Acanthoclada, Bergquist, 1970

Morrow, Christine, Cárdenas, Paco, Boury-Esnault, Nicole, Picton, Bernard, Mccormack, Grace, Soest, Rob Van, Collins, Allen, Redmond, Niamh, Maggs, Christine, Sigwart, Julia & Allcock, Louise A., 2019, Integrating morphological and molecular taxonomy with the revised concept of Stelligeridae (Porifera: Demospongiae), Zoological Journal of the Linnean Society 187, pp. 31-81 : 34-36

Acanthoclada

Desmoxya Didiscus Halicnemia Heteroxya Higginsia Julavis Microxistyla Myrmekioderma Negombo Parahigginsia Halicnemia * Higginsia * Paratimea * Stelligera * Acanthoclada Alloscleria Desmoxya Heteroxya * Julavis Microxistyla Myrmekioderma Negombo Parahigginsia Didiscus *

Adreus *

Axos * Hemiasterella Acanthoclada Alloscleria Alveospngia Desmoxya Didiscus Heteroxya Julavis Microxistyla Myrmekioderma Negombo Parahigginsia Acanthoclada *

Halicnemia *

Higginsia *

Paratimea *

Stelligera *

Plenaster †

Heteroxyidae Alloscleria

( Axinellida )

Alveospongia

Heteroxya *

Julavis

Raspailiidae ( Axinellida ) Desmoxyidae ( Poecilosclerida ) Hemiasterellidae ( Tethyida ) Microxistyla Myrmekioderma pars.* Negombo Parahigginsia Myrmekioderma pars.* Didiscus *

Desmoxya *

Adreus *

Axos *

Hemiasterella

With the exception of Hooper & Van Soest (2002), these studies were not complete taxonomic reviews but rather faunas of a particular region, and therefore they include only the genera that were represented in their chosen area.

*There are supporting molecular data.

† Molecular data do not support the allocation of Plenaster to Stelligeridae (taxonomic affinities will be discussed in a separate manuscript).

Dendy (1905) established the subfamily Heteroxyinae for Heteroxya Topsent, 1898 and his new genus Acanthoxifer . Later, Dendy (1922) added Higginsia and Halicnemia to a section, Heteroxyeae, in the subfamily Axinellinae . Dendy (1922) agreed with Topsent (1897) and treated Bubaris constellata as a species of Halicnemia ; he stated: ‘Nor do I imagine that H. constellata , on account of its pseudasters, need be considered as the type of a distinct genus’. Topsent (1928) proposed the family Astraxinellidae Dendy, 1905 for the group Halicnemia , Higginsia , Vibulinus Gray, 1867 (= Stelligera ; synonymy by Hooper, 2002) and Hemiasterella and retained Heteroxyinae for Heteroxya and Acanthoxifer . He assigned four families to Halichondrina Vosmaer, 1887: Axinellidae ; Astraxinellidae ; Heteroxyidae Dendy, 1905 ; and Bubaridae Topsent, 1894 . Dendy (1905) introduced the name Astraxinellidae briefly and rather flippantly:

Some of the old group Axinellidae (e. g. Vibulinus , with asterose microscleres) must likewise be included in this sub-order, and it may prove necessary to institute a new family – Astraxinellidae – for their reception.

Given that there is no genus Astraxinella , the family name Astraxinellidae is invalid. For reasons of priority, Morrow et al. (2012) resurrected Stelligeridae for the clade that contained Halicnemia , Paratimea and Stelligera . Lendenfeld (1898) gave the following diagnosis for Stelligeridae :

Euastrosa with a spongin skeleton, which either just glues the megascleres together or is highly abundant like in Axinella . This family includes the two genera, Stelligera (with monactine megascleres and additional diactines) and Hemiasterella (with only diactine megascleres). In the Adriatic, they are represented by genus Stelligera .

Laubenfels (1936) subdivided Axinellidae into two subfamilies, Axinellinae and Higginsinae, for genera that are similar to typical Axinellidae but possess microsclere oxeas that are frequently spiny. Lévi (1955) synonymized Higginsinae with Desmoxyinae because Higginsia , the type genus of Laubenfels’ subfamily, was included in Desmoxyinae and Desmoxya , the type genus of Desmoxyinae, was included in Higginsinae by Laubenfels. Lévi (1955) also then raised Desmoxyinae to family rank. Lévi (1953) erected a new order, Axinellida , containing the family Axinellidae , which had previously been classified in Halichondrida ( Topsent, 1928; Laubenfels, 1936) and assigned Desmoxyidae Hallmann, 1917 , Heteroxyidae and Astraxinellidae (for axinellids with asterose microscleres) to it. Later, Lévi (1973) used Hemiasterellidae , rather than Astraxinellidae , for axinellids with asters ( Hemiasterella , Adreus , Paratimea and Stelligera ).

Bergquist (1970) and Wiedenmayer (1977) adopted the classification of Lévi (1955) and used Desmoxyidae for axinellids with microscleres in the form of spined or smooth microxeas. Bergquist (1970) established the genus Acanthoclada and assigned it to Desmoxyidae .

Van Soest et al. (1990), in a study using morphocladistics, called for the abandonment of Axinellida and the allocation of Desmoxyidae to Halichondrida and Hemiasterellidae to Hadromerida Topsent, 1894 (see Morrow et al., 2013). Van Soest et al. (1990) assigned Myrmekioderma Ehlers, 1870 and Didiscus Dendy, 1922 to Halichondriidae Gray, 1867 , a classification followed by Díaz et al. (1993). Later, Van Soest & Lehnert (1997) returned Myrmekioderma and Didiscus to Desmoxyidae .

The monotypic genus, Desmoxya Hallmann, 1917 , was created for Higginsia lunata Carter, 1885 . Hooper & Lévi (1993) synonymized Desmoxya and Dendropsis Ridley & Dendy, 1886 with Higginsia Higgin, 1877 based on the shared apomorphy of spined microxeas, but they highlighted that there were major skeletal differences and that Desmoxya might need to be resurrected to accommodate Higginsia- like species that lack any evidence of axial and extra-axial skeletons, such as Higginsia lunata and their new species, Higginsia anfractuosa . The discovery of another species, Desmoxya pelagiae Van Soest & Hooper, 2005 , from the cold-water coral reefs of the North Atlantic led to the resurrection of Desmoxya . Van Soest & Hooper (2005) considered Desmoxyidae synonymous with Heteroxyidae and changed the family to which Halicnemia , Higginsia , Desmoxya etc. were assigned from Desmoxyidae to Heteroxyidae .