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''Eriococcus orariensis'', commonly known as the mānuka blight, is a felt
scale insect Scale insects are small insects of the order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient grouping than th ...
in the genus '' Eriococcus''. It is native to Australia, but was discovered in New Zealand in 1937 after being accidentally introduced to the country.


Description

To identify and differentiate ''Eriococcus orariensis'' from similar ''Eriococcus'' species a microscope is needed. ''E. orariensis'' is
sexually dimorphic Sexual dimorphism is the condition where the sexes of the same animal and/or plant species exhibit different morphological characteristics, particularly characteristics not directly involved in reproduction. The condition occurs in most ani ...
. Adult females are typically light brown and an oval shape that tapers towards the anal lobes. Adult females lack wings and average 1.25mm in length and 0.84mm in width. Their antennae are six-segmented, the third segment being the longest. Female egg sacs are also greyish-white, closely felted, and open towards the anal end. ''Eriococcus orariensis'' female dorsal (upper) and ventral (lower) body surfaces are membranous and irregularly covered in small, inconspicuous, bristle-like structures called
seta In biology, setae (singular seta ; from the Latin word for "bristle") are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Annelid setae are stiff bristles present on the body. T ...
e; although the setae extend in semi-regular rows along the abdominal segments. There is no noticeable marginal fringe of setae on the dorsum, again excepting the distinct abdominal segments. A small number of quinquelocular sessile pores exist on the ventral abdominal segments in association with respiratory openings called spiracles. Contrastingly, long tubular ducts ending in cup-shaped pores cover the dorsum in moderate numbers, also concentrating on abdominal segments. ''Eriococcus orariensis'' female anal lobes are cylindrical for two-thirds their length before tapering towards the base of much longer caudal (tail-like) setae. A prominent feature used to differentiate ''E. orariensis'' from other ''Eriococcus'' species in a key created by the entomologist, Dr James Mather Hoy, are three dorsal setae on the anal lobe: two towards the base and one two-thirds the distance up towards the inner margin. The anal ring has eight setae. Legs in both mature sexes are well developed, but relatively small for the species' size. Back leg tarsi are longer than tibiae. Curved claws at the end of each leg have a small tooth-like projection. ''Eriococcus orariensis'' adult males are smaller than females, averaging 0.84mm in length. They are typically reddish-yellow with an abdominal spike and a pair of long filaments extending from the second to last abdominal segment. They have iridescent wings fringed with hairs, and
halteres ''Halteres'' (; singular ''halter'' or ''haltere'') (from grc, ἁλτῆρες, weights held in the hands to give an impetus in leaping) are a pair of small club-shaped organs on the body of two orders of flying insects that provide infor ...
with a hook-like projection called a hamulus. Male puparia are oval, white, cottony and not as closely felted as females. Male antennae are ten-segmented with many setae, but the third segment is also longest. They have four light-detecting organs called
ocelli A simple eye (sometimes called a pigment pit) refers to a form of eye or an optical arrangement composed of a single lens and without an elaborate retina such as occurs in most vertebrates. In this sense "simple eye" is distinct from a multi-le ...
: two close together on the mid-ventral surface, and two on the lateral margins. Adult males also have no mouthparts. Their legs are very setose, with a longer spine-like seta on each tibia. The first
instar An instar (, from the Latin '' īnstar'', "form", "likeness") is a developmental stage of arthropods, such as insects, between each moult (''ecdysis''), until sexual maturity is reached. Arthropods must shed the exoskeleton in order to grow o ...
nymphs of ''Eriococcus orariensis'' are pale pink or brown, with an average a 0.37mm length and 0.17mm width. They are most similar to adult females, with slight differences. For example, nymph anal lobes are more cone-like, their dorsal abdominal setae are more spine-like and arranged in rows lengthwise, and they only have six setae on their anal ring.


Range

''Eriococcus orariensis'' is native to Australia. There they are widely distributed through Southern and Eastern Australia, including Tasmania, but populations exist in low numbers there due to an abundance of natural predators and parasitoid species. ''Eriococcus orariensis'' was first discovered in New Zealand in 1937 at Orari Gorge, Canterbury. It was introduced from Australia, but exactly when and how is unknown. The most likely theory is that it was accidentally introduced on imported ornamental ''
Leptospermum scoparium ''Leptospermum scoparium'', commonly called mānuka, () mānuka myrtle, New Zealand teatree, broom tea-tree, or just tea tree, is a species of flowering plant in the myrtle family Myrtaceae, native to New Zealand (including the Chatham Islan ...
'' (mānuka) plant material. Initially, ''Eriococcus orariensis'' appeared restricted to Orari Gorge, with a slow spread through surrounding Geraldine through the 1940s. However, in the late 1940s-50s ''E. orariensis'' was deliberately spread to farms across the country due to its ability to kill mānuka within years of attack, which was considered a major pasture weed at the time. By 1954 it was widespread through mānuka populations on both main islands, likely aided by its release from natural enemies only found in Australia. Today, although exact distributions of ''Eriococcus orariensis'' in New Zealand are unknown, it appears their population has been dramatically reduced in range to several sites in Nelson, Otago, Fiordland, and the West Coast, and replaced by the scale insects '' Acanthococcus campbelli'' (Hoy) and ''Acanthococcus leptospermi'' (Maskell). This is likely due to parasitism by the entomogenous fungi, ''Angatia thwaitesii'' Petch, 1924, discovered in New Zealand in 1957.


