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''Percnodaimon merula'', the black mountain ringlet, is a satyrid butterfly in the family
Nymphalidae The Nymphalidae are the largest family of butterflies, with more than 6,000 species distributed throughout most of the world. Belonging to the superfamily Papilionoidea, they are usually medium-sized to large butterflies. Most species ha ...
. It is currently the only recognised species in the
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...
genus ''Percnodaimon'',
endemic Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
to New Zealand, although there may be other undescribed species in the genus. The black mountain ringlet is notable for living exclusively in rocky areas of New Zealand's
Southern Alps The Southern Alps (; officially Southern Alps / Kā Tiritiri o te Moana) are a mountain range extending along much of the length of New Zealand, New Zealand's South Island, reaching its greatest elevations near the range's western side. The n ...
, usually above 1200 m. Its eggs are laid on rocks, its larvae feed on mountain ''
Poa ''Poa'' is a genus of about 570 species of Poaceae, grasses, native to the temperate regions of both hemispheres. Common names include meadow-grass (mainly in Europe and Asia), bluegrass (mainly in North America), tussock (some New Zealand spe ...
'' species, and it pupates under a stone. It has distinctive dark velvety wings and a zig-zag flight pattern over the scree slopes on which it lives.


Taxonomy

This species has had a complicated taxonomic history. It was originally described as ''Erebia pluto'' by Richard W. Fereday in 1872 from the Craigieburn Range in the South Island, and was moved to the new genus ''Percnodaimon'' by
Butler A butler is a person who works in a house serving and is a domestic worker in a large household. In great houses, the household is sometimes divided into departments, with the butler in charge of the dining room, wine cellar, and pantries, pantr ...
in 1876. It was known as ''Percnodaimon pluto'' for many years, and Wise in 1967 regarded this as the correct name for the species, but was eventually synonymised with ''P. merula'', under which name it is usually referred to today. Most sources consider ''Percnodaimon''
monotypic In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unisp ...
, but in 2012
mitochondrial gene Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the DNA contained in ...
sequencing by Hamish Patrick for his Lincoln University Hons thesis suggested it was a
species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
, with up to six species in the genus. In their 2012 guide to South Pacific butterflies, Brian and Hamish Patrick stated there are up to eight species of ''Percnodaimon'' still to be described, based on wing shape, pattern, and colour, size, behaviour, and season of emergence. Their species list was as follows: * ''Percnodaimon pluto'' (Fereday, 1872) – the commonest black mountain butterfly of the eastern
South Island The South Island ( , 'the waters of Pounamu, Greenstone') is the largest of the three major islands of New Zealand by surface area, the others being the smaller but more populous North Island and Stewart Island. It is bordered to the north by ...
* ''Percnodaimon micans'' (new status) – described by
Augustus Hamilton Augustus Hamilton (1 March 1853 – 12 October 1913) was a New Zealand ethnologist, biologist and museum director. He was born in Poole, Dorset, England on 1 March 1853. He wrote on the fishing and seafoods of the ancient Māori people. He also w ...
in 1909 as ''Erebia pluto'' var. ''micans'', and found in the Harris and Richardson mountains of
Otago Otago (, ; ) is a regions of New Zealand, region of New Zealand located in the southern half of the South Island and administered by the Otago Regional Council. It has an area of approximately , making it the country's second largest local go ...
* ''Percnodaimon'' sp.nov. 1 –
Fiordland Fiordland (, "The Pit of Tattooing", and also translated as "the Shadowlands"), is a non-administrative geographical region of New Zealand in the south-western corner of the South Island, comprising the western third of Southland. Most of F ...
mountains from western Otago to
Fox Glacier Fox Glacier (; officially Fox Glacier / Te Moeka o Tuawe) is a temperate maritime glacier located in Westland Tai Poutini National Park on the West Coast of New Zealand's South Island. Like nearby Franz Josef Glacier, Fox Glacier is one of th ...
* ''Percnodaimon'' sp.nov. 2 – Pisa Range, Otago * ''Percnodaimon'' sp.nov. 3 – Eyre, Livingstone, and Takitiku Mountains to the south of Lake Wakitipu * ''Percnodaimon'' sp.nov. 4 – Mountains of
South Canterbury South Canterbury is the area of the Canterbury Region of the South Island of New Zealand bounded by the Rangitata River in the north and the Waitaki River (the border with the Otago Region) to the south. The Pacific Ocean and ridge of the S ...
and
North Otago North Otago is an area in New Zealand that covers the area of the Otago region between Shag Point and the Waitaki River, and extends inland to the west as far as the village of Omarama (which has experienced rapid growth as a developing centre f ...
Although the Patricks stated in 2012 that
nuclear DNA Nuclear DNA (nDNA), or nuclear deoxyribonucleic acid, is the DNA contained within each cell nucleus of a eukaryotic organism. It encodes for the majority of the genome in eukaryotes, with mitochondrial DNA and plastid DNA coding for the rest. ...
was being studied with a view to formally naming and describing these species, as of 2022 that had yet to happen, and ''P. merula'' is still considered the only species in the genus. Its
Māori Māori or Maori can refer to: Relating to the Māori people * Māori people of New Zealand, or members of that group * Māori language, the language of the Māori people of New Zealand * Māori culture * Cook Islanders, the Māori people of the Co ...
name, ''pepe pouri'', means "dark moth", and is applied to several ringlet species, including the forest ringlet ('' Dodonidia helmsii'') and Butler's ringlet ('' Erebiola butleri'').


