The Haplogynae or haplogynes are one of the two main groups into which araneomorph spiders have traditionally been divided, the other being the Entelegynae. Morphological phylogenetic studies suggested that the Haplogynae formed a clade; more recent molecular phylogenetic studies refute this, although many of the ecribellate haplogynes do appear to form a clade, Synspermiata.
Unlike the Entelegynae, haplogynes lack hardened (sclerotized) female genitalia (epigynes).
Most of the species within this group have six eyes, as opposed to most other spiders. Spiders in the genus Tetrablemma (Tetrablemmidae) have only four eyes, as do some members of the family Caponiidae; caponiids may even have only two eyes. However, spiders in the family Plectreuridae have the normal eight eyes. [1]
The Haplogynae are one of the two major groups into which araneomorph spiders were traditionally divided, the other being the Entelegynae. In 2005, Coddington summarized the relationships of these groups as suggested by morphological phylogenetic studies: [2]
Araneomorphae |
| ||||||||||||||||||
Subsequent molecular phylogenetic studies have confirmed the monophyly of some of the groups suggested by morphological studies, while rejecting many others. [3] [4] [5]
A study published in 2015 suggested that two families formerly placed in the Haplogynae do not belong there. Filistatidae groups with Hypochilidae at the base of the Haplogynae; Leptonetidae is basal to the Entelegynae. [4] The similarity of some morphological features of Leptonetidae to those of entelegynes had already been noted. [6]
| Haplogynae sensu Coddington (2005) |
In 2016, a large molecular phylogenetic study was published online that included 932 spider species, representing all but one of the then known families. It "refutes important higher-level groups", [5] including Paleocribellatae, Neocribellatae, Araneoclada and Haplogynae. In the preferred cladogram, the "Haplogynae" are divided among a number of clades basal to the Entelegynae, forming at most a grade. "Haplogynae" in the sense of Coddington (2005) are shaded yellow in the cladogram below; Entelegynae in the same sense are shaded blue. The clade Synspermiata comprises all the ecribellate haplogynes and is consistently recovered, but with low support. [5]
Araneomorphae |
| ||||||||||||||||||||||||||||||||||||||||||
As shown above, Filistatidae and Leptonetidae are placed outside the traditional haplogynes in the analysis by Wheeler et al. (2017). Traditional haplogyne families they place in Synspermiata are: [5]
Telemidae, traditionally placed in Haplogynae, was not included in the analysis by Wheeler et al. [5] However, it is placed in Synspermiata in other studies. [7] The recently discovered haplogyne family Trogloraptoridae [8] was also placed in Synspermiata. [5]
Several extinct families have also been placed into the haplogynes.
Crevice weaver spiders (Filistatidae) comprise cribellate spiders with features that have been regarded as "primitive" for araneomorph spiders. They are weavers of funnel or tube webs. The family contains 18 genera and more than 120 described species worldwide.
Spider taxonomy is the part of taxonomy that is concerned with the science of naming, defining and classifying all spiders, members of the Araneae order of the arthropod class Arachnida, which has more than 48,500 described species. However, there are likely many species that have escaped the human eye as well as specimens stored in collections waiting to be described and classified. It is estimated that only one-third to one half of the total number of existing species have been described.
Cribellum literally means "little sieve", and in biology the term generally applies to anatomical structures in the form of tiny perforated plates.
The Dionycha are a clade of spiders (Araneomorphae:Entelegynae), characterized by the possession of two tarsal claws with tufts of hairs (setae) beside them, which produce strong adhesion, enabling some species to climb glass. The circumscription of the group has varied widely; a 2021 analysis resulted in about 20 families, including Salticidae, Gnaphosidae, and Clubionidae.
Lycosoidea is a clade or superfamily of araneomorph spiders. The traditional circumscription was based on a feature of the eyes. The tapetum is a reflective layer at the back of the eye, thought to increase sensitivity in low light levels. Lycosoids were then defined by having a "grate-shaped" tapetum. Research from the late 1990s onwards suggests that this feature has evolved more than once, possibly as many as five times, so that the original Lycosoidea is paraphyletic. Studies published in 2014 and 2015 suggest that a smaller group of families does form a clade.
The Palpimanoidea or palpimanoids, also known as assassin spiders, are a group of araneomorph spiders, originally treated as a superfamily. As with many such groups, its circumscription has varied. As of September 2018, the following five families were included:
Scytodoidea is a taxon of araneomorph spiders, at the rank of superfamily. It contains four families:
The Leptonetoidea are a superfamily of haplogyne araneomorph spiders with three families. Phylogenetic studies have provided weak support for the relationship among the families. The placement of one of the families within the Haplogynae has been questioned.
