TUBIPHYTIDAE Vachard, Krainer and Lucas, 2012

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 : 22-24

Family TUBIPHYTIDAE Vachard, Krainer and Lucas, 2012

Description. Microproblematica probably constituted by the weakly to strongly transformed consortial association of cyanobacterial algae and Miliolata foraminifers. Proloculus and juvenarium are very poorly known but occasionally illustrated. Poorly transformed chambered tubes of Tubiphytes are called Cordiformis Güvenç¸ 1965 (nomen nudum; see Vachard and Montenat, 1981 and Vachard and Ferrière, 1991) and Nodophthalmidium Macfadyen, 1939 . Typical Tubiphytes appear as masses of a rounded, thick-walled taxon with a small rounded cavity (e.g., Razgallah and Vachard, 1991; Senowbari-Daryan and Flügel, 1993); atypical Tubiphytes exhibit large cavities, integrated small tubes and other transformations of the initial foraminifer for which the specimen illustrated by Riding and Guo (1992, plate 1, figure 4) is the best example.

Composition. Tubiphytes Maslov, 1956 (= Nigriporella Rigby, 1958 = Tubiella sensu Rauzer-Chernousova, 1949 (or 1951 fide Roux, 1985, p. 620) preoccupied = Shamovella in the sense of Rauzer-Chernousova, 1950 nomen dubium); Plexoramea Mello, 1977; Crescentiella Senowbari-Daryan et al., 2008 ; Latitubiphytes Vachard et al., 2012 ; Cordiformis Güvenç¸ 1965 (nomen nudum; = Incertae sedis 5 in the sense of Nguyen Duc Tien, 1979 = Foraminifer gen. et sp. indet. in the sense of Senowbari-Daryan and Flügel, 1993); one of the Vervilleina bradyi ( Spandel, 1901) illustrated by Filimonova (2010, plate 5, figure 11); Porferitubus Senowbari-Daryan, 1984; Nodophthalmidium Macfadyen, 1939 ; Ramovsia Kochansky-Devidé, 1973 (= Dorudia Jenny and Jenny-Deshusses, 1978 = Apterinella in the sense of Berczi-Makk and Kochansky-Devidé, 1981 non Cushman and Waters, 1928b);? Nubeculinella Cushman, 1930 .

Remarks. As indicated by many authors ( Elias, 1959; Riding and Guo, 1992), the genera Shamovella and Tubiphytes share the same stratigraphic level and type area, in the pre-Kungurian of the Urals in the Ishimbajevo oil-field, near the town of Sterlitamak, probably in the bioconstruction (shikhan) of Nizhni-Irginsk re-studied by Vennin et al. (1997). An origin in the Sylvinsk reefs, near the town of Kungur, where Tubiphytes is very common ( Chuvashov, 1983), is also possible. Furthermore, the type material of Tubiphytes was collected by Maslov and Rauzer-Chernousova together ( Riding, 1993). During this joint field-trip, Rauzer-Chernousova, who had created Shamovella a few years before had a valuable opportunity to promote it to Maslov. However, in his diagnosis of Tubiphytes, Maslov did not refer to Shamovella , but instead to Tubiella Rauzer-Chernousova as a preoccupied name which specifically must be replaced by Tubiphytes . After that, Rauzer-Chernousova has never contested Tubiphytes and proposed its replacement by Shamovella . We can deduce that, as soon as 1956, a consensus about Tubiphytes existed among the Russian micropalaeontologists, and, ipso facto, the unique argument of Riding (1993) and Riding and Barkham (1999) to replace Tubiphytes by Shamovella is irrelevant.

On the other hand, our studies in New Mexico confirm the synonymy of Nigriporella magna Rigby, 1958 , defined in the lower Leonard Formation, with Tubiphytes obscurus (synonymy first suggested by Elliott, 1962).

The external surface of the wall of Tubiphytes is probably relatively soft (or firm, at the maximum) because, as mentioned by Vachard (1980) and Vennin et al. (1997), Tubiphytes is rarely encrusted and only by Tubiphytes , Archaeolithoporella Endo, 1959 , tuberitinid microproblematica, and/or fistuliporid bryozoans. For this reason, the soft/firm wall of Tubiphytes was probably able to integrate some elements of the sea bottom: generally, sponge spicules but also some fragments of bryozoans, rare foraminifers and fragments of ungdarellacean algae ( Maslov, 1956; Homann, 1972; Vachard, 1980; Vachard and Montenat, 1981; Vennin et al., 1997). Due to this character, the foraminifer included in Tubiphytes and illustrated by Krainer et al. (2009, plate 3, figure 2), which is probably a Nodosinelloides netschajewi (compare with plate 5, figures 1, 2, 3, 4 in the same article), is not a boring foraminifer as suggested by Schlagintweit et al. (2013), but an included foraminifer. Another nodosariate incorporated in Tubiphytes has been illustrated by Razgallah and Vachard (1991, plate 2, figures 7, 8). After this soft/firm stage with the possibility of inclusion, Tubiphytes rapidly becomes strongly hard and lithified because many fragments of Tubiphytes can be reworked near the colonies in situ ( Sano et al., 1990), and because these fragments can be covered by early diagenetic fibrous cements (Vennin et al., 1997).

No new data can be provided here concerning the assignment of Tubiphytes . The interpretation of Bernier (1984), Vachard et al. (2001), and Vachard and Krainer (2001b) is fundamentally different from that of Maslov (1956) and Senowbari-Daryan and Flügel (1993), because the former authors consider the internal cavity as an important part of the organism; i.e., inherited from an ancestral Miliolata foraminifer. During evolution in the late Moscovian, this ancestor (probably a species of Palaeonubecularia Reitlinger, 1950 ) becomes intimely associated with a cyanobacterium (in a narrow assemblage simila to a lichen, for example). During this evolution, the shape of the foraminiferal cavity trends to disappear; for example, from Latitubiphytes in the Late Pennsylvanian (Kasimovian- Gzhelian) to true Tubiphytes in the Early Permian (Asselian-Kungurian), but it can reappear in the Late Permian with the “stages” of Tubiphytes obscurus illustrated by Senowbari-Daryan and Flügel, 1993 (text-figure 10 A-D, plate 3, figures 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16; plate 4, figures 1, 2, 5, 6) and Vachard et al., 2003 (plate 27, figures 1, 4, 5), and called Cordiformis (nomen nudum) by Güvenç (1965). Similarly, the resemblance of the cavity of Tubiphytes (now Crescentiella ) morronensis Crescenti, 1969 with that of nubecularioid foraminifers has often been stressed ( Bernier, 1984; Senowbari-Daryan et al., 2008).

Occurrence. Questionable references to tubiphytids in Bashkirian and early Moscovian deposits seem more related to “tolypamminid” (in fact, porcelaneous homeomorphs of these latter) foraminifers (Vachard et al., 2012). The family truly appears in the late or latest Moscovian ( Chuvashov et al., 1993; Vachard et al., 2012), during which Palaeonubecularia rustica Reitlinger, 1950 , true primitive tubiphytids ( Latitubiphytes ), and transitional forms between both genera can be found together. The typical tubiphytids are present from the Late Pennsylvanian (Kasimovian) to Early Cretaceous (for the Mesozoic forms see Senowbari-Daryan et al., 2008).