family pet, domesticated from grey wolf, Canis lupus. Still loosely carnivorous and behaviour controlled by pack mentality in the human pack home.
Ecological notes: Much to owners’ disgust, will readily eat horse and other dung when out walking. This may be due to vague savoury scents and now well-ingrained habit of eating shapeless mush provided from a tin. However, there are well-documented studies of coprophagy in carnivores where such behaviour is thought to help them mask their carnivore smell from potential prey. Will also roll in dung to smear the fur.
Badger, Meles meles
Size: To 90 cm long, 30 cm high, weighing 15–17 kg.
Description: Stocky, short-legged, broad-shouldered but narrow-snouted mustelid, mostly grey blend of black and white hairs, but with distinctive black and white striped head.
Life history: Large deep burrow nest in the soil all year. Young born in almost any month, though spring peak. Nocturnal foraging after fruit, roots, insects, other small animals, wasp and bee nests.
Ecological notes: Not a dung-feeder, but taste for earthworms means they turn over or dig into old dry cow pats, especially in damp meadows, taking any insect grubs they find too.
Rook, Corvus frugilegus
Size: Length 40–47 cm, wingspan to 90 cm.
Description: Large, handsome, black bird with bluish tinge to feathers. Face bare, grey. Beak narrow, grey. Leg bases shaggily feathered giving impression of wearing ragamuffin trousers.
Life history: Nests high in trees, usually in large groups (rookeries). 3–9 eggs laid in March or April, young fledged in 5–6 weeks. Scavenges for whatever it can find – insects, worms, carrion, fruit, seeds.
Ecological notes: Often seen pecking at seasoned cow pats, where it appears to search for larvae of the noon fly, Mesembrina meridiana. Probably eats other dung-dwellers it comes across. Crows, jackdaws, choughs and magpies have similar behaviour.
Egyptian vulture, Neophron percnopterus
Size: Length to 65 cm, wingspan to 165 cm, weight to 2.8 kg.
Description: South-west Europe, North Africa, Arabian Peninsula, India. Large pale grey, or dusty brown, to bright white bird with black flight feathers in the wings. Unfeathered face bright yellow.
Life history: Scavengers, eating flesh from animal carcasses, also vegetable material, insects and dung.
Ecological notes: Often seen pecking at cow or goat dung, earning it the local name churretero or moniguero, meaning ‘dung-eater’ in Spain. The birds obtain carotinoids, yellow pigments, from the faeces which give their faces the bright colour.
Sitatunga, or marshbuck, Tragelephus spekei
Size: Length 115–170 cm, height 75–125 cm at the shoulder.
Description: Rather shaggy, slim-faced antelope; chestnut red to brown, males with rough main and pale dorsal stripe. Face with white bar below eyes. Males with prominent spiral horns. Hooves splayed.
Life history: Central African. Secretive, in the dense vegetation of the marshy waterways of the Congo, Cameroon, etc., adapted to swamp dwelling by waterproof shaggy coat and splayed hooves which allow them to walk well on mud and floating islands of reed, papyrus, and other water plants.
Ecological notes: Although, like other antelopes, they browse leaves, stems and shoots, sitatunga spend a significant part of their time foraging in elephant dung, eating the barely digested leaf material and also seeds which pass through the elephant digestive tract.
Alabama cave fish, Speoplatyrhinus poulsoni
Size: up to 6 cm.
Description: Slim, unpigmented fish, appearing translucent or slightly pinkish. Blind, lacking eyes. Head blunt, snout flattened. Lacks branching fin rays, fin membranes deeply cut in.
Life history: Occurs only in Key Cave, Lauderdale, Alabama. Poorly known. First discovered in 1974. Population may be less than 100.
Ecological notes: May feed on guano dropped by grey bat, Myotis grisescens, which represents the only biomatter entering the cave. Also likely to eat the other cave-dwelling animals which rely on the nutrient-rich guano, mostly crayfish, isopods and copepods.
Grotto salamander, Eurycea spelaea
Size: Length to 13.5 cm.
Description: Adult pinkish white, but larvae brown or purple, often flecked with yellow. High tail fin, external gills.
Life history: Only known from caves in the Ozark Mountains of central USA. Larvae (tadpoles) live in pools and springs near the cave entrances, are pigmented and have functional eyes. After 2–3 years they metamorphose into adults, lose colours and their eyelids fuse or partially shut.
Ecological notes: Although they probably feed on whatever else they can find in the cave pools, including invertebrates and scavenged material washed in during storm floods, they have often been observed feeding directly on bat guano, and have been measured obtaining nutrition from the droppings. Other amphibians have been observed eating guano, but this was thought to be merely to derive gut bacteria and other micro-organisms.