Pettalidae
Encyclopedia
The Pettalidae are a family of harvestmen with 52 described species
Species
In biology, a species is one of the basic units of biological classification and a taxonomic rank. A species is often defined as a group of organisms capable of interbreeding and producing fertile offspring. While in many cases this definition is adequate, more precise or differing measures are...

 and subspecies in eleven genera
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...

. Several undescribed species are known or assumed in some genera.

Name

Pettalus is a name from Greek mythology
Greek mythology
Greek mythology is the body of myths and legends belonging to the ancient Greeks, concerning their gods and heroes, the nature of the world, and the origins and significance of their own cult and ritual practices. They were a part of religion in ancient Greece...

 that appears in Ovid
Ovid
Publius Ovidius Naso , known as Ovid in the English-speaking world, was a Roman poet who is best known as the author of the three major collections of erotic poetry: Heroides, Amores, and Ars Amatoria...

's Metamorphoses.

Description

All species except the cave-dwelling South African Speleosiro argasiformis spend their entire life cycle in leaf litter.

They are two to five millimeters long, usually with an oval shaped body.

Although all Pettalidae except Parapurcellia have eyes, these were long thought to be absent in the family, mainly because they cannot be seen by Scanning Electron Microscopy
Scanning electron microscope
A scanning electron microscope is a type of electron microscope that images a sample by scanning it with a high-energy beam of electrons in a raster scan pattern...

. They are often incorporated at the base of the ozophore
Ozophore
An ozophore is an elevated cone present in the harvestman suborder Cyphophthalmi. It carries the openings of the defensive glands, called ozopores, which are present in many harvestmen....

s and typically lack lenses.

Distribution

The members of this family are distributed throughout former temperate Gondwana
Gondwana
In paleogeography, Gondwana , originally Gondwanaland, was the southernmost of two supercontinents that later became parts of the Pangaea supercontinent. It existed from approximately 510 to 180 million years ago . Gondwana is believed to have sutured between ca. 570 and 510 Mya,...

, with genera in Chile
Chile
Chile ,officially the Republic of Chile , is a country in South America occupying a long, narrow coastal strip between the Andes mountains to the east and the Pacific Ocean to the west. It borders Peru to the north, Bolivia to the northeast, Argentina to the east, and the Drake Passage in the far...

, South Africa
South Africa
The Republic of South Africa is a country in southern Africa. Located at the southern tip of Africa, it is divided into nine provinces, with of coastline on the Atlantic and Indian oceans...

, Madagascar
Madagascar
The Republic of Madagascar is an island country located in the Indian Ocean off the southeastern coast of Africa...

, Sri Lanka
Sri Lanka
Sri Lanka, officially the Democratic Socialist Republic of Sri Lanka is a country off the southern coast of the Indian subcontinent. Known until 1972 as Ceylon , Sri Lanka is an island surrounded by the Indian Ocean, the Gulf of Mannar and the Palk Strait, and lies in the vicinity of India and the...

, eastern and western Australia
Australia
Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

, and New Zealand
New Zealand
New Zealand is an island country in the south-western Pacific Ocean comprising two main landmasses and numerous smaller islands. The country is situated some east of Australia across the Tasman Sea, and roughly south of the Pacific island nations of New Caledonia, Fiji, and Tonga...

, where they are most diverse by far, with 29 species and subspecies found in three genera.

Relationships


Phylogeny of most Pettalidae

(after Boyer & Giribet 2007)

The family Pettalidae is monophyletic, although it is at the moment (2007) unclear what the nearest relatives are. It probably originated in the southern part of Gondwana. Parsimony analysis
Maximum parsimony
Parsimony is a non-parametric statistical method commonly used in computational phylogenetics for estimating phylogenies. Under parsimony, the preferred phylogenetic tree is the tree that requires the least evolutionary change to explain some observed data....

 suggests it could be a sister group to the remaining Cyphophthalmi
Cyphophthalmi
The Cyphophthalmi are a suborder of harvestmen, with about 36 genera, and more than hundred described species.The six families are currently grouped into two infraorders, the Tropicophthalmi and the Temperophthalmi; however, these are not supported by modern phylogenetic analysis.They are smaller...

