Myiasis Producing Flies

INSECTA PARASITES

Introduction

Myiasis is the parasitism of living animals by dipteran larvae. Myiasis can be obligatory or facultative (optional) and is described as cutaneous, nasal or somatic.

Oestridae

Warble Fly - Geo. H. Carpenter, Cambridge University Press 1913 - Wikimedia Commons

• The larvae of Oestridae spp. are obligatory parasites

• The three important species of veterinary interest
  • Hypoderma
  • Oestrus
  • Gasterophilus

Hypoderma spp.

• Also known as warble flies

• Important cattle parasites
  • Also parasitise horses, small ruminants and sometimes humans

• The two main species in cattle are
  • H. bovis
  • H. lineatum

• H. diana parasitises deer

• Infestation is largely confined to the Northern Hemisphere

Recognition

• Large 13-15mm long

• Similar to bumble bees in appearance
  • Yellow abdomen
  • Band of black hairs

• 1 pair of wings

Life cycle

• Adult flies emerge on warm, sunny days between June and August

• White eggs are laid around the belly and legs of the animal
  • Eggs are attached to hairs by cement and a small terminal clasp
  • H. lineatum lays a whole row of eggs as it approaches the animal stealthily
  • H. bovis only deposits a single egg before the animal runs away (gadding) as it makes a loud buzzing noise

• The adult lives for 3 weeks

• Females can lay several hundred eggs in their life

• A few days after eggs are laid, larvae emerge and crawl down the hairs into the hair follicles

• Larvae penetrate the skin through wounds made by biting flies

• Larvae migrate through the subcutaneous tissue towards the diaphragm
  • Enzymatic secretions and mouth hooks aid larval migration

• After 3 months larvae reach winter resting sites where they remain from November to February/March whilst moulting to the L2 stage. These sites are different between the 2 species
  • Epidural fat in the spinal canal for H. bovis
  • Wall of the oesophagus for H. lineatum

• Larvae then migrate back to the subcutis along the midline of the back where they bore a breathing hole through the skin and moult to the L3 stage
  • Larvae are palpable as distinct swellings called warbles

• L3 larvae emerge after 4-10 weeks where they fall to the ground and pupate under loose vegetation

• Adult flies emerge from the pupa after 1 month where they copulate, lay eggs and die within 2 weeks

• H. lineatum are on the wing 6-8 weeks before H. bovis

• There is only 1 generation of warble flies per year

Pathogenesis

• Causesmyositis

• Production losses
  • Condemnation and down-grading of hides
  • Reduced milk yield and reduced weight gain
  • Injury from stock panic
  • Trimmed meat losses from H. lineatum
    • 'Butcher's Jelly' around warbles which is green due to mass eosinophil attraction

• Paraplegia resulting from
  • Toxin release
  • Pressure on the spinal cord (H. bovis)
  • Bloat from pressure on the oesophageal wall (H. lineatum)

Control

• Total eradication should be aimed for

• Old methods include popping out warbles
  • But this could lead to anaphylactic shock

• Ectoparasiticides
  • Systemic organophosphorus insecticides in pour-on formula
  • Avermectins and milbemycins in pour-on and injectible formulations

• Timing is crucial for treatment
  • Larvae residing in winter resting sites, if killed, can lead to bloat and paraplegia
  • It is safe to treat in the autumn before larvae reach their winter resting sites and in the spring when the warbles have migrated to the midline of the back
  • Ivermectin can be given at any time without risking host infection as larval antigen is released much slower

Legislation in the UK

• 'Warble Fly Order 1978' requires all clinically affected animals to be treated

• Notifiable disease

• 'Warble Fly Infected Area Order 1983'

• For more information on the warble fly orders, see here

Oestrus ovis

• Also known as the sheep nasal bot fly

• Larvae parasitise the nasal chambers of sheep and goats

• Found in most sheep rearing areas of the world

Recognition

• 13-15mm long

• Grey colouring

• Black spots on abdomen

• Clear wings

• Larvae have distinct black bands on each body segment

Life cycle

• Larvae are squirted into the nostrils of sheep in a jet of liquid by the adult fly

• The larvae crawl caudally into the nasal cavity and feed on the nasal mucosa and mature before returning to the nostrils

• Larval development takes up to 2 months
  • Larvae can overwinter in the nasal cavity if deposited late in the summer

• Once the larvae have developed they are sneezed out and pupate on the ground

• The adult fly emerges 1 months later

• Adult flies only live for 2-3 weeks

Pathogenesis

• Adult flies cause annoyance
  • Interrupts feeding
  • Leads to a decreased weight gain

• Larvae cause nasal irritation, nasal discharge and sneezing
  • Irritate the nasal mucosa with oral hooks and spines causing a viscous exudate to be produced from which they feed
  • Heavy infestations lead to erosion of the bones in the sinuses (turbinate bones)
  • Penetration of the brain leads to false gid (high stepping gait and incoordination)

• Pathology in the nasal cavity

Control

• Systemic insecticides can be used in heavy infestations

• In warmer countries, strategic prophylactic treatment can be used

Gasterophilus spp.

