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 (see below). , over 6,650 species of harvestmen have been discovered w ...

with 75 described

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

in 10

genera Genus (; : genera ) is a taxonomic rank above species and below family as used in the biological classification of living and fossil organisms as well as viruses. In binomial nomenclature, the genus name forms the first part of the binomial s ...

.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 Greek mythology is the body of myths originally told by the Ancient Greece, ancient Greeks, and a genre of ancient Greek folklore, today absorbed alongside Roman mythology into the broader designation of classical mythology. These stories conc ...

that appears in

Ovid Publius Ovidius Naso (; 20 March 43 BC – AD 17/18), known in English as Ovid ( ), was a Augustan literature (ancient Rome), Roman poet who lived during the reign of Augustus. He was a younger contemporary of Virgil and Horace, with whom he i ...

Metamorphoses The ''Metamorphoses'' (, , ) is a Latin Narrative poetry, narrative poem from 8 Common Era, CE by the Ancient Rome, Roman poet Ovid. It is considered his ''Masterpiece, magnum opus''. The poem chronicles the history of the world from its Cre ...

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 has fallen to the ground. This detritus or dead organic material and its constituen ...

. 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 A scanning electron microscope (SEM) is a type of electron microscope that produces images of a sample by scanning the surface with a focused beam of electrons. The electrons interact with atoms in the sample, producing various signals that ...

. 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. Th ...

s and typically lack lenses.

Distribution

The members of this family are distributed throughout former temperate

Gondwana Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...

, with genera in

Chile Chile, officially the Republic of Chile, is a country in western South America. It is the southernmost country in the world and the closest to Antarctica, stretching along a narrow strip of land between the Andes, Andes Mountains and the Paci ...

South Africa South Africa, officially the Republic of South Africa (RSA), is the Southern Africa, southernmost country in Africa. Its Provinces of South Africa, nine provinces are bounded to the south by of coastline that stretches along the Atlantic O ...

Madagascar Madagascar, officially the Republic of Madagascar, is an island country that includes the island of Madagascar and numerous smaller peripheral islands. Lying off the southeastern coast of Africa, it is the world's List of islands by area, f ...

Sri Lanka Sri Lanka, officially the Democratic Socialist Republic of Sri Lanka, also known historically as Ceylon, is an island country in South Asia. It lies in the Indian Ocean, southwest of the Bay of Bengal, separated from the Indian subcontinent, ...

, eastern and western

Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...

, and

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 ...

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

Relationships

The family Pettalidae is

monophyletic In biological cladistics for the classification of organisms, monophyly is the condition of a taxonomic grouping being a clade – that is, a grouping of organisms which meets these criteria: # the grouping contains its own most recent co ...

, although it is at the moment (2007) unclear what the nearest relatives are. It probably originated in the southern part of Gondwana.

Parsimony analysis In phylogenetics and computational phylogenetics, maximum parsimony is an optimality criterion under which the phylogenetic tree that minimizes the total number of character-state changes (or minimizes the cost of differentially weighted charact ...

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, S ...

, though this could also be the case for the Stylocellidae, or it could be related to the

Sironidae The Sironidae are a family of harvestmen with more than 60 described species. The family shows a Laurasian distribution, with most species found in temperate Europe and the west coast of North America. The only exception is ''Suzukielus sauteri' ...

, or specifically to the sironid genus '' Suzukielus''. It is unrelated to the

Troglosironidae Troglosironidae is a family of harvestmen with seventeen described species in a single genus, ''Troglosiro'', which is found on the island of New Caledonia, in the Pacific Ocean. Name The name of the genus giving the family its name is a combina ...

that are

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 ...

New Caledonia New Caledonia ( ; ) is a group of islands in the southwest Pacific Ocean, southwest of Vanuatu and east of Australia. Located from Metropolitan France, it forms a Overseas France#Sui generis collectivity, ''sui generis'' collectivity of t ...

. The main lineages of the family may have arisen rapidly, possibly during the rapid expansion of ''

Glossopteris ''Glossopteris'' (etymology: from Ancient Greek γλῶσσα (glôssa, " tongue ") + πτερίς (pterís, " fern ")) is the largest and best-known genus of the extinct Permian order of seed plants known as Glossopteridales (also known as Ar ...

'' 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 (also known as the Antarctic Ocean), it contains the geographic South Pole. ...

, which formed a land bridge between Australia and South America up until circa 50

million years ago Million years ago, abbreviated as Mya, Myr (megayear) or Ma (megaannum), is a unit of time equal to (i.e. years), or approximately 31.6 teraseconds. Usage Myr is in common use in fields such as Earth science and cosmology. Myr is also used w ...

(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, found across southern South America (Chile, Argentina) and east and southeast Australia, New Zealand, New Guin ...

'' 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 (geology), epoch of the Jurassic Period, and it spans the geologic time scale, geologic time from 161.5 ± 1.0 to 143.1 ± 0.8 million years ago (Ma), which is preserved in Upper Jurassic stratum, strata.Owen ...

, 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 Thomas Bayes ( ; c. 1701 – 1761) was an English statistician, philosopher, and Presbyterian Presbyterianism is a historically Reformed Protestant tradition named for its form of church government by representative assemblies of elde ...

, but not in direct optimization parsimony analyses.

Species

The family Pettalidae contains the following genera and species: * '' Aoraki'' Boyer & Giribet, 2007 (New Zealand) :* '' Aoraki calcarobtusa'' ( Forster, 1952) :* '' Aoraki crypta'' (Forster, 1948) :* '' Aoraki denticulata'' (Forster, 1948) ::* '' Aoraki denticulata denticulata'' (Forster, 1948) ::* '' Aoraki denticulata major'' (Forster, 1952) :* '' Aoraki granulosa'' (Forster, 1952) :* '' Aoraki healyi'' (Forster, 1948) :* '' Aoraki inerma'' (Forster, 1948) :* '' Aoraki longitarsa'' (Forster, 1952) :* '' Aoraki stephenesis'' (Forster, 1952) :* '' Aoraki tumidata'' (Forster, 1948) :* '' Aoraki westlandica'' (Forster, 1952) * '' Archaeopurcellia'' Giribet, Shaw, Lord & Derkarabetian, 2022 :* '' Archaeopurcellia eureka'' Giribet, Shaw, Lord & Derkarabetian, 2022 * '' Austropurcellia'' Juberthie, 1988 :* '' 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'' (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 (

) :* '' Chileogovea jocasta'' Shear, 1993 :* '' Chileogovea oedipus'' Roewer, 1961 * '' Karripurcellia'' Giribet, 2003 (Australia:

Western Australia Western Australia (WA) is the westernmost state of Australia. It is bounded by the Indian Ocean to the north and west, the Southern Ocean to the south, the Northern Territory to the north-east, and South Australia to the south-east. Western Aust ...

) :* '' 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 :* '' Parapurcellia transvaalica'' (Lawrence, 1963) * '' 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 Sørensen () is a Danish- Norwegian patronymic surname meaning "son of Søren" (given name equivalent of Severin). , it is the eighth most common surname in Denmark. Immigrants to English-speaking countries often changed the spelling to ''Sorense ...

, 1904 :* '' Purcellia lawrencei'' de Bivort & Giribet, 2010 :* '' Purcellia leleupi'' Starega, 2008 * ''

Rakaia Rakaia is a town sited on 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 the townsh ...

'' Hirst, 1926 (New Zealand) :* '' Rakaia antipodiana'' Hirst, 1926 :* '' Rakaia australis'' Forster, 1952 :* '' Rakaia collaris'' Roewer, 1942 :* '' Rakaia digitata'' Forster, 1952 :* '' Rakaia dorothea'' Phillipps & Grimmett, 1932 :* '' Rakaia florensis'' Forster, 1948 :* '' Rakaia insula'' Forster, 1952 :* ''

Rakaia isolata Rakaia is a town sited on 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 the township ...

'' Forster, 1952 :* '' Rakaia lindsayi'' Forster, 1952 :* '' Rakaia macra'' Boyer & Giribet, 2003 :* '' Rakaia magna'' Forster, 1948 :* '' Rakaia media'' Forster, 1948 :* '' Rakaia minutissima'' Forster, 1948 :* '' Rakaia pauli'' Forster, 1952 :* '' Rakaia solitaria'' Forster, 1948 :* '' Rakaia sorenseni'' Forster, 1952 :* '' Rakaia stewartiensis'' Forster, 1948 :* '' Rakaia uniloca'' Forster, 1952

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