Habitat

''Eriococcus orariensis'' is host-specific to six species of ''
Leptospermum ''Leptospermum'' is a genus of shrubs and small trees in the myrtle family Myrtaceae commonly known as tea trees, although this name is sometimes also used for some species of ''Melaleuca''. Most species are endemic to Australia, with the great ...
'' plants in Australia. In New Zealand it is only found on ''
Leptospermum scoparium ''Leptospermum scoparium'', commonly called mānuka, () mānuka myrtle, New Zealand teatree, broom tea-tree, or just tea tree, is a species of flowering plant in the myrtle family Myrtaceae, native to New Zealand (including the Chatham Islan ...
'' (mānuka) and ''
Kunzea ericoides ''Kunzea ericoides'', commonly known as kānuka, kanuka, white tea-tree or burgan, is a tree or shrub in the myrtle family, Myrtaceae and is endemic to New Zealand. It has white or pink flowers similar to those of '' Leptospermum'' and from its ...
'' (kānuka - previously ''Leptospermum ericoides''). It can survive in all places those trees grow, but consistently establishes at higher rates on mānuka, as mānuka bark does not exfoliate in papery sheets like kānuka does, so offers many more establishing sites for the insect. ''E. orariensis'' occupies the specific niche of mānuka bark crevices in trunks and branches to avoid competition with other scale insects like ''Eriococcus leptospermi'' which occupies new plant growth. While male puparia sacs tend to cover the surface of sooty mold and bark on mānuka plants, female egg sacs and early ''E. orariensis'' stages tend to stay sheltered under the bark.


Ecology


Life cycle

''Eriococcus orariensis'' eggs are laid around February in New Zealand inside a sac that the female produces around herself. A single egg is laid approximately every eight hours, until an average of 47 eggs are laid per female, although females often die before all eggs are laid. Typically, eggs hatch within 15 minutes of being laid, regardless of external factors. Nymphs then remain inactive for a variable period, depending on environmental temperatures. If air temperatures are above 21 °C, nymphs will begin rapidly moving across the host plant surface 20 minutes after hatching in search of a feeding site. Using their antennae to feel the bark surface, this search can take hours to days until the nymph finds a suitable crevice to wedge itself into and insert its mouthparts. First-stage nymphs do not tend to travel once attached to a feeding site. They feed and grow for several months until they increase by about 1.7x in length and 1.9x in width. The feeding period can vary depending on when nymphs hatch, but is finished by September. When finished, nymphs remove their mouthparts from the host plant. If female, within 2–3 days of finishing the feeding period around September, the nymph will moult and an intermediate female stage will emerge and reinsert its mouthparts to continue feeding and growing within 6–8 hours. They do not usually travel, unless the plant material in their crevice has dried out. Around October they moult again, and an adult female emerges, reinserts their mouthparts within 6 hours, and continues feeding. When male nymphs finish the feeding period, second stage males travel across the host plant to find a site to build a
pupa A pupa ( la, pupa, "doll"; plural: ''pupae'') is the life stage of some insects undergoing transformation between immature and mature stages. Insects that go through a pupal stage are holometabolous: they go through four distinct stages in their ...
rium. They appear to prefer sheltered sites like bark crevices, but can settle on any surface if a plant is overly populated. Puparium sacs are completed from waxy filaments excreted from dorsal tubular ducts 4–6 days after a site is found. Prepupa emerge 2–3 days after completion and adult males emerge between October–January. Mating occurs October–January. Males may mate multiple times, but do not feed, so die within days. Females only mate once, so only one generation occurs per year. Within two hours of fertilisation, females drastically increase production of waxy threads from their sessile pores and tubular ducts, encasing themselves in a sac within 4–6 days, ready for egg laying.


Diet and foraging

''Eriococcus orariensis'' is host-specific to a small number of plant species, only feeding on ''
Leptospermum scoparium ''Leptospermum scoparium'', commonly called mānuka, () mānuka myrtle, New Zealand teatree, broom tea-tree, or just tea tree, is a species of flowering plant in the myrtle family Myrtaceae, native to New Zealand (including the Chatham Islan ...
'' (mānuka) and ''
Kunzea ericoides ''Kunzea ericoides'', commonly known as kānuka, kanuka, white tea-tree or burgan, is a tree or shrub in the myrtle family, Myrtaceae and is endemic to New Zealand. It has white or pink flowers similar to those of '' Leptospermum'' and from its ...
'' (kānuka) in New Zealand, and several other ''
Leptospermum ''Leptospermum'' is a genus of shrubs and small trees in the myrtle family Myrtaceae commonly known as tea trees, although this name is sometimes also used for some species of ''Melaleuca''. Most species are endemic to Australia, with the great ...
'' species in Australia. ''E. orariensis'' are sap feeders for several life cycle stages, the most important being the feeding stage of first-stage nymphs. Once a suitable feeding site is found, the nymph will wedge itself firmly into the crevice. It then inserts its piercing, sucking mouthparts into the plant tissue, arching its body and thrusting forwards to push itself deeper. It then feeds in the sap stream for several months, staying in the same spot. Because plant sap is mainly sugar, the insects must feed continuously to get required nutrients like nitrogen and excrete excess sugar through their anal opening. Intermediate-stage and adult females also feed continuously in this manner, usually not moving from the feeding site they selected as nymphs. Adult males do not feed again, as they lack mouthparts after emerging from their puparium.


Predators, parasites, and diseases

Despite initial devastation of ''Eriococcus orariensis'' on mānuka populations across New Zealand, its numbers plummeted from the late 1950s onwards, following the likely accidental introduction of the entomogenous
fungi A fungus (plural, : fungi or funguses) is any member of the group of Eukaryote, eukaryotic organisms that includes microorganisms such as yeasts and Mold (fungus), molds, as well as the more familiar mushrooms. These organisms are classified ...
, ''Angatia thwaitesii'', from Australia. Once fungal mycelia enter the insect, they quickly grow through the body and kill the insect, using up its nutrients until a black mass of hyphae remain. All insect life stages are vulnerable to ''A. thwaitesii'', although adult females are attacked most frequently. Interestingly, ''A. thwaitesii'' is likely a specific pathogen of ''E. orariensis'' as no other ''Eriococcus'' species in New Zealand or Australia are attacked. This partially explains why species like ''E. campbelli'' and ''E. leptospermi'' have replaced ''E. orariensis'' as successful mānuka
scale insect Scale insects are small insects of the order Hemiptera, suborder Sternorrhyncha. Of dramatically variable appearance and extreme sexual dimorphism, they comprise the infraorder Coccomorpha which is considered a more convenient grouping than th ...
s nationwide. The Australian ladybeetle, ''
Rhyzobius ventralis ''Rhyzobius ventralis'', common names including black lady beetle, gumtree scale ladybird, is a ladybird species endemic to Tasmania and all the mainland states of Australia except the Northern Territory. It is also found in New Zealand, but not ...
'' Erichson, 1843, also predates ''Eriococcus oraiensis'', however, it is mainly associated with the gum tree scale, ''
Eriococcus coriaceus Eriococcus coriaceus is a scale insect of the genus Eriococcus. Its common name In biology, a common name of a taxon or organism (also known as a vernacular name, English name, colloquial name, country name, popular name, or farmer's name) ...
''. As a result, its effect on ''E. oraiensis'' is minimal.


Cultural uses

Perhaps the most interesting thing about ''Eriococcus orariensis'', apart from its drastic
population bottleneck A population bottleneck or genetic bottleneck is a sharp reduction in the size of a population Population typically refers to the number of people in a single area, whether it be a city or town, region, country, continent, or the world. Go ...
associated with ''Angatia thwaitesii'', is the lengths people took to spread a known plant blight. When ''E. orariensis'' was first noted in the late 1930s, mānuka trees died within several years of insect invasion. By the late 1940s, it was supposedly hard to find live mānuka anywhere in South Canterbury. This was because large, uncontrolled populations of ''E. orariensis'' remove too many nutrients for a tree to survive. Additionally, their sugar excretions promote the growth of the
sooty mold Sooty mold (also spelled sooty mould) is a collective term for different Ascomycete fungi, which includes many genera, commonly '' Cladosporium'' and '' Alternaria''. It grows on plants and their fruit, but also environmental objects, like fences ...
, ''Capnodium walteri'' Sacc, 1893, which reduces plant photosynthetic capacity. Far from causing concern as the discovery of a new blight might today, ''Eriococcus orariensis'' was quickly and widely adopted by farmers to remove mānuka, which was perceived as a pesky, economically damaging weed. The
New Zealand Department of Scientific and Industrial Research The Department of Scientific and Industrial Research (DSIR) is a now-defunct government science agency in New Zealand, founded in 1926 and broken into Crown Research Institutes in 1992. Foundation DSIR was founded in 1926 by Ernest Marsden aft ...
even suggested "mānuka blight" was the most efficient biological control of a plant ever seen in New Zealand. Infected material was frantically sold, distributed via post, and even aerial dropped in the North Island.


References

{{Taxonbar, from=Q10488022 Eriococcidae Insects described in 1954 Hemiptera of Australia Hemiptera of New Zealand