Ricinulei

Distribution

Of the three living genera, Ricinoides inhabits tropical West Africa from The Gambia and Senegal, eastwards to the Congo and Gabon, Pseudocellus inhabits North and Central America from Panama to Southern Texas, as well as the Caribbean (including Cuba), while Cryptocellus ranges from Rondônia in western Brazil northwards to Honduras in Central America.

Behavior and life history

Ricinuleids inhabit the leaf litter of rainforest floors, as well as caves, where they search for prey with their elongate sensory second leg pair. Ricinulei feed on other small invertebrates, although details of their natural prey are sparse. Relatively little is known about their courtship and mating habits, but males have been observed using their modified third pair of legs to transfer a spermatophore to the female. The eggs are carried under the mother's hood, until the young hatch into six-legged larva, which later molt into their eight-legged adult forms. The six-legged larva is a feature they share with Acari (see Relationships). Despite the scarce number of studies about the biology of this group, recent studies have reported nocturnal habits, as well as novel behaviors for this group, which include interactions between individuals different than mating. Ricinuleids are often found in large congregations, the exact purpose of which is unknown.

Fossil record

Ricinulei are unique among arachnids in that the first one to be discovered was a fossil, described in 1837 by the noted English geologist William Buckland; albeit misinterpreted as a beetle. Further fossil species were added in subsequent years by, among others, Samuel Hubbard Scudder, Reginald Innes Pocock and Alexander Petrunkevitch.

Fifteen of the twenty species of fossil ricinuleids discovered so far originate from the late Carboniferous (Pennsylvanian) coal measures of Europe and North America. They were revised in detail in 1992 by Paul Selden, who placed them in a separate suborder, Palaeoricinulei.

The fossils are divided into four families: Curculioididae, Poliocheridae, Primoricinuleidae, and Sigillaricinuleidae. The poliocherids are more like modern ricinuleids in having an opisthosoma with a series of three large, divided tergites. Curculioidids, by contrast, have an opisthosoma without obvious tergites, but with a single median sulcus; a dividing line running down the middle of the back. This superficially resembles the elytra of a beetle and explains why Buckland originally misidentified the first fossil species. Five species: ?Poliochera cretacea, Primoricinuleus pugio, Hirsutisoma acutiformis, H. bruckschi, H. grimaldii and H. dentata, are known from the Cenomanian (~99 million years old) Burmese amber of Myanmar; Curculioides bohemondi, the largest of all Ricinulei, was a member of the Curculioididae. Monooculricinuleus incisus and M. semiglobosus from Burmese amber were originally described as members of Ricinulei, but they might belong to Opiliones instead.

Some Carboniferous genera of Palaeoricinulei exceed modern Ricinulei in size, with bodies 24 millimetres (0.94 in) in length, and many appear to have had eyes, unlike modern representatives which are completely blind. It is likely they had a surface dwelling ecology, unlike that of modern Ricinulei. The fossil genera from the Cretaceous Burmese amber are referred to the extinct order Primoricinulei, and are thought to have had a different ecology than modern species as tree-dwelling predators that crawled on bark.

Genera

As of September 2022, the World Ricinulei Catalog accepts the following genera:

• Ricinoididae Ewing, 1929 (103+ species)
  • Cryptocellus Westwood, 1874
  • Pseudocellus Platnick, 1980
  • Ricinoides Ewing, 1929

In 2026, five new genera of African Ricinulei were described, including some species previously attributed to Ricinoides: Alantakun, Bambara, Gizoo, Tautau, and Ududo.

• † Curculioididae Cockerell, 1916 (12 species, Carboniferous)
  • † Amarixys Selden, 1992
  • † Curculioides Buckland, 1837
• † Hirsutisomidae Wunderlich, 2017 (4 species, Burmese amber)
  • † Hirsutisoma Wunderlich, 2017
• † Poliocheridae Scudder, 1884 (5 species, Carboniferous, ?Burmese amber)
  • † Poliochera Scudder, 1884
  • † Terpsicroton Selden, 1992
• † Primoricinuleidae Wunderlich, 2015 (1 species, Burmese amber)
  • † Primoricinuleus Wunderlich, 2015
• † Sigillaricinuleidae _{Wunderlich, 2022} (1 species, Burmese amber)
  • † Sigillaricinuleus Wunderlich, 2022

Relationships

Early work

In 1665, Robert Hooke described a large crab-like mite he observed with a microscope, he published a description of it in his book; Micrographia. The first living ricinuleid described using Linnaean taxonomy was from West Africa by Félix Édouard Guérin-Méneville in 1838, i.e. one year after the first fossil. This was followed by a second living example collected by Henry Walter Bates in Brazil and described by John Obadiah Westwood in 1874, and a third from Sierra Leone by Tamerlan Thorell in 1892. In these early studies ricinuleids were thought to be unusual harvestmen (Opiliones), and in his 1892 paper Thorell introduced the name "Ricinulei" for these animals as a suborder of the harvestman. Ricinuleids were subsequently recognized as an arachnid order in their own right in the 1904 monograph by Hansen & Soerensen. These authors recognised a group called "Arachnida micrura", comprising spiders, whip spiders, whip scorpions and ricinuleids, which they defined as having a rather narrow join between the prosoma and opisthosoma and a small 'tail end' to the opisthosoma.

Ricinuleids and mites

Morphological studies of arachnid relationships have largely concluded that ricinuleids are most closely related to Acari (mites and ticks) though more recent phylogenomic studies refute this. L. van der Hammen placed ricinuleids in a group called "Cryptognomae", together with the anactinotrichid mites only. Peter Weygoldt and Hannes Paulus referred to ricinuleids and all mites as "Acarinomorpha". Jeffrey Shultz used the name "Acaromorpha". This hypothesis recognizes that both ricinuleids and mites hatch with a larval stage with only six legs, rather than the usual eight seen in arachnids. The additional pair of legs appears later during development. Some authors have also suggested that the gnathosoma, a separate part of the body bearing the mouthparts, is also a unique character for ricinuleids and mites, but this feature is rather complex and difficult to interpret and other authors would restrict the presence of a gnathosoma sensu stricto to mites only.

Ricinuleids and trigonotarbids

In 1892, Ferdinand Karsch suggested that ricinuleids were the last living descendants of the extinct arachnid order Trigonotarbida. This hypothesis was widely overlooked, but was reintroduced by Jason Dunlop in 1996. Characteristics shared by ricinuleids and trigonotarbids include the division of the tergites on the opisthososma into median and lateral plates and the presence of an unusual 'locking mechanism' between the two halves of the body. A further study subsequently recognised that the tip of the pedipalp in both ricinuleids and trigonotarbids ends in a similar small claw. Ricinuleids as sister group of trigonotarbids was also recovered in the 2002 study by Gonzalo Giribet and colleagues.

Phylogenomic studies

Recent phylogenomic studies have recovered different relationships than those previously suggested. An analysis in early 2019 suggested the sister group of the ricinuleids may be Xiphosura, the arthropod order containing horseshoe crabs. In response to this work, a more recent study placed Ricinulei and Opiliones as sister taxa.