, though this could also be the case for the Stylocellidae
Stylocellidae
The Stylocellidae are a family of harvestmen with about 30 described species, all of which occur from India to New Guinea. Members of this family are from one to seven millimeters long...

, or it could be related to the Sironidae
Sironidae
The Sironidae are a family of harvestmen with more than 30 described species.The family shows a Laurasian distribution, with most species found in temperate Europe and the west coast of North America...

, or specifically to the sironid genus Suzukielus. It is unrelated to the Troglosironidae
Troglosironidae
The Troglosironidae are a family of harvestmen with thirteen described species in a single genus, Troglosiro. Several species, including six undescribed ones, were collected around the island, and more are expected to be found....

 that are endemic
Endemic (ecology)
Endemism is the ecological state of being unique to a defined geographic location, such as an island, nation or other defined zone, or habitat type; organisms that are indigenous to a place are not endemic to it if they are also found elsewhere. For example, all species of lemur are endemic to the...

 to New Caledonia
New Caledonia
New Caledonia is a special collectivity of France located in the southwest Pacific Ocean, east of Australia and about from Metropolitan France. The archipelago, part of the Melanesia subregion, includes the main island of Grande Terre, the Loyalty Islands, the Belep archipelago, the Isle of...

.

The main lineages of the family may have arisen rapidly, possibly during the rapid expansion of Glossopteris
Glossopteris
Glossopteris is the largest and best-known genus of the extinct order of seed ferns known as Glossopteridales ....

forests that were predominant in temperate Gondwana. Pettalidae were likely present throughout the forests of Antarctica, which formed a land bridge between Australia and South America up until circa 50 million years ago (mya).

The Australian genera Austropurcellia (Eastern Australia: Queensland) and Karripurcellia (Western Australia) are not sister groups. It is possible that the Cyphophthalmi dispersed across Australia while the central region was covered with Nothofagus
Nothofagus
Nothofagus, also known as the southern beeches, is a genus of 35 species of trees and shrubs native to the temperate oceanic to tropical Southern Hemisphere in southern South America and Australasia...

rainforest (until 37 mya), or that the ancestors of the two genera independently dispersed from adjacent landmasses now separate from Australia.

Parapurcellia from eastern South Africa is sister to all other Pettalidae, while Purcellia from western South Africa is sister to the Chilean Chileogovea. Western South Africa and southern South America were last connected during the Late Jurassic
Late Jurassic
The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 161.2 ± 4.0 to 145.5 ± 4.0 million years ago , which is preserved in Upper Jurassic strata. In European lithostratigraphy, the name "Malm" indicates rocks of Late Jurassic age...

, about 150 mya. Likewise, the monotypic Neopurcellia from New Zealand appears as the sister group to all Pettalidae except for Parapurcellia, instead of being monophyletic with the other two New Zealand genera, which themselves appear as sister groups in Bayesian analysis, but not in direct optimization parsimony analyses.

Species

  • Aoraki Boyer & Giribet, 2007 (New Zealand
    New Zealand
    New Zealand is an island country in the south-western Pacific Ocean comprising two main landmasses and numerous smaller islands. The country is situated some east of Australia across the Tasman Sea, and roughly south of the Pacific island nations of New Caledonia, Fiji, and Tonga...

    )
  • Aoraki calcarobtusa (Forster
    Raymond Robert Forster
    Raymond Robert Forster was a spider expert.He wrote his first paper on spiders at the age of 17. He studied at Victoria University...

    , 1952)
    Wellington
    Wellington
    Wellington is the capital city and third most populous urban area of New Zealand, although it is likely to have surpassed Christchurch due to the exodus following the Canterbury Earthquake. It is at the southwestern tip of the North Island, between Cook Strait and the Rimutaka Range...

    , North Island
    North Island
    The North Island is one of the two main islands of New Zealand, separated from the much less populous South Island by Cook Strait. The island is in area, making it the world's 14th-largest island...

  • Aoraki calcarobtusa westlandica (Forster, 1952)Buller
    Buller District
    200px|rightBuller District, administered by the Buller District Council is an administrative region in the West Coast Region of New Zealand. It covers Westport, Karamea, Reefton and Inangahua Junction. Its land area is 7,953.12 km²...

    , South Island
    South Island
    The South Island is the larger of the two major islands of New Zealand, the other being the more populous North Island. It is bordered to the north by Cook Strait, to the west by the Tasman Sea, to the south and east by the Pacific Ocean...

    • Aoraki crypta (Forster, 1948)Coromandel
      Coromandel Peninsula
      The Coromandel Peninsula lies in the North Island of New Zealand. It is part of the Waikato Region and Thames-Coromandel District and extends 85 kilometres north from the western end of the Bay of Plenty, forming a natural barrier to protect the Hauraki Gulf and the Firth of Thames in the west...

       & Bay of Plenty
      Bay of Plenty
      The Bay of Plenty , often abbreviated to BOP, is a region in the North Island of New Zealand situated around the body of water of the same name...

      , North Island
    • Aoraki denticulata (Forster, 1948)Nelson
      Nelson, New Zealand
      Nelson is a city on the eastern shores of Tasman Bay, and is the economic and cultural centre of the Nelson-Tasman region. Established in 1841, it is the second oldest settled city in New Zealand and the oldest in the South Island....

      , Marlborough
      Marlborough, New Zealand
      Marlborough is one of the regions of New Zealand, located in the northeast of the South Island. Marlborough is a unitary authority, both a region and a district, and its council is located at Blenheim. Marlborough is known for its dry climate, the picturesque Marlborough Sounds, and sauvignon blanc...

      , and Buller, South Island
  • Aoraki denticulata major (Forster, 1952)Arthur's Pass
    Arthur's Pass
    Arthur's Pass is a mountain pass in the Southern Alps of the South Island of New Zealand. It marks part of the boundary between the West Coast and Canterbury regions, 140 km from Christchurch and 95 km from Greymouth. The pass lies in a saddle between the valleys of the Otira River, a...

    , South Island
    • Aoraki granulosa (Forster, 1952)Waikato
      Waikato
      The Waikato Region is a local government region of the upper North Island of New Zealand. It covers the Waikato, Hauraki, Coromandel Peninsula, the northern King Country, much of the Taupo District, and parts of Rotorua District...

      , Coromandel, and Bay of Plenty, North Island
    • Aoraki healyi (Forster, 1948) — Marlborough and Marlborough Sounds
      Marlborough Sounds
      The Marlborough Sounds are an extensive network of sea-drowned valleys created by a combination of land subsidence and rising sea levels at the north of the South Island of New Zealand...

      , South Island
    • Aoraki inerma (Forster, 1948) — Waikato, Tongariro, and Bay of Plenty, North Island
  • Aoraki inerma stephenensis (Forster, 1952)Stephen's Island
    Stephens Island, New Zealand
    Stephens Island is at the northern most tip of the Marlborough Sounds in the South Island of New Zealand. It lies two kilometres to the northeast of Cape Stephens, the northernmost point of D'Urville Island. The Māori call the island Takapourewa but Stephens Island is the commonly used name...

    , Marlborough, South Island
    • Aoraki longitarsa (Forster, 1952) — Mount Cook, Canterbury
      Canterbury, New Zealand
      The New Zealand region of Canterbury is mainly composed of the Canterbury Plains and the surrounding mountains. Its main city, Christchurch, hosts the main office of the Christchurch City Council, the Canterbury Regional Council - called Environment Canterbury - and the University of Canterbury.-...

      , South Island
    • Aoraki tumidata (Forster, 1948)Cuvier Island
      Cuvier Island
      Cuvier Island is a small island off the east coast of the North Island of New Zealand. It lies on the seaward end of the Colville Channel, north of the Mercury Islands and approximately south-east of Great Barrier Island...

      , Coromandel, North Island

  • Austropurcellia Shear, 1980 (Queensland
    Queensland
    Queensland is a state of Australia, occupying the north-eastern section of the mainland continent. It is bordered by the Northern Territory, South Australia and New South Wales to the west, south-west and south respectively. To the east, Queensland is bordered by the Coral Sea and Pacific Ocean...

    , Australia
    Australia
    Australia , officially the Commonwealth of Australia, is a country in the Southern Hemisphere comprising the mainland of the Australian continent, the island of Tasmania, and numerous smaller islands in the Indian and Pacific Oceans. It is the world's sixth-largest country by total area...

    )
  • Austropurcellia arcticosa (Cantrell, 1980)
  • Austropurcellia capricornia (Todd Davies, 1977)
  • Austropurcellia daviesae (Juberthie, 1989)
  • Austropurcellia forsteri (Juberthie, 2000)
  • Austropurcellia scoparia Juberthie, 1988
  • Austropurcellia woodwardi (Forster, 1955)

  • Chileogovea Roewer, 1961 (Chile
    Chile
    Chile ,officially the Republic of Chile , is a country in South America occupying a long, narrow coastal strip between the Andes mountains to the east and the Pacific Ocean to the west. It borders Peru to the north, Bolivia to the northeast, Argentina to the east, and the Drake Passage in the far...

    )
  • Chileogovea jocasta Shear, 1993
  • Chileogovea oedipus Roewer
    Carl Friedrich Roewer
    Carl Friedrich Roewer was a German arachnologist. He concentrated on harvestmen, where he described almost a third of today's known species, but also almost 700 taxa of spiders and numerous Solifugae....

    , 1961

  • Karripurcellia Giribet, 2003 (Western Australia
    Western Australia
    Western Australia is a state of Australia, occupying the entire western third of the Australian continent. It is bounded by the Indian Ocean to the north and west, the Great Australian Bight and Indian Ocean to the south, the Northern Territory to the north-east and South Australia to the south-east...

    )
  • Karripurcellia harveyi Giribet, 2003
  • Karripurcellia peckorum Giribet, 2003
  • Karripurcellia sierwaldae Giribet, 2003

  • Manangotria Shear & Gruber, 1996 (Madagascar
    Madagascar
    The Republic of Madagascar is an island country located in the Indian Ocean off the southeastern coast of Africa...

    )
  • Manangotria taolanaro Shear & Gruber, 1996

  • Neopurcellia Forster, 1948 (New Zealand: South Island)
  • Neopurcellia salmoni Forster, 1948

  • Parapurcellia Rosas Costa, 1950 (eastern South Africa
    South Africa
    The Republic of South Africa is a country in southern Africa. Located at the southern tip of Africa, it is divided into nine provinces, with of coastline on the Atlantic and Indian oceans...

    )
  • Parapurcellia fissa (Lawrence, 1939)
  • Parapurcellia monticola (Lawrence, 1939)
  • Parapurcellia rumpiana (Lawrence, 1933)
  • Parapurcellia silvicola (Lawrence, 1939)

  • Pettalus Thorell, 1876 (Sri Lanka
    Sri Lanka
    Sri Lanka, officially the Democratic Socialist Republic of Sri Lanka is a country off the southern coast of the Indian subcontinent. Known until 1972 as Ceylon , Sri Lanka is an island surrounded by the Indian Ocean, the Gulf of Mannar and the Palk Strait, and lies in the vicinity of India and the...

    )
  • Pettalus brevicauda Pocock
    Reginald Innes Pocock
    Reginald Innes Pocock F.R.S. was a British zoologist.Pocock was born in Clifton, Bristol, the fourth son of Rev. Nicholas Pocock and Edith Prichard. He began showing interest in natural history at St. Edward's School, Oxford. He received tutoring in zoology from Sir Edward Poulton, and was allowed...

    , 1897
  • Pettalus cimiciformis (O. P-Cambridge
    Octavius Pickard-Cambridge
    The Reverend Octavius Pickard-Cambridge FRS was an English clergyman and zoologist.Pickard-Cambridge was born in Bloxworth rectory, Dorset, the fifth son of Revd George Pickard, rector and squire of Bloxworth: the family changed their name to Pickard-Cambridge in 1848...