Horse Bot Fly - Dennis Ray - Wikimedia Commons

• Also known as the horse bot fly

• Obligate parasites of equids

• Spend most of life cycle in the equine stomach
  • Of little pathogenic significance

• Three important species (in the UK)
  • G. intestinalis which is the most common
  • G. nasalis
  • G. haemorrhoidalis which is rare

• Two other important veterinary species
  • G. nigricornis
  • G. inermis

Recognition

• Medium to large flies at 10-20mm long

• Look similar to drone bumble bees
  • Body covered with dense yellow hair
  • Dark coloured hairs produce a banding pattern

• Clear wings with brown patches

Life cycle

• Adults are most active in late summer

• Eggs are laid on the body and hatch spontaneously or are stimulated to hatch through an increase in warmth and moisture from the animal self-grooming. They are laid in different areas according to species
  • G. intestinalis eggs are laid in the hair of the shoulders and fore legs
  • G. nasalis eggs are laid in the intermandibular area
  • G. haemorrhoidalis eggs are laid around the lips

• Creamy-white eggs

• 1-2mm in length

• Larvae crawl into the mouth and penetrate the tissues of the tongue and buccal mucosa

• Larvae then emerge after a couple of weeks and are swallowed

• Larvae pass into the stomach and attach to the gastric mucosa
  • Larvae are now known as bots

• Each species attaches to a specific part of the stomach
  • G. intestinalis attaches to the cardiac region
  • G. nasalis attaches to the pylorus

• After 10-12 months in the stomach, the larvae detach and are passed out in the faeces
  • G. haemorrhoidalis attaches to the rectal mucosa before being passed out

• Larvae pupate on the ground

• Adults hatch after 1-2 months and survive from a few days to 2 weeks
  • Adults have non-functional mouthparts so cannot feed

• There is only 1 generation per year in temperate regions of the world

Pathogenesis

• Adults cause annoyance when egg laying
  • Disturbance and panic can ensue

• Larvae cause a marked inflammatory reaction when attached to the gastric mucosa
  • Ring like thickening around the base of each attached larvae

• Large numbers of larvae may interfere with the passage of food and action of gastric sphincters

• G. haemorrhoidalis can cause mild irritation to the rectal wall

• Host reaction to larvae in the mouth is minimal

Control

• Treatment of horses with insecticides over winter
  • Breaks the life cycle as all the population are present as bots in the stomach

• If eggs are present in late summer, the horse's coat can be sponged with an insecticide
  • Stimulates hatching
  • Kills larvae

Dermatobia hominis

Human Bot Fly - Captain R. Goodman, United States Air Force

Human Bot Fly Larvae - Captain R. Goodman, United States Air Force

• Also called the human bot fly

• Larvae are important parasites of both humans and animals

• Specifically found in South America

Recognition

• Adult can grow up to 25mm in length

• Similar to Calliphora in appearance
  • Blue/black
  • Yellow/orange head and legs

• Larvae are distinctive as they taper towards the posterior end

Life Cycle

• Eggs laid on blood sucking flies
  • E.g. On mosquitos, which hatch when the mosquito next lands on a warm blooded animal

• Larvae penetrate skin causing painful swellings

• Larvae emerge after 35-42 days and fall to ground to pupate

• 4 month life cycle

Pathogenesis

• In humans, the larvae are most often found in swellings on the head and limbs

• Larvae cause painful swellings and distress to cattle

• Larvae cause production losses

• Wounds caused by exiting larvae can increase the prevalence of attack by other myiasis flies

Calliphoridae

Pollenia rudis (Calliphoridae) - Richard Bartz, Munich - Wikimedia Commons

Lucilia cuprina - Wikimedia Commons

Blowfly Head Close Up - Martin Pot - Wikimedia Commons

• The Calliphoridae family are facultative parasites

• Cause blowfly strike

• If the fly lays eggs on an animal, the animal is said to be blown

• The damage the larvae cause to the animal is known as strike

• Worldwide

• Affects sheep mostly
  • Rabbits can also be affected

Species of Veterinary importance

• Lucilia serrata
  • Greenbottles

• Phormia terra-novae
  • Blackbottle

• Calliphora erythrocephala
  • Bluebottle

• Callipohora vomitoria
  • Bluebottle

• Lucilia cuprina
  • South Africa, Australia

• Chrysomya spp.
  • Africa, Asia, Australia

• Wohlfahrtia
  • Fleshfly

Recognition of Adults

• Medium sized flies under 10mm long

• Metallic sheen to abdomen
  • Colour depends on species

• Clear wings

Recognition of Larvae

• Smooth maggots

• 10-15mm long

• Spiracles and stigmatic plates on the tail can be used for species differentiation

Life Cycle

• Eggs laid in wounds, soiled fleece and on carrion
  • Females attracted by the odour emitted
  • Clusters of yellow/cream eggs laid

