Algospongia is a class of small, calcified fossil organisms of uncertain taxonomic position, assigned in a comprehensive 2010 review to "Animalia" incertae sedis (possibly Protista), but both prior to and post that to an unnamed phylum of Algae; other workers simply list them as Problematica (or Microproblematica). They occur in carbonate rocks of the Paleozoic era and their last representatives occur in the Late Permian geological period. Characteristic genera include Aoujgalia, Moravammina and the early-appearing Wetheredella, although the taxonomic validity (and algosponge affinity) of the last named genus has been disputed.
Background and possible taxonomic affinity
Algospongia (vernacular name: algosponges) is a taxon of calcified fossil organisms comprising around 90 accepted genera and several hundred species,[1] treated taxonomically as a single class in an unspecified phylum. Originally considered to be fossil sponges or "pseudo algae",[2] an assignment now refuted, their taxonomic position is somewhat unresolved. Of the three most prominent recent researchers, B. Mamet (Belgium) assigns certain families (e.g. Palaeoberesellaceae, Beresellaceae and Issinellaceae) to Chlorophyta (green algae) as an order of that phylum (Palaeosiphonocladales), at least one other family (Ungdarellaceae) to Rhodophyta (red algae), and some other genera (Groenlandella, Labyrinthoconus) simply to "Microproblematica".[3][4] Meanwhile, D. Vachard (France) and P. Cózar (Spain) treated Algospongia (such as the family Calcifoliaceae) as Algae incertae sedis, before deciding (in 2010) against any algal affinity and reassigning them to the [zoological] protists, as a group possibly paralleling the Foraminifera.[1] Since that paper, Cózar has continued to use zoological terminology in his published works, while Vachard has once again assigned the group to the Algae incertae sedis, lying hypothetically mid-way between the extant Rhodophyta and Chlorophyta.[5][6] A number of the genera allocated to Algospongia in the present treatment (which follows Vachard & Cózar, 2010 in the main) were recently (2021) treated as either Foraminifera, or as red or green algae, in the World Register of Marine Species, a situation that is currently under revision.[a] For convenience in the present article, the published summary of Vachard & Cózar (2010) is presented below (with modifications as necessary), which treats the relevant genera as a single class (not split e.g. among different botanical phyla) under zoological nomenclature, however also recognizing that the botanical treatment now preferred by Vachard may in fact be correct, an approach recently adopted in the Interim Register of Marine and Nonmarine Genera (IRMNG)[8] and reflected in the taxon box supplied with this article.
Members of the group occur in the fossil record from the Ordovician to the Late Permian periods, although only Wetheredella is prominent prior to the start of the Devonian.[1] They share a common wall appearance (frequently perforated), possess functional apertures between successive chambers or cells, and appear to have had a sessile or attached mode of life, at depths considered too deep for green algae but possibly overlapping those of red algae, while morphologically they do not exactly resemble members of either of those groups. The most comprehensive recent taxonomic treatment of the group is by Vachard & Cózar, 2010, in which they are treated as a single class divided into 2 orders (Aoujgalida and Moravamminida, corresponding to Aoujgaliales and Moravamminales in botanical nomenclature) typified by the genera Aoujgalia and Moravammina, respectively, plus a number of suborders; this treatment is reproduced below, together with adjustments based on more recent literature of relevance.
Morphology
Algosponges are described as a group of sessile or attached microorganisms with characteristic calcified walls described as "yellowish, apparently granular" that are frequently perforated, and possess either lateral or terminal apertures between successive chambers or cells. The order Aoujgalida (bot.: Aoujgaliales) is characterised by Vachard & Cózar as follows (emended description):
Attached, laminar to cylindrical or conical, bifurcated tests, composed of rows of chambers forming laminae, with a concentric or uniseriate growth. Encrusting or erect test. Irregular shape, generally subconical, occasionally cylindrical, ramified. Endoskeleton constituted by rows of chambers connected with a central or basal communication. Chambers quadratic to hemispherical, generally irregular in shape. The system of attachment is generally undifferentiated but can be preserved as "baskets" (e.g., Ungdarella). Wall calcitic, yellowish, hyaline and granular, generally compact or finely perforate (Pseudostacheoides, Costacheoides gen. nov.). Some interruptions of the chamber wall can exist (Ungdarella). In the [suborder] Calcifoliina, the interlaminar space is firstly thinner and then absent because the laminae evolve to petaloid forms, which finally contain filaments.
