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The Pettalidae are a family of
harvestmen The Opiliones (formerly Phalangida) are an Order (biology), order of arachnids Common name, colloquially known as harvestmen, harvesters, harvest spiders, or daddy longlegs. , over 6,650 species of harvestmen have been discovered worldwide, alth ...
with 75 described
species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of ...
in 10
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial ...
.Boyer & Giribet 2007 Several undescribed species are known or assumed in some genera. (2007): Pettalidae Shear, 1980. In: Pinto-da-Rocha ''et al.'' 2007: 99ff


Name

''Pettalus'' is a name from
Greek mythology A major branch of classical mythology, Greek mythology is the body of myths originally told by the ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the origin and nature of the world, the lives and activities of ...
that appears in
Ovid Pūblius Ovidius Nāsō (; 20 March 43 BC – 17/18 AD), known in English as Ovid ( ), was a Roman poet who lived during the reign of Augustus. He was a contemporary of the older Virgil and Horace, with whom he is often ranked as one of the ...
's
Metamorphoses The ''Metamorphoses'' ( la, Metamorphōsēs, from grc, μεταμορφώσεις: "Transformations") is a Latin narrative poem from 8 CE by the Roman poet Ovid. It is considered his '' magnum opus''. The poem chronicles the history of the ...
.


Description

All species except the cave-dwelling South African ''Speleosiro argasiformis'' spend their entire life cycle in
leaf litter Plant litter (also leaf litter, tree litter, soil litter, litterfall or duff) is dead plant material (such as leaves, bark, needles, twigs, and cladodes) that have fallen to the ground. This detritus or dead organic material and its constituent ...
. 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. They are often incorporated at the base of the
ozophore An ozophore is an elevated cone present in the harvestman ("daddy long-legs") suborder Cyphophthalmi. It carries the openings, called ozopores, of the defensive glandsPinto-da-Rocha ''et al.'' 2007: 22f that are present in many harvestmen. The nam ...
s and typically lack lenses.


Distribution

The members of this family are distributed throughout former temperate
Gondwana Gondwana () was a large landmass, often referred to as a supercontinent, that formed during the late Neoproterozoic (about 550 million years ago) and began to break up during the Jurassic period (about 180 million years ago). The final st ...
, with genera in
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the eas ...
,
South Africa South Africa, officially the Republic of South Africa (RSA), is the southernmost country in Africa. It is bounded to the south by of coastline that stretch along the South Atlantic and Indian Oceans; to the north by the neighbouring count ...
,
Madagascar Madagascar (; mg, Madagasikara, ), officially the Republic of Madagascar ( mg, Repoblikan'i Madagasikara, links=no, ; french: République de Madagascar), is an island country in the Indian Ocean, approximately off the coast of East Africa ...
,
Sri Lanka Sri Lanka (, ; si, ශ්‍රී ලංකා, Śrī Laṅkā, translit-std=ISO (); ta, இலங்கை, Ilaṅkai, translit-std=ISO ()), formerly known as Ceylon and officially the Democratic Socialist Republic of Sri Lanka, is an ...
, eastern and western Australia, and
New Zealand New Zealand ( mi, Aotearoa ) is an island country in the southwestern Pacific Ocean. It consists of two main landmasses—the North Island () and the South Island ()—and over 700 List of islands of New Zealand, smaller islands. It is the ...
, where they are most diverse by far, with 29 species and subspecies found in three genera.


