Costelytra Giveni
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''Costelytra giveni'' (mischaracterized as '' Costelytra zealandica'' prior to 2016), commonly known as New Zealand grass grub, is a
scarab beetle The family Scarabaeidae, as currently defined, consists of over 35,000 species of beetles worldwide; they are often called scarabs or scarab beetles. The classification of this family has undergone significant change. Several groups formerly tre ...
that is endemic to
New Zealand New Zealand () is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and List of islands of New Zealand, over 600 smaller islands. It is the List of isla ...
and is a prevalent
pasture Pasture (from the Latin ''pastus'', past participle of ''pascere'', "to feed") is land used for grazing. Types of pasture Pasture lands in the narrow sense are enclosed tracts of farmland, grazed by domesticated livestock, such as horses, c ...
pest.


Taxonomy

''Costelytra giveni'' was first described by B.B. Given in 1952, though he mischaracterized specimens as '' C. zealandica''. In 2016, Coca-Abia and Romero-Samper found differences in syntype specimens between White's (1846) ''C. zealandica'' and Given's (1952) description, and revised the species name of the latter to ''C. giveni'' after Given''.''


Identification/description

New Zealand grass grubs are one of the most common insects in New Zealand. Grass grubs belong to the
holometabolous Holometabolism, also called complete metamorphosis, is a form of insect development which includes four life stages: egg, larva, pupa, and imago (or adult). Holometabolism is a synapomorphic trait of all insects in the clade Holometabola. Immatur ...
insect group, this means that they have a life cycle with four stages: egg, larva, pupa, adult. The adult beetle can grow up to a mature length of 13 mm. It has a shiny brown colouring with a thick wing covering called an
elytra An elytron (; ; : elytra, ) is a modified, hardened forewing of beetles (Coleoptera), though a few of the true bugs (Hemiptera) such as the family Schizopteridae are extremely similar; in true bugs, the forewings are called hemelytra (sometime ...
. Adult beetles also have a prominent plate-like piece covering the
thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main di ...
called the
pronotum The prothorax is the foremost of the three segments in the thorax of an insect, and bears the first pair of legs. Its principal sclerites (exoskeletal plates) are the pronotum (dorsal), the prosternum (ventral), and the propleuron (lateral) on e ...
. These adults can be seen flying in the evenings during the summer months. Female adults lay up to 100 eggs at a time into wet or loose soil, 10–15 days after mating. Eggs are small, white, oval shaped and around 1.5 mm in diameter. 3–4 days after being laid, the eggs turn smooth, spherical, and start to swell by absorbing moisture. Once the eggs are mature they turn a dark brown/black colour. On hatching
larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
have a whitish/grey C-shaped segmented body, with a darker coloured head and dark brown jaws (
mandibles In jawed vertebrates, the mandible (from the Latin ''mandibula'', 'for chewing'), lower jaw, or jawbone is a bone that makes up the lowerand typically more mobilecomponent of the mouth (the upper jaw being known as the maxilla). The jawbone i ...
). The larvae have three pairs of legs (true legs) attached to the underside of the segment behind the head, called the
thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main di ...
. The length of the larvae ranges from 5 mm, when newly hatched, to 25 mm at larval maturity. Newly hatched larvae weigh 2–3 mg and grow up to 200 mg. The most obvious characteristic of larva is that the abdomen is very swollen and looks a lot bigger than any other part of the body. There are many thin fine hairs along both sides of the body, especially around the abdomen. Occasionally, grass grub larvae can be found that are purple/yellow or green in colour, these are diseased. The
pupae A pupa (; : 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 life cycle, the stages th ...
start off the same creamy white colour as the larvae, but slowly turn to a dark brown during pupation. They have a soft body and are generally shorter and thicker than the grass grub larvae, they grow to between 10–30 mm in length. The pupae have a thin translucent skin called the
epidermal The epidermis is the outermost of the three layers that comprise the skin, the inner layers being the dermis and hypodermis. The epidermal layer provides a barrier to infection from environmental pathogens and regulates the amount of water relea ...
layer, through which the developing wings and legs can be seen.


Distribution

''Costelytra giveni'' is endemic to and found throughout New Zealand. There are, however, similar species found in other countries like '' Acrossidius tasmaniae'', the Tasmanian grass grub found in
Tasmania, Australia Tasmania (; palawa kani: ''Lutruwita'') is an island States and territories of Australia, state of Australia. It is located to the south of the Mainland Australia, Australian mainland, and is separated from it by the Bass Strait. The sta ...
. Grass grubs prefer altitudes below 1200m, with favored soil types being free-draining loam soils, but they are also found in peaty or sandy soils.


Life cycle/phenology

New Zealand grass grub has an annual life cycle, taking typically one year under normal climatic conditions but can take up to two years in adverse weather such as drought or significant cold periods (this also includes periods where the soil moisture is very high), or due to a food shortage. Over that one-year period the adult beetles live for 4–6 weeks emerging in spring (October – December) when the air temperature is above 10 degrees Celsius, and undertake two main flights, the first flight is to mate and the second and consecutive are to feed and lay clutches of eggs. Females are the last to emerge from the soil, this leads to localized infestations due to them being unable to reach a significant distance before being mated. Egg cluster size can range from 5-80 eggs and range from 80–250 mm below the surface of the soil. Females can lay two to three clusters over a two to three week period. ''Costelytra giveni'' larvae hatch between November and March (depending on the time they were laid). They start to feed on roots of
clover Clovers, also called trefoils, are plants of the genus ''Trifolium'' (), consisting of about 300 species of flowering plants in the legume family Fabaceae originating in Europe. The genus has a cosmopolitan distribution with the highest diversit ...
and pasture plants from the time they hatch until September. They undergo three stages (
instar An instar (, from the Latin '' īnstar'' 'form, likeness') is a developmental stage of arthropods, such as insects, which occurs between each moult (''ecdysis'') until sexual maturity is reached. Arthropods must shed the exoskeleton in order to ...
s) before pupation. During the summer months, the grubs moult and reach their second instar phase. By early autumn, they are usually fully grown and have reached their third instar phase. This is when they are the most detrimental to pastures as it is the final feeding phase before winter. In late autumn and during winter, they retreat down from the top 5 cm of soil and burrow down between 50–200 mm into the soil. During this phase, the grubs undergo a colour change from grayish/white into a yellow/cream colour. Once they reach the appropriate depth, the grub empties its stomach and starts to form a smooth oval shaped cell. The developing wings and legs can be seen through the then-translucent skin (epidermal layer) as develops. The size range of pupae is 10–30 mm in length. This procedure takes place over 30–40 days. The change from pupa to mature beetle takes place over 4–6 weeks, but they remain in the soil for up to a week to let the newly formed outer skin harden, then it proceeds to dig its way up to the surface of the pasture. The grass grub populations usually increase in a 3-5 year cycle before crashing, usually due to a buildup of disease in the soil.


Diet and foraging

The larvae of ''C. giveni'' tend to feed on fine roots of pasture plants like perennial ryegrass (''
Lolium perenne ''Lolium perenne'', common name perennial ryegrass, English ryegrass, winter ryegrass, or ray grass, is a grass from the family Poaceae. It is native to Europe, Asia and northern Africa, but is widely cultivated and naturalised around the world. ...
'') and white clover (''
Trifolium repens ''Trifolium repens'', the white clover, is a herbaceous perennial plant in the bean family Fabaceae (otherwise known as Leguminosae). It is native to Europe, including the British Isles, and central Asia and is one of the most widely cultivated ...
''). Originally, only native species of grasses such as tussocks were affected, but they have subsequently adapted to feed on the exotic species that have been brought into New Zealand, as their abundance continues to grow. Adult beetles feed on leaves of trees, shrubs,
brassica ''Brassica'' () is a genus of plants in the cabbage and mustard family (Brassicaceae). The members of the genus are informally known as cruciferous vegetables, cabbages, mustard plants, or simply brassicas. Crops from this genus are sometim ...
s and foliage of pasture crops. If there is a concentrated group of beetles flying in one area, there is potential to defoliate seedling crops and fruit trees.


Predators, parasites, and diseases

There are some species of pasture grasses, legumes, and herbs that are naturally tolerant of grass grubs including cocksfoot (''
Dactylis glomerata ''Dactylis glomerata'' is a species of flowering plant in the grass family Poaceae, known as cock's-foot,Interactive Flora of NW Europ''Dactylis glomerata'' (Cock's-foot)/ref> also colloquially as orchard grass, or cat grass (due to its populari ...
''), tall fescue (''
Festuca arundinacea ''Lolium arundinaceum'', commonly known as tall fescue, is a cool-season, perennial C3 grass species native to Europe and introduced to North America and other parts of the world. It naturally occurs in grasslands and coastal marshes.Texas Park ...
'') and chicory (''
Cichorium intybus Common chicory (''Cichorium intybus'') is a somewhat woody, perennial herbaceous plant of the family Asteraceae, usually with bright blue flowers, rarely white or pink. Native to Europe, it has been introduced to the Americas and Australia. M ...
'') and some that are resistant like Phalaris (''
Phalaris aquatica ''Phalaris aquatica'', known by the common names bulbous canary-grass and Harding grass, is a species of grass in the genus ''Phalaris'' of the family Poaceae. Description It is an erect, waist-high, stout perennial bunch grass, with grayish to ...
''), lucerne, (
Alfalfa Alfalfa () (''Medicago sativa''), also called lucerne, is a perennial plant, perennial flowering plant in the legume family Fabaceae. It is cultivated as an important forage crop in many countries around the world. It is used for grazing, hay, ...
), (''
Medicago sativa Alfalfa () (''Medicago sativa''), also called lucerne, is a perennial flowering plant in the legume family Fabaceae. It is cultivated as an important forage crop in many countries around the world. It is used for grazing, hay, and silage, as we ...
'') and lotus ('' Lotus uliginosus''), ('' Lotus major''). ;Predators Because they are insects, grass grubs fall prey to almost all bird species present in the same areas. Great black backed gulls (''
Larus marinus The great black-backed gull (''Larus marinus'') is the largest member of the gull family. It is a very aggressive hunter, pirate, and scavenger which breeds on the coasts and islands of the North Atlantic in northern Europe and northeastern Nort ...
'') prey on the grubs especially in areas undergoing cultivation. Starlings (''
Sturnus vulgaris The common starling (''Sturnus vulgaris''), also known simply as the starling in Great Britain and Ireland, and as European starling in North America, is a medium-sized passerine bird in the starling family, Sturnidae. It is about long and ha ...
'') and black birds ('' Turdus merula'') seem to favour grass grub larvae in their third instar phase, as well as the adult beetle, and are often seen actively digging through the soil in areas with a high grub population. Another common predator is the hedgehog (''
Erinaceus europaeus The European hedgehog (''Erinaceus europaeus''), also known as the West European hedgehog or common hedgehog, is a hedgehog species native to Europe from Iberia and Italy northwards into Scandinavia and westwards into the British Isles.Harris, S. ...
'') whose intake of grass grubs can be up to 40% of the total diet of an adult hedgehog. There are a few invertebrate predators but they are not common in pastures. ;Parasites There have been recordings of
Diptera Flies are insects of the order Diptera, the name being derived from the Greek δι- ''di-'' "two", and πτερόν ''pteron'' "wing". Insects of this order use only a single pair of wings to fly, the hindwings having evolved into advance ...
species being parasites and predators of New Zealand grass grub, an example is ''
Proscissio cana ''Proscissio'' is a genus of flies in the family Tachinidae. Species *'' Proscissio albiceps'' Malloch, 1938 *'' Proscissio cana'' Hutton, 1901 *'' Proscissio lateralis'' Malloch, 1938 *'' Proscissio milleri'' Malloch, 1938 *'' Proscissio m ...
'' (
Tachinidae The Tachinidae are a large and variable family of true fly, flies within the insect order Fly, Diptera, with more than 8,200 known species and many more to be discovered. Over 1,300 species have been described in North America alone. Insects in t ...
) which parasitizes larvae in the third instar phase and is present in the Canterbury High country, but it only parasitizes around 20% of larvae present. In November 2012, researchers in New Zealand discovered an unknown maggot preying on grass grub larvae, with 35-50% of the grubs in the nearby area being parasitized. DNA samples from these maggots were not matched to any known species so a sample was collected and allowed to pupate, and they were finally identified as '' Ostenia robusta'' (
Dolichopodidae Dolichopodidae, the long-legged flies, are a large, cosmopolitan family of true flies with more than 8,000 described species in about 250 genera. The genus '' Dolichopus'' is the most speciose, with some 600 species. Dolichopodidae generally a ...
). This is the sole species in the genus ''Ostenia,'' is endemic to New Zealand and not much is known about its biology or ecology. ;Diseases Because grass grub build up large colonies in the soil, disease can spread quickly and be devastating to the population. The most common disease of the New Zealand Grass Grub is Amber disease. Amber disease is a chronic infection of the stomach of grass grub larvae, and is caused by two strains of bacteria, '' Serratia entomophila'' and '' S. proteamaculans''. The bacteria ''S. entomophila'' is available for purchase commercially as a biocontrol agent. Once these pathogenic bacteria have been ingested, the infected larvae stop eating within 2–5 days, and gain an amber colouring due to the stomach emptying, and the levels of major digestive enzymes (
trypsin Trypsin is an enzyme in the first section of the small intestine that starts the digestion of protein molecules by cutting long chains of amino acids into smaller pieces. It is a serine protease from the PA clan superfamily, found in the dig ...
and
chymotrypsin Chymotrypsin (, chymotrypsins A and B, alpha-chymar ophth, avazyme, chymar, chymotest, enzeon, quimar, quimotrase, alpha-chymar, alpha-chymotrypsin A, alpha-chymotrypsin) is a digestive enzyme component of pancreatic juice acting in the duodenu ...
) in the mid-gut decreasing. However, the infected larvae take 1–3 months to die after becoming infected. Amber disease can carry over into following generations as it is applied at a subsurface level of soil. It was first isolated in Mid Canterbury, New Zealand in 1981. '' Yersinia entomophaga'' was first isolated from diseased grass grub larvae and is also being studied for
biological pest control Biological control or biocontrol is a method of controlling pests, whether pest animals such as insects and mites, weeds, or pathogens affecting animals or plants by using other organisms. It relies on predation, parasitism, herbivory, or ot ...
.


Cultural uses

The Māori name for ''C. giveni'' is Tutaeruru. Grass grubs are very damaging to agricultural communities. Loss in crop productivity can be the cause of economic loss. Previous findings show that the cost of crop damage due to ''C. giveni'' is said to be around $89 million per year.


References

{{Taxonbar, from=Q28967017 Beetles described in 1846 Endemic beetles of New Zealand Melolonthinae