    , 1875)
  • Pettalus lampetides Sharma & Giribet, 2006

  • Purcellia Hansen & Sørensen, 1904 (western South Africa)
  • Purcellia illustrans Hansen & Sørensen, 1904
  • Purcellia peregrinator Lawrence, 1963
  • Purcellia transvaalica Lawrence, 1963

  • Rakaia Hirst, 1925 (New Zealand)
  • Rakaia antipodiana Hirst, 1925Rakaia Gorge
    Rakaia Gorge
    The Rakaia Gorge is located on the Rakaia River in inland Canterbury in New Zealand's South Island.Like its neighbour, the Waimakariri River, the Rakaia runs through wide shingle beds for much of its length, but is forced through a narrow canyon as it approaches the Canterbury Plains.In the 1870s,...

    , Canterbury, South Island
  • Rakaia dorothea (Phillipps & Grimmett, 1932) — Wellington and Wairarapa
    Wairarapa
    Wairarapa is a geographical region of New Zealand. It occupies the south-eastern corner of the North Island, east of metropolitan Wellington and south-west of the Hawke's Bay region. It is lightly populated, having several rural service towns, with Masterton being the largest...

    , North Island
  • Rakaia florensis (Forster, 1948) — Nelson, South Island
  • Rakaia isolata Forster, 1952 — North Canterbury, South Island
  • Rakaia lindsayi Forster, 1952 — Stewart Island
  • Rakaia macra Boyer & Giribet, 2003Waipori
    Waipori
    Waipori may refer to either:*Waipori, New Zealand, a township within Dunedin, New Zealand's city limits* The Waipori River in Otago in the South Island of New Zealand, or to* Lake Waipori, an area of wetlands draining into the Waipori River...

    , Otago
    Otago
    Otago is a region of New Zealand in the south of the South Island. The region covers an area of approximately making it the country's second largest region. The population of Otago is...

    , South Island
  • Rakaia magna Forster, 1948 — Wellington, North Island
  • Rakaia magna australis Forster, 1952Lewis Pass
    Lewis Pass
    Lewis Pass is a mountain pass in the South Island of New Zealand.The northernmost of the three main passes across the Southern Alps, it is higher than the Haast Pass, but slightly lower than Arthur's Pass...

    , South Island
    • Rakaia media Forster, 1948 — Tongariro and Bay of Plenty, North Island
  • Rakaia media insula Forster, 1952 — Little Barrier Island, Auckland, North Island
    • Rakaia minutissima (Forster, 1948) — Waikato, South Island
    • Rakaia pauli Forster, 1952 — Mid-Canterbury, South Island
    • Rakaia solitaria Forster, 1948 — Opouawa Gully, West Wairarapa, North Island
    • Rakaia sorenseni Forster, 1952Southland
      Southland Region
      Southland is New Zealand's southernmost region and is also a district within that region. It consists mainly of the southwestern portion of the South Island and Stewart Island / Rakiura...

       and Dunedin
      Dunedin
      Dunedin is the second-largest city in the South Island of New Zealand, and the principal city of the Otago Region. It is considered to be one of the four main urban centres of New Zealand for historic, cultural, and geographic reasons. Dunedin was the largest city by territorial land area until...

      , South Island
  • Rakaia sorenseni digitata Forster, 1952 — Chaslands, Otago, South Island
    • Rakaia stewartiensis Forster, 1948 — Stewart Island
    • Rakaia uniloca Forster, 1952 — Marlborough Sounds, South Island

  • Speleosiro Lawrence, 1931 (South Africa)
  • Speleosiro argasiformis Lawrence, 1931


The monotypic Ankaratra franzi Shear & Gruber, 1996 from Madagascar is sometimes placed in this family, but is as of 2007 considered incertae sedis
Incertae sedis
, is a term used to define a taxonomic group where its broader relationships are unknown or undefined. Uncertainty at specific taxonomic levels is attributed by , , and similar terms.-Examples:*The fossil plant Paradinandra suecica could not be assigned to any...

within the Cyphophthalmi.
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