• Larvae hatch 24 hours later and crawl down onto the skin
  • Feed rapidly
  • Grow rapidly
  • Moult twice before becoming full maggots
  • Process takes 1-2 weeks

• Larvae falls to the ground to pupate

• Adult emerges in under a week (during summer)

• The female reaches sexual maturity after a protein meal
  • Lays eggs in batches of 100-200

• Adult flies survive for one month

• Between May and September four generations can develop
  • In warmer climates, up to 9 or 10 generations can develop per year

• Flies can survive the winter as pupae and emerge the next spring

Epidemiology

• Divided into categories depending on their ability to initiate strike
  • Primary flies are capable of initiating a strike on living sheep so larvae can penetrate intact skin
    • E.g. Lucilia, Phormia and Calliphora
  • Secondary flies cannot initiate a strike, larvae attack an area already struck or damaged, extending it
    • E.g. Calliphora and Chrysoma (in warmer climates)
  • Tertiary flies attack lesions on carcasses which have become dry
    • E.g. Musca and Sarcophaga spp.

• Temperature will affect the prevalence of strike
  • Temperature in the spring will determine when the overwintering larvae hatch
  • High temperature and humidity will create a microclimate in the fleece, attracting adult flies to lay eggs

• Persistent rainfall will make the fleece microclimate attractive to adult flies
  • Females lay eggs after the rain ceases
  • Breeds with long, fine wool are the most susceptible

• Host susceptibility is increased when putrefactive odours develop on the fleece due to bacterial decomposition of organic matter
  • Merino sheep have a narrow breech area with excessive wrinkling making them more susceptible to soiling
  • A narrow opening of the penile sheath in rams and wethers may result in accumulation of urine and increase blowfly strike in this area
  • Cuts during shearing, fighting and barbed wire will also increase the incidence of blowfly strike

Pathogenesis

• Severe skin damage
  • Larvae lacerate skin with oral hooks and liquefy host tissue by secreting proteolytic enzymes

• Skin lesions are extended and deepened
  • Secondary blowfly attack
  • Flies attracted to odour of decomposing tissue

• Secondary bacterial infection

• Production losses
  • Irritation and distress associated with skin lesions
  • Poor weight gain (often the first clinical sign)

• Risk of strike is highest in warm, moist weather

Clinical Signs

• Anorexia, listlessness, animals standing apart from flock

• Fleece may appear darker, be damp and have a foul smell

• Strike is classified according to where the body is affected

Control

• Prophylactic insecticide treatment
  • Must kill larvae and remain in fleece to prevent flies from laying eggs
  • Spraying, dipping, spray race or jetting

• Insect growth regulators
  • Pour-on
  • 2-4 month protection depending upon the product used

• Effective worm control
  • To minimise diarrhoea and therefore soiled fleece

• Crutching

• 'Mule's operation'
  • Surgical removal of breech skin in Merino breeds

• Tail docking of lambs

• Proper carcass disposal
  • Eliminates fly breeding sites

• Vaccination

Screw Worm Myiasis

Screw worm larvae - John Kucharski - Wikimedia Commons

• C. bezziana cause myiasis in both animals and humans

• Located mainly in tropical regions

• Larvae are obligate parasites

Recognition

• Similar to Calliphora
  • Iridescent
  • Clear wings
  • Blue abdomen

• Longitudinal stripes on thorax

• Larvae have bands of spines
  • Look like screws

Life Cycle

• Eggs laid in wounds or body cavities

• Larvae feed as colonies

• Larvae drop to the ground to pupate

Pathogenesis

• Spiracles are exposed as larvae feed which expands the wound
  • Creates a foul smelling lesion

• Cause irritation and pyrexia

Control

• In the USA
  • Mass eradication through the release of sterile males
  • Currently only persists where flies have migrated across the Mexican border

• In Africa
  • Introduced into Libya through the importation of infested livestock
  • Sterile males released
  • Eradication occurred in 1991

Maggot Debridement Therapy

Maggot therapy on a wound - Wikimedia Commons

• Human medicine
  • Sterile Lucilia sericata maggots used to treat infected and necrotic wounds
  • Larvae secrete proteolytic enzymes and antimicrobial agents into the wound
    • Larvae do not burrow under the skin or attack healthy tissue

• Veterinary medicine
  • Published reports rare
  • Recently used successfully to treat a suppurative wound in a donkey that did not respond to conventional methods and surgery

Wohlfahrtia sp.

• Obligatory parasite

• Occurs in North America

• Parasite of mink and sometimes humans

Recognition

• Large 8-14mm long

• Pale grey

• Black stripes on thorax

• Black spots on abdomen

Life Cycle

• Larvae deposited directly onto host
  • No egg stage

• Larvae moult twice on host then fall to ground to pupate

Pathogenesis

• Larvae penetrate intact skin
  • Cause boil like swellings

Links

• Viruses

• Ectoparasiticides

• Blowfly strike