Cyphophthalmi is a suborder 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 ...

, 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, Scopulophthalmi, and Sternophthalmi.

Description

Cyphophthalmi are smaller than the more familiar "daddy long-legs" harvestmen, with adults ranging from 1 to 7 mm in length. Moreover, their legs are relatively short compared to most other harvestmen, typically shorter than the body. Some superficially resemble

mite Mites are small arachnids (eight-legged arthropods) of two large orders, the Acariformes and the Parasitiformes, which were historically grouped together in the subclass Acari. However, most recent genetic analyses do not recover the two as eac ...

s, which is where they get their common name. Their coloration is almost always some shade of brown, with a heavily sclerotized body, and they are quite inconspicuous, residing 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 ...

or in

cave Caves or caverns are natural voids under the Earth's Planetary surface, surface. Caves often form by the weathering of rock and often extend deep underground. Exogene caves are smaller openings that extend a relatively short distance undergrou ...

s. Many Cyphophthalmi are eyeless, and presumably rely on olfactory cues to find food and mates. Very little is known about their behavior, though they likely subsist mostly by scavenging and preying on minute arthropods. They have low dispersal rates and consequently high

endemism 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 ...

Diagnostic features

Cyphophthalmi differ from other harvestmen in a number of key ways. Like all harvestmen, they have a pair of ozopores located on the sides of the prosoma. Unlike other harvestmen, however, the ozopores in Cyphophthalmi are located on elevated cone-shaped structures known as ozophores. Although members of most families are eyeless, most members of the families Pettalidae and Stylocellidae do have eyes located on or near the ozophores. These eyes, however, are not homologous to the eyes of other harvestmen, and are instead derived from ancestral lateral eyes, instead of ancestral median eyes, as is the case for other harvestmen. The male genitalia of Cyphophthalmi is unique among harvestmen, as males lack a true penis, and instead have a short ''spermatopositor'', a structure which is not inserted inside the female, but used to deposit a spermatophore. In addition, they are the only harvestmen to lack a genital operculum, and instead exhibit a completely open gonopore. Mating in Cyphophthalmi is not well studied, but in some species at least, males and females do engage in direct contact. They further differ from most harvestmen in that the first 8 opisthosomal tergites and the prosomal carapace are fused together in what is known as a ''scutum completum'', although this feature is also known from the unrelated family Sandokanidae.

Sexual dimorphisms

Like most animals, Cyphophthalmi express morphological distinctions between male and female individuals. Male Cyphophthalmi possess a structure on their fourth pair of tarsi known as an adenostyle. The adenostyle usually appears as a small hornlike projection, but can take a variety of shapes, depending on species. The function of the adenostyle is currently unknown, but is presumably associated with chemical glands. Aside from the adenostyle, males in many species possess glandular pores on either the underside of the opisthosoma or the anal region, although these glands are not present in all species. Males in several families also possess structural modifications of the anal plate and males of certain species in the family Stylocellidae possess a modified patch of cuticle near the adenostyle known as the Rambla's organ. The exact function of these glands and structures is currently unknown.

Chemical defenses

Like most harvestmen, Cyphophthalmi rely heavily on chemical secretions for defense. This defensive character is particularly well developed in Cyphophthalmi, when compared to the more familiar Eupnoi and Dyspnoi. Cyphophthalmi possess more elaborate musculature associated with the defensive glands than other harvestmen, and the secretions, which primarily consist of a variety of naphthoquinones and ketones, are thought to be more potent than in Eupnoi and Dyspnoi.

Distribution

Cyphophthalmids are found on all continents, with the exception 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. ...

, where they probably also once lived. As they did not disperse onto any oceanic islands, and it is believed that they did not travel between separate landmasses, they make an interesting subject for

biogeography Biogeography is the study of the species distribution, distribution of species and ecosystems in geography, geographic space and through evolutionary history of life, geological time. Organisms and biological community (ecology), communities o ...

. Each of the six currently recognized families has a distinct distribution: * Stylocellidae: found from

India India, officially the Republic of India, is a country in South Asia. It is the List of countries and dependencies by area, seventh-largest country by area; the List of countries by population (United Nations), most populous country since ...

New Guinea New Guinea (; Hiri Motu: ''Niu Gini''; , fossilized , also known as Papua or historically ) is the List of islands by area, world's second-largest island, with an area of . Located in Melanesia in the southwestern Pacific Ocean, the island is ...