while the order Moravamminida (bot.: Moravamminales) is described as follows (emended description):
Tests generally tubular, sometimes bifurcated, rarely laminar, divided by foraminifer-like septa or pseudosepta. The system of attachment, generally unknown or constituted by a whorl of chambers around the substrate (e.g., Kettnerammina, Moravammina); by "bracelets" (Dil et al., 1977) (Exvotarisella, Ardengostella); or rarely, it is encrusting (Evlania). Wall calcareous, hyaline (granular to prismatic), generally perforated, with relatively common visible polysynthetic twinning of calcite, and exceptionally monocristalline extinction. Perforations are generally simple, aspondyl pores, in some taxa with up to third order of subdivisions. Pores vary from rare and sporadic to abundant and concentrated in specialized sectors. Generally functional and communicating with the exterior ([suborder] Moravamminina), they become blind ([suborder] Beresellina).[1]
Classification
The treatment below is based on that published by Vachard & Cózar in 2010, (alternative, botanical treatment indicated separated by "/"); more recent genera (post 2010) and, in some cases, corrections have been added per other available sources. If the group is considered zoological (i.e. non-Algal), nomenclature should follow the provisions of the International Code of Zoological Nomenclature (ICZN); if treated as Algae incertae sedis as per Vachard et al., 2016 and Vachard, 2021, the botanical Code (now International Code of Nomenclature for algae, fungi, and plants or ICNafp) applies.
Kingdom incertae sedis (Animalia or Protista, or Plantae, inferred from Vachard, 2021)
Phylum incertae sedis ("Algae incertae sedis" in Vachard, 2021)
Class †Algospongia Termier, Termier & Vachard, 1977
Order †Aoujgaliida Termier, Termier & Vachard, 1975 / bot.: †Aoujgaliales Termier et al., 1975 ex Vachard & Cózar, 2010
Suborder †Aoujgaliina G. Termier et al., 1975, nomen translat. Vachard & Cózar, 2010 / bot.: -inae?
†Valuzieria Termier, Termier & Vachard, 1977 (listed under accepted name †Aoujgalia in Vachard & Cózar, 2010, Table 3, but reinstated as a distinct genus by Vachard et al., 2016[5]
Family †Cuneiphycidae Vachard & Cózar, 2010 / bot.: †Cuneiphycaceae D. Vachard & P. Cózar, 2010 ('Cuneiphycidae')
†Ungdarella V.P. Maslov, 1956 (syn.: †Pseudoungdarella Ivanova, 1999; †Suundukella B. Chuvashov & A. Anfimov, 2007; †Ungdarelloides B. Chuvashov & A. Anfimov, 2007; †Urtasimella B. Chuvashov & A. Anfimov, 2007)
†Calcicaulis V.P. Shuysky & D.I. Schirschova in V.P. Shuysky, 1987
†Dreesenulella Vachard, 1991 (syn.: †Baculella Conil & Dreesen in Dreesen et al., 1985) note: synonymy reversed from that given in Vachard & Cózar, 2010, per Vachard et al., 2016[9]
†Palachemonella H. Beckmann, 1953 (misspelling: †Palaschemonella H. Beckmann, 1953)
†Pseudoissinella B.L. Mamet & B. Rudloff, 1972 (listed under accepted name Kettnerammina in Vachard & Cózar, 2010, Table 3, but only as "possible synonym" in text)
†Wetheredella Wood, 1948 (syn.: †Catena V.P. Maslov, 1956; †Cateniphycus V.P. Maslov in J.A. Orlov, 1963; Catena Maslov not listed in Vachard & Cózar, 2010)
Geological occurrence and biostratigraphic value
The stratigraphic range of selected algosponge genera is summarised in Figure 15 of Vachard & Cózar (2010). From this Figure it is apparent that Wetheredella is the sole representative of the group from the Upper Ordovician through the Silurian, being joined by Asphaltinoides a little before the Silurian ends. The majority of other genera then develop within, and/or are restricted to, the Devonian through Carboniferous periods, with a relatively small number (16) persisting through the Permian before all becoming extinct by the end of that period. The same authors also comment (their "Conclusion" no. 4) that "Biostratigraphically, the algospongia can become the most important group in the Tournaisian biostratigraphy with the conodonts, due to the rarity of foraminifera and true dasycladales at least in Europe (western Palaeo-Tethys) and Gondwana."[1]
Phylogeny
Vachard & Cózar (2010) present a suggested phylogeny for algosponge suborders and families in their Figure 17, which shows the Wetheredellina as the basal suborder giving rise to both the Moravamminina and the Donezellina, the Moravamminina then giving rise to the Beressellina, and Donezellina to the Aoujgaliina which in turn give rise to the Calcifoliina.
Alternative taxonomic opinions
As mentioned above, members of the claimed single class Algospongia have been assigned to different taxonomic groups over time, with no clear consensus emerging. More recently Wetheredella, treated as the earliest algosponge genus to appear in the fossil record by Vachard & Cózar, has been re-interpreted as the same as (and therefore junior synonym) of the incertae sedis taxon Allonema (and thus, presumably, not an algosponge) by Jarochowska & Munnecke (2014),[10] