CHLOROPHYCOPHYTA Papenfuss, 1955

Vachard, D, Krainer, K & Lucas, SG, 2015, Late Early Permian (late Leonardian; Kungurian) algae, microproblematica, and smaller foraminifers from the Yeso Group and San Andres Formation (New Mexico; USA), Palaeontologia Electronica (English ed.) 3 (8), pp. 1-77 : 10-12

Phylum CHLOROPHYCOPHYTA Papenfuss, 1955 Class BRYOPSIDOPHYCEAE Bessey, 1907 Order BRYOPSIDALES Schaffner, 1922

Description. See e.g., Chuvashov, Luchinina, Shuysky, Shaikin, Berchenko, Ishchenko, Saltovskaya and Shirshova, 1987.

Remarks. The bryopsidales (codiales or caulerpales auctorum) appeared in the Ordovician and were abundant and diverse in back-reef environments during the Late Silurian and Early to Middle Devonian ( Rothpletz, 1908; Poncet, 1982, 1990; Roux, 1985; Chuvashov et al., 1987; Mamet and Préat, 1994; Vachard, 1993). However, unquestionable bryopsidalean genera are very rare during Carboniferous and Permian times (e.g., Vachard et al., 1989b, 2001; Parvizi et al., 2013), whereas many Middle-Late Permian taxa belong to Gymnocodiaceae or “phylloid algae”, the botanical assignment of which is discussed. These questionable genera are Succodium Konishi, 1955 ; Aphroditicodium Elliott, 1970 ; Tauridium Güvenç, 1966 ; Tibeticodium Mu, 1982; Nanjinophycus Mu and Riding, 1983 ; and Thaiporella Endo, 1969 , which, although described as codiaceans, instead belong to gymnocodiaceans. Pia (1937), Elliott (1955) and Kochansky-Devidé and Sliskovic (1969) interpreted the gymnocodiaceans as galaxauracean red algae, based on the shape and subcortical location of the conceptacles, and the intraskeletal tubular elements identified as long files of cells. More recently, the gymnocodiaceans have been transferred to the green algae ( Mu and Riding, 1983; Chuvashov et al., 1987; Bucur, 1994; Radoicic, 2004; and Schlagintweit and Sanders, 2007). Morphologically, it is evident that the files of cells are rarely visible (except for Tauridium , perhaps) and that the deltoid terminations of many genera are similar to the utricles of the codiaceans.

Phylogenetically, the assignment to the galaxauraceans leads to many problems with a very problematical and puzzling history: 1) unquestionable pre-Middle Permian ancestors are unknown; the Devonian “ Gymnocodium ” devonicum Mamet et al., 1993, is most probably a fossil ulotrichale (according to the criteria proposed by Kozlowski and Kazmierczak, 1968); 2) productivity was huge in Middle-Late Permian times followed by an almost total absence during the Triassic and the Early and Middle Jurassic; 3) a Lazarus effect occurs from Late Jurassic to Palaeocene; 4) a second Lazarus effect occurs in the Recent. Several derivations from one or several stocks of bryopsidales are most easy to imagine.

We speculate here that the phylogeny codiaceans-gymnocodiaceans really exists, and that our late Early Permian material has been particularly interesting for confirming this relationship with possible transitional forms similar to and/or ancestral to Boueina ? described from our material.

Similarly, the “phylloid algae” are questionable codiacean algae. According to Schlagintweit (2010) and Granier (2012), the phylloid algae sensu lato can be considered as an informal botanical group. Indeed, the group defined by Pray and Wray (1963) is heterogenous, since it encompassed codiaceans (see Roux, 1985; Mamet et al., 1987; Forsythe et al., 2002; Torres, 2003), ancestral corallinales (see Wray, 1971; Vachard et al., 1989a; Corrochano et al., 2013), and algospongia (see Vachard et al., 1989a, 1989b, 2001, 2012; Vachard and Cózar 2010; Granier 2012; Parvizi et al., 2013). Furthermore, some representatives are probably the result of complex taphonomic interactions on algal thalli (Vachard et al., 1989a, 1989b, 2001). According to our published and unpublished data, the phylloid algae are to interpret as follows: 1) Eugonophyllum Konishi and Wray, 1961 ; Ivanovia Khvorova, 1946 ; Anchicodium Johnson, 1946 ; Neoanchicodium Endo in Endo and Kanuma, 1954; Kansaphyllum Baars, 1992 ; Calcipatera Torres, West and Sawin, 1992 ; and Iranicodium Senowbari-Daryan and Rashidi, 2010 most probably belong to the codiaceans (moreover, there are many generic synonymies; see discussion in Parvizi et al. (2013)); 2) Archaeolithophyllum Johnson, 1956 is typically a red alga; and perhaps, the unique unquestionable Palaeozoic calcareous red alga (Vachard et al., 1989a, 2001; Kirkland et al., 1991, 1993; Dawson, 1992; Moshier and Kirkland, 1993; Pintigore, 1994; Corrochano et al., 2013); 3) Calcifolium Maslov, 1956 and the calcifoliids differ totally and belong to the incertae sedis algosponges ( Vachard and Cózar, 2010).

Occurrence. Ordovician to Recent; cosmopolitan in subtropical seas.