* Ogoveidae: found exclusively in West Africa * Neogoveidae: found in Neotropical ecosystems between

Florida Florida ( ; ) is a U.S. state, state in the Southeastern United States, Southeastern region of the United States. It borders the Gulf of Mexico to the west, Alabama to the northwest, Georgia (U.S. state), Georgia to the north, the Atlantic ...

and

Ecuador Ecuador, officially the Republic of Ecuador, is a country in northwestern South America, bordered by Colombia on the north, Peru on the east and south, and the Pacific Ocean on the west. It also includes the Galápagos Province which contain ...

, and in western equatorial

Africa Africa is the world's second-largest and second-most populous continent after Asia. At about 30.3 million km2 (11.7 million square miles) including adjacent islands, it covers 20% of Earth's land area and 6% of its total surfac ...

* Pettalidae: found in

South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

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 ...

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, ...

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 ...

, demonstrating a

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 ...

n distribution. * Sironidae: found in temperate

Europe Europe is a continent located entirely in the Northern Hemisphere and mostly in the Eastern Hemisphere. It is bordered by the Arctic Ocean to the north, the Atlantic Ocean to the west, the Mediterranean Sea to the south, and Asia to the east ...

North America North America is a continent in the Northern Hemisphere, Northern and Western Hemisphere, Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South Ameri ...

, and

Japan Japan is an island country in East Asia. Located in the Pacific Ocean off the northeast coast of the Asia, Asian mainland, it is bordered on the west by the Sea of Japan and extends from the Sea of Okhotsk in the north to the East China Sea ...

, demonstrating a

Laurasia Laurasia () was the more northern of two large landmasses that formed part of the Pangaea supercontinent from around ( Mya), the other being Gondwana. It separated from Gondwana (beginning in the late Triassic period) during the breakup of Pa ...

n distribution * Troglosironidae: found exclusively in

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 ...

Colombia Colombia, officially the Republic of Colombia, is a country primarily located in South America with Insular region of Colombia, insular regions in North America. The Colombian mainland is bordered by the Caribbean Sea to the north, Venezuel ...

shows the largest diversity of cyphophthalmids among countries in South America. This could reflect the large number of

ecosystem An ecosystem (or ecological system) is a system formed by Organism, organisms in interaction with their Biophysical environment, environment. The Biotic material, biotic and abiotic components are linked together through nutrient cycles and en ...

s found there, but could also be due to a high rate of sampling in this country.

Fossil record

A fossil cyphophthalmid assigned to a modern genus, ''Siro platypedibus'' Dunlop & Giribet, 2003 (Sironidae), was described from

Paleogene The Paleogene Period ( ; also spelled Palaeogene or Palæogene) is a geologic period and system that spans 43 million years from the end of the Cretaceous Period Ma (million years ago) to the beginning of the Neogene Period Ma. It is the fir ...

aged Bitterfeld amber of eastern

Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...

. Another of this genus was described from

Eocene The Eocene ( ) is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (Ma). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene'' comes ...

Baltic amber as ''Siro balticus'' Dunlop & Mitov, 2011. A third fossil, ''Palaeosiro burmanicum'' Poinar, 2008, has been described from the

Cretaceous The Cretaceous ( ) is a geological period that lasted from about 143.1 to 66 mya (unit), million years ago (Mya). It is the third and final period of the Mesozoic Era (geology), Era, as well as the longest. At around 77.1 million years, it is the ...

(

Cenomanian The Cenomanian is, in the International Commission on Stratigraphy's (ICS) geological timescale, the oldest or earliest age (geology), age of the Late Cretaceous epoch (geology), Epoch or the lowest stage (stratigraphy), stage of the Upper Cretace ...

) aged

Burmese amber Burmese amber, also known as Burmite or Kachin amber, is amber from the Hukawng Valley in northern Myanmar. The amber is dated to around 100 million years ago, during the latest Albian to earliest Cenomanian ages of the mid-Cretaceous period. Th ...

. It was also assigned to Sironidae but is now thought to belong to Stylocellidae (a more typical Asian family). As the fossil record for Cyphophthalmi is so sparse, recent studies have attempted to predict the diversification time of the suborder by using molecular clocks and comparing gene sequences among living taxa. One 2012 study estimated the earliest diversification of the suborder at approximately 332 MYA, in the Carboniferous. Another study, in 2014, recovered a diversification age of approximately 340 MYA. A more recent study in 2017, however, recovered the diversification of the modern Cyphophthalmi lineages to have occurred more recently, during the Jurassic, with the emergence of the suborder as a whole at less than 300 MYA.

Relationships

Cyphophthalmi is one of the two major lineages of harvestmen; the other, containing the

Laniatores Laniatores is the largest suborder of the arachnid order Opiliones with over 4,200 described species worldwide. The majority of the species are highly dependent on humid environments and usually correlated with tropical and temperate forest habit ...

Dyspnoi Dyspnoi is a suborder of harvestmen, currently comprising 43 extant genera and 356 extant species, although more species are expected to be described in the future. The eight families are currently grouped into three superfamilies: the Acropsopil ...

and

Eupnoi The Eupnoi are a suborder of harvestmen, with more than 200 genera, and about 1,700 described species. They consist of two superfamilies, the Phalangioidea with many long-legged species common to northern temperate regions, and the small group ...

, is known as Phalangida.Shultz, Jeffrey W. & Pinto-da-Rocha, Ricardo (2007): Morphology and Functional Anatomy. In: Pinto-da-Rocha ''et al.'' 2007: 15ff The extinct suborder,

Tetrophthalmi Tetrophthalmi is an extinct suborder of Opiliones (commonly known as harvestmen or daddy-longlegs) that had both median and lateral eyes. First described in 2014, it is known from two extinct species. Phylogenetic analysis suggests that this e ...

, shares several features in common with Cyphophthalmi, and these two suborders may represent sister taxa. They are grouped into the following infraorders: Boreophthalmi, Scopulophthalmi, and Sternophthalmi. The Boreophthalmi and Sternophthalmi together form a monophyletic clade, and comprise 5 of the 6 families of Cyphphthalmi. The Scopulophthalmi, with only 1 family, the Pettalidae, forms the sister group to all the other families. Within the Sternophthalmi, the families Ogoveidae and Neogoveidae are most closely related, and form the superfamily Ogoveoidea, which is the sister group to the family Troglosironidae. The systematics of the Boreophthalmi, however, is not yet fully resolved. In one recent analysis, the Boreophthalmi were recovered as paraphyletic with respect to the Sternophthalmi. Even within the Boreophthalmi, the Sironidae has been recovered either as sister family to Stylocellidae, or as paraphyletic with respect to Stylocellidae. In addition, there are 3 genera of Cyphophthalmi with uncertain affinities that have not been placed in any family or infraorder.

Taxonomy

History of taxonomy

Originally, Cyphophthalmi was recognized as a single family consisting of two subfamilies, Sironini and Stylocellini. This classification scheme existed until 1980, when a new taxonomy was proposed, separating 5 families into the now-defunct infraorders Temperophthalmi (consisting of Petallidae, Sironidae and, later, Troglosironidae) and Tropicophthalmi (consisting of Stylocellidae, Ogoveidae and Neogoveidae). These two infraorders were never recovered as monophyletic in subsequent studies, and, following a 2012 phylogenetic analysis, a new Cyphophthalmi taxonomy was proposed.

Current taxonomy

The Cyphophthalmi are currently represented by the following taxa, which represent approximately 200 species, although there are a significant number of undescribed species that have been collected, and the current number of species is believed to be a severe under-representation. The monophyly of most subgroups of Cyphophthalmi is well supported, although both Boreophthalmi and Sironidae have been recovered as paraphyletic in recent analyses. * Cyphophthalmi ''incertae sedis'' *:* Family Parasironidae Karaman, Mitov & Snegovaya, 2024 (1 genus, 3 species) *::*'' Ankaratra franzi'' Shear & Gruber, 1996 *::*'' Marwe coarctata'' Shear, 1985 *::*'' Sheargovea mexasca'' Giribet, 2011 * Infraorder Boreophthalmi Giribet, 2012 ::* Family Sironidae Simon, 1879 (9 genera, 60 species) ::* Family Stylocellidae Hanson and Sørenson, 1904 (6 genera extinct 41 species) * Infraorder Scopulophthalmi Giribet, 2012 ::* Family Pettalidae Shear, 1980 (11 genera, 80 species) * Infraorder Sternophthalmi Giribet, 2012 ::* Family Troglosironidae Shear, 1993 (1 genus, 17 species) :* Superfamily Ogoveoidea Shear, 1980 :** Family Neogoveidae Shear, 1980 (11 genera, 40 species) :** Family Ogoveidae Shear, 1980 (1 genus, 3 species)