Relationships

The family Pettalidae is
monophyletic In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. 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 suggests it could be a sister group to the remaining
Cyphophthalmi Cyphophthalmi is a suborder of harvestmen, colloquially known as mite harvestmen. Cyphophthalmi comprises 36 genera, and more than two hundred described species. The six families are currently grouped into three infraorders: the Boreophthalmi, Sc ...
, though this could also be the case for the Stylocellidae, or it could be related to the Sironidae, or specifically to the sironid genus '' Suzukielus''. It is unrelated to the Troglosironidae that are
endemic Endemism is the state of a species being found 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 found els ...
to New Caledonia. The main lineages of the family may have arisen rapidly, possibly during the rapid expansion of ''
Glossopteris ''Glossopteris'' tymology: from Ancient Greek γλῶσσα (glôssa, " tongue ") + πτερίς (pterís, " fern ")is the largest and best-known genus of the extinct Permian order of seed ferns known as Glossopteridales (also known as Arber ...
'' forests that were predominant in temperate Gondwana. Pettalidae were likely present throughout the forests of
Antarctica Antarctica () is Earth's southernmost and least-populated continent. Situated almost entirely south of the Antarctic Circle and surrounded by the Southern Ocean, it contains the geographic South Pole. Antarctica is the fifth-largest co ...
, which formed a land bridge between Australia and South America up until circa 50
million years ago The abbreviation Myr, "million years", is a unit of a quantity of (i.e. ) years, or 31.556926 teraseconds. Usage Myr (million years) is in common use in fields such as Earth science and cosmology. Myr is also used with Mya (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'', also known as the southern beeches, is a genus of 43 species of trees and shrubs native to the Southern Hemisphere in southern South America (Chile, Argentina) and Australasia (east and southeast Australia, New Zealand, New Gu ...
'' 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 The Late Jurassic is the third epoch of the Jurassic Period, and it spans the geologic time from 163.5 ± 1.0 to 145.0 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic strata.Owen 1987. In European lithostratigraphy, the ...
, 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 Bayesian inference is a method of statistical inference in which Bayes' theorem is used to update the probability for a hypothesis as more evidence or information becomes available. Bayesian inference is an important technique in statistics, and ...
, but not in direct optimization parsimony analyses.


Species

The family Pettalidae contains the following genera and species: * ''
Aoraki Aoraki / Mount Cook is the highest mountain in New Zealand. Its height, as of 2014, is listed as . It sits in the Southern Alps, the mountain range that runs the length of the South Island. A popular tourist destination, it is also a favourit ...
'' Boyer & Giribet, 2007 (New Zealand) :* ''
Aoraki calcarobtusa Aoraki / Mount Cook is the highest mountain in New Zealand. Its height, as of 2014, is listed as . It sits in the Southern Alps, the mountain range that runs the length of the South Island. A popular tourist destination, it is also a favouri ...
'' Forster, 1952 ::* ''Aoraki calcarobtusa calcarobtusa'' Forster, 1952 ::* ''Aoraki calcarobtusa westlandica'' Forster, 1952 :* ''
Aoraki crypta Aoraki / Mount Cook is the highest mountain in New Zealand. Its height, as of 2014, is listed as . It sits in the Southern Alps, the mountain range that runs the length of the South Island. A popular tourist destination, it is also a favouri ...
'' Forster, 1948 :* ''
Aoraki denticulata Aoraki / Mount Cook is the highest mountain in New Zealand. Its height, as of 2014, is listed as . It sits in the Southern Alps, the mountain range that runs the length of the South Island. A popular tourist destination, it is also a favouri ...
'' Forster, 1948 ::* ''Aoraki denticulata denticulata'' Forster, 1948 ::* ''Aoraki denticulata major'' Forster, 1952 :* ''
Aoraki granulosa Aoraki / Mount Cook is the highest mountain in New Zealand. Its height, as of 2014, is listed as . It sits in the Southern Alps, the mountain range that runs the length of the South Island. A popular tourist destination, it is also a favouri ...
'' Forster, 1952 :* '' Aoraki healyi'' Forster, 1948 :* ''
Aoraki inerma Aoraki / Mount Cook is the highest mountain in New Zealand. Its height, as of 2014, is listed as . It sits in the Southern Alps, the mountain range that runs the length of the South Island. A popular tourist destination, it is also a favouri ...
'' Forster, 1948 ::* ''Aoraki inerma inerma'' Forster, 1948 ::* ''Aoraki inerma stephenensis'' Forster, 1952 :* ''
Aoraki longitarsa Aoraki / Mount Cook is the highest mountain in New Zealand. Its height, as of 2014, is listed as . It sits in the Southern Alps, the mountain range that runs the length of the South Island. A popular tourist destination, it is also a favourite ...
'' Forster, 1952 :* ''
Aoraki tumidata Aoraki / Mount Cook is the highest mountain in New Zealand. Its height, as of 2014, is listed as . It sits in the Southern Alps, the mountain range that runs the length of the South Island. A popular tourist destination, it is also a favouri ...
'' Forster, 1948 * '' Austropurcellia'' Shear, 1980 (Australia:
Queensland ) , nickname = Sunshine State , image_map = Queensland in Australia.svg , map_caption = Location of Queensland in Australia , subdivision_type = Country , subdivision_name = Australia , established_title = Before federation , established_ ...
) :* '' Austropurcellia absens'' Boyer & Popkin-Hall, 2015 :* '' Austropurcellia acuta'' Popkin-Hall & Boyer, 2014 :* '' Austropurcellia alata'' Boyer & Reuter, 2012 :* '' Austropurcellia arcticosa'' Cantrell, 1980 :* '' Austropurcellia barbata'' Popkin-Hall & Boyer, 2014 :* '' Austropurcellia cadens'' Baker & Boyer, 2015 :* '' Austropurcellia capricornia'' Todd Davies, 1977 :* '' Austropurcellia clousi'' Boyer, Baker & Popkin-Hall, 2015 :* '' Austropurcellia culminis'' Boyer & Reuter, 2012 :* '' Austropurcellia daviesae'' Juberthie, 1989 :* '' Austropurcellia despectata'' Boyer & Reuter, 2012 :* '' Austropurcellia finniganensis'' Popkin-Hall, Jay & Boyer, 2016 :* '' Austropurcellia forsteri'' Juberthie, 2000 :* '' Austropurcellia fragosa'' Popkin-Hall, Jay & Boyer, 2016 :* '' Austropurcellia giribeti'' Boyer & Quay, 2015 :* '' Austropurcellia megatanka'' Jay, Coblens & Boyer, 2016 :* '' Austropurcellia monteithi'' Jay, Popkin-Hall, Coblens & Boyer, 2016 :* '' Austropurcellia nuda'' Popkin-Hall, Jay & Boyer, 2016 :* '' Austropurcellia riedeli'' Jay, Coblens & Boyer, 2016 :* '' Austropurcellia scoparia'' Juberthie, 1988 :* '' Austropurcellia sharmai'' Boyer & Quay, 2015 :* '' Austropurcellia superbensis'' Popkin-Hall & Boyer, 2014 :* '' Austropurcellia tholei'' Baker & Boyer, 2015 :* '' Austropurcellia vicina'' Boyer & Reuter, 2012 :* '' Austropurcellia woodwardi'' Forster, 1955 * '' Chileogovea'' Roewer, 1961 (
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the eas ...
) :* '' Chileogovea jocasta'' Shear, 1993 :* '' Chileogovea oedipus'' Roewer, 1961 * '' Karripurcellia'' Giribet, 2003 (Australia:
Western Australia Western Australia (commonly abbreviated as WA) is a state of Australia occupying the western percent of the land area of Australia excluding external territories. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to ...
) :* '' Karripurcellia harveyi'' Giribet, 2003 :* '' Karripurcellia peckorum'' Giribet, 2003 :* '' Karripurcellia sierwaldae'' Giribet, 2003 * '' Manangotria'' Shear & Gruber, 1996 (Madagascar) :* '' Manangotria taolanaro'' Shear & Gruber, 1996 * '' Neopurcellia'' Forster, 1948 (New Zealand: South Island) :* '' Neopurcellia salmoni'' Forster, 1948 * '' Parapurcellia'' Rosas Costa, 1950 (eastern South Africa) :* '' Parapurcellia amatola'' de Bivort & Giribet, 2010 :* '' Parapurcellia convexa'' de Bivort & Giribet, 2010 :* '' Parapurcellia fissa'' Lawrence, 1939 :* '' Parapurcellia minuta''de Bivort & Giribet, 2010 :* '' Parapurcellia monticola'' Lawrence, 1939 :* '' Parapurcellia natalia'' de Bivort & Giribet, 2010 :* '' Parapurcellia peregrinator'' Lawrence, 1963 :* '' Parapurcellia rumpiana'' Lawrence, 1933 :* '' Parapurcellia silvicola'' Lawrence, 1939 :* '' Parapurcellia staregai'' de Bivort & Giribet, 2010 * '' Pettalus'' Thorell, 1876 (Sri Lanka) :* '' Pettalus brevicauda'' Pocock, 1897 :* '' Pettalus cimiciformis'' O. P-Cambridge, 1875 :* '' Pettalus lampetides'' Sharma & Giribet, 2006 :* '' Pettalus thwaitesi'' Sharma, Karunarathna & Giribet, 2009 * '' Purcellia'' Hansen & Sørensen, 1904 (western South Africa) :* '' Purcellia argasiformis'' Lawrence, 1931 :* '' Purcellia griswoldi'' de Bivort & Giribet, 2010 :* '' Purcellia illustrans'' Hansen & Sørensen, 1904 :* '' Purcellia lawrencei'' de Bivort & Giribet, 2010 :* '' Purcellia leleupi'' Starega, 2008 :* '' Purcellia transvaalica'' Lawrence, 1963 * ''
Rakaia Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on State Highway 1 and the Main South Line. Immediately north o ...
'' Hirst, 1925 (New Zealand) :* ''
Rakaia antipodiana Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on State Highway 1 and the Main South Line. Immediately north of ...
'' Hirst, 1925 :* ''
Rakaia dorothea Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on State Highway 1 and the Main South Line. Immediately north of ...
'' Phillipps & Grimmett, 1932 :* ''
Rakaia florensis Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on New Zealand State Highway 1, State Highway 1 and the Main South Li ...
'' Forster, 1948 :* ''
Rakaia isolata Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on State Highway 1 and the Main South Line. Immediately north of ...
'' Forster, 1952 :* ''
Rakaia lindsayi Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on State Highway 1 and the Main South Line. Immediately north of ...
'' Forster, 1952 :* ''
Rakaia macra Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on State Highway 1 and the Main South Line. Immediately north of ...
'' Boyer & Giribet, 2003 :* ''
Rakaia magna Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on State Highway 1 and the Main South Line. Immediately north of ...
'' Forster, 1948 ::* ''Rakaia magna australis'' Forster, 1952 ::* ''Rakaia magna magna'' Forster, 1948 :* ''
Rakaia media Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on State Highway 1 and the Main South Line. Immediately north of ...
'' Forster, 1948 ::* ''Rakaia media insula'' Forster, 1952 ::* ''Rakaia media media'' Forster, 1948 :* ''
Rakaia minutissima Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on State Highway 1 and the Main South Line. Immediately north of ...
'' Forster, 1948 :* ''
Rakaia pauli Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on New Zealand State Highway 1, State Highway 1 and the Main South Li ...
'' Forster, 1952 :* ''
Rakaia solitaria Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on State Highway 1 and the Main South Line. Immediately north of ...
'' Forster, 1948 :* ''
Rakaia sorenseni Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on State Highway 1 and the Main South Line. Immediately north of ...
'' Forster, 1952 ::* ''Rakaia sorenseni digitata'' Forster, 1952 ::* ''Rakaia sorenseni sorenseni'' Forster, 1952 :* ''
Rakaia stewartiensis Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on State Highway 1 and the Main South Line. Immediately north of ...
'' Forster, 1948 :* ''
Rakaia uniloca Rakaia is a town seated close to the southern banks of the Rakaia River on the Canterbury Plains in New Zealand's South Island, approximately 57 km south of Christchurch on New Zealand State Highway 1, State Highway 1 and the Main South Li ...
'' Forster, 1952


See also

* Environment of Sri Lanka


Footnotes


References

* Joel Hallan's Biology Catalog
Pettalidae

Checklist of the Cyphophthalmi species of the World
(with pictures) * (1971): Les opilions cyphophthalmes cavernicoles. Notes sur ''Speleosiro argasiformis'' Lawrence. ''Bull. Mus. Natl Hist. Nat.'' 42: 864–871. * (eds.) (2007): Harvestmen - The Biology of Opiliones. ''Harvard University Press'' * (2007): A new model Gondwanan taxon: systematics and biogeography of the harvestman family Pettalidae (Arachnida, Opiliones, Cyphophthalmi), with a taxonomic revision of genera from Australia and New Zealand. ''Cladistics'' 23(4): 337-361. {{Taxonbar, from=Q733898 Harvestmen Harvestman families