The Caponioidea or caponioids are a group of haplogyne araneomorph spiders that have been treated as superfamily with two members, the families Caponiidae and Tetrablemmidae. Phylogenetic studies from 1991 onwards have shown that the group is not monophyletic, being composed of two basal members of a larger clade. The precise members of that clade differ from study to study; one hypothesis is shown below.
The Dysderoidea are a clade or superfamily of araneomorph spiders. The monophyly of the group, initially consisting of the four families Dysderidae, Oonopidae, Orsolobidae and Segestriidae, has consistently been recovered in phylogenetic studies. In 2014, a new family, Trogloraptoridae, was created for a recently discovered species Trogloraptor marchingtoni. It was suggested that Trogloraptoridae may be the most basal member of the Dysderoidea clade. However, a later study found that Trogloraptoridae was placed outside the Dysderoidea and concluded that it was not part of this clade.
The Eresoidea or eresoids are a group of araneomorph spiders that have been treated as a superfamily. As usually circumscribed, the group contains three families: Eresidae, Hersiliidae and Oecobiidae. Studies and reviews based on morphology suggested the monophyly of the group; more recent gene-based studies have found the Eresidae and Oecobiidae to fall into different clades, placing doubt on the acceptability of the taxon. Some researchers have grouped Hersiliidae and Oecobiidae into the separate superfamily Oecobioidea, a conclusion supported in a 2017 study, which does not support Eresoidea.
The Deinopoidea or deinopoids are group of cribellate araneomorph spiders that may be treated as a superfamily. As usually circumscribed, the group contains two families: Deinopidae and Uloboridae.
The Agelenoidea or agelenoids are a superfamily or informal group of entelegyne araneomorph spiders. Phylogenetic studies since 2000 have not consistently recovered such a group, with more recent studies rejecting it.
The Dictynoidea or dictynoids are a group of araneomorph spiders that have been treated as a superfamily. The composition of the group has varied. Phylogenetic studies in the 21st century have failed to confirm the monophyly of the dictynoids as originally defined.
The Austrochiloidea or austrochiloids are a group of araneomorph spiders, treated as a superfamily. The taxon contains two families of eight-eyed spiders:
The Entelegynae or entelegynes are a subgroup of araneomorph spiders, the largest of the two main groups into which the araneomorphs were traditionally divided. Females have a genital plate (epigynum) and a "flow through" fertilization system; males have complex palpal bulbs. Molecular phylogenetic studies have supported the monophyly of Entelegynae.
Plectreuridae, also called plectreurid spiders, is a small spider family confined to the Southwestern United States, Mexico, and the Caribbean. Only two living genera are known—the nominate genus Plectreurys and Kibramoa. In the past, the family was more widespread, with the Jurassic genus Eoplectreurys known from China, the Eocene Palaeoplectreurys baltica from Baltic amber and the Miocene Plectreurys pittfieldi from Dominican amber.
Caponiidae is a family of ecribellate haplogyne spiders that are unusual in a number of ways. They differ from other spiders in lacking book lungs and having the posterior median spinnerets anteriorly displaced to form a transverse row with the anterior lateral spinnerets. Most species have only two eyes, which is also unusual among spiders. A few species of Caponiidae variously have four, six or eight eyes. In some species the number of eyes will increase when the spiderling changes its skin as it grows towards adulthood.
Orbiculariae is a potential clade of araneomorph spiders, uniting two groups that make orb webs. Phylogenetic analyses based on morphological characters have generally recovered this clade; analyses based on DNA have regularly concluded that the group is not monophyletic. The issue relates to the origin of orb webs: whether they evolved early in the evolutionary history of entelegyne spiders, with many groups subsequently losing the ability to make orb webs, or whether they evolved later, with fewer groups having lost this ability. As of September 2018, the weight of the evidence strongly favours the non-monophyly of "Orbiculariae" and hence the early evolution of orb webs, followed by multiple changes and losses.
Synspermiata is a clade of araneomorph spiders, comprising most of the former "haplogynes". They are united by having simpler genitalia than other araneomorph spiders, lacking a cribellum, and sharing an evolutionary history of synspermia – a particular way in which spermatozoa are grouped together when transferred to the female.