Description

The wingspan of the ''Percnodaimon''
imago In biology, the imago (Latin for "image") is the last stage an insect attains during its metamorphosis, its process of growth and development; it is also called the ''imaginal'' stage ("imaginal" being "imago" in adjective form), the stage in wh ...
can be between 35 and 55 mm, and its broad velvety wings are usually black or brown across the dorsal surface, sometimes with a purple reflection. Forewings have characteristic black and white circles inside a patch of brown at their distal ends. There is little sexual dimorphism: males and females are similar in appearance, with females a little larger. The row of large pale spots often found on the underside of the hindwings is more noticeable in females. Black mountain ringlets fly slowly in zigzag patterns, especially on a sunny day, taking advantage of
thermals A thermal column (or thermal) is a rising mass of buoyant air, a convective current in the atmosphere, that transfers heat energy vertically. Thermals are created by the uneven heating of Earth's surface from solar radiation, and are an example ...
and often gliding in a "V" position.
"When disturbed this insect flies often with considerable rapidity and thus often eludes the net, so that the capture of a good series of specimens on a rugged mountain-slope is usually very exciting, if not actually dangerous work." — George V. Hudson (1928)
Wing colour plays an important physiological role: the dark wings with a large surface area absorb sunlight and store heat, essentially acting like
solar panels A solar panel is a device that converts sunlight into electricity by using photovoltaic (PV) cells. PV cells are made of materials that produce excited electrons when exposed to light. These electrons flow through a circuit and produce direct ...
, an adaptation to the cool climate of the
Southern Alps The Southern Alps (; officially Southern Alps / Kā Tiritiri o te Moana) are a mountain range extending along much of the length of New Zealand, New Zealand's South Island, reaching its greatest elevations near the range's western side. The n ...
. The butterfly sits with its wings open and turned towards the sun, flying while the sun is shining and hiding amongst rocks when clouds intervene. At night they shelter deep in rocky crevices.


Distribution

The genus ''Percnodaimon'' is restricted to the dry eastern mountains in the South Island of New Zealand, on
greywacke Greywacke or graywacke ( ) is a variety of sandstone generally characterized by its hardness (6–7 on Mohs scale), dark color, and Sorting (sediment), poorly sorted angular grains of quartz, feldspar, and small rock fragments or sand-size Lith ...
scree Scree is a collection of broken rock fragments at the base of a cliff or other steep rocky mass that has accumulated through periodic rockfall. Landforms associated with these materials are often called talus deposits. The term ''scree'' is ap ...
slopes and amongst rockfalls. It occurs at altitudes ranging from 800 to 2500 metres, up to 3100 m in the northern mountains of the South Island. Although the butterfly is less common below 1200 m, it can be found down to 800 m on valley floors and passes if the scree habitat is present. It can be quite common and even seasonally abundant in suitable habitat.


Life cycle


Ovum

The adult female black mountain ringlet deposits her eggs atop stones on alpine slopes, rather than on a larval food plant. When the sun is out it heats the stone's exposed surface, incubating the eggs. This behaviour is unique among New Zealand butterflies and is only seen in a handful of butterfly species occupying the Himalayas and European Alps. The eggs are initially blue, eventually turning speckled and brown, which allows them to blend in with the stony background. About two days before hatching this colouration is lost and the brown head of the larvae becomes visible. The hatching process takes around 12 days, or longer if the eggs are at higher altitudes. To sustain itself the newly-hatched larva consumes its egg casings, which have sufficient nutrients for early larval stages.


Larvae

''Percnodaimon'' larvae vary from dull grey to brown and have black anterior
setae In biology, setae (; seta ; ) are any of a number of different bristle- or hair-like structures on living organisms. Animal setae Protostomes Depending partly on their form and function, protostome setae may be called macrotrichia, chaetae ...
. The larvae have five
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, each of which take about a month in pleasant summer temperatures, but up to eight months over winter. For this reason the butterfly can stay in its larval stage for up to two years, and can be found almost year-round in an instar stage. The alpine grasses ''
Poa colensoi ''Poa colensoi'', the blue tussock, is a species of cool-season grass in the family Poaceae, endemic to New Zealand. It is considered the native grass species with the highest potential for use in high altitude livestock grazing In agricult ...
'', '' P. buchananii'', and other ''
Poa ''Poa'' is a genus of about 570 species of Poaceae, grasses, native to the temperate regions of both hemispheres. Common names include meadow-grass (mainly in Europe and Asia), bluegrass (mainly in North America), tussock (some New Zealand spe ...
'' species are the larval food plant. These tussocks are common throughout the Southern Alps, but larvae are present only where the grass grows adjacent to rocky areas. Black mountain ringlet larvae are night feeders to avoid predators, which can easily spot them feeding on the tips of tussock blades. They spend little time feeding, but instead conceal themselves in nearby hiding spots.


Pupa

At the start of pupation the black mountain ringlet turns grey with brown speckles to match the surrounding stones. Unlike most butterflies pupae are suspended horizontally rather than vertically, and are attached to the undersurface of a rock with a large cremaster, a hook-shaped protuberance on the abdomen. Pupae have been found up to 3 feet away from the nearest food plant. The adult butterflies emerge 2 to 3 weeks later.


Imago

Adult black mountain ringlets are found in summer, from December to February, and the Patricks note that early versus late emergence is one of the characteristics distinguishing the undescribed species. This species is considered to live about a year in its adult stage, but as the larvae can take two or three summers to grow to full size there is a possibility that it can live up to four. This slow growth may reflect the extreme climate and conditions of New Zealand's Southern Alps.


See also

*
Butterflies of New Zealand Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossil ...


References


External links

*Black mountain ringlet discussed on
RNZ Radio New Zealand (), commonly known as RNZ or Radio NZ, is a New Zealand public service broadcaster and Crown entity. Established under the Radio New Zealand Act 1995, it operates news and current affairs station, RNZ National, and a classica ...
''
Critter of the Week ''Critter of the Week'' is a weekly RNZ National programme about endangered and neglected native plants and animals of New Zealand. Beginning in 2015, ''Critter of the Week'' is an approximately 15-minute discussion between Nicola Toki (origi ...
''
2 December 2022
{{Taxonbar, from=Q1758376 Satyrini Butterflies of New Zealand Monotypic butterfly genera Taxa named by Arthur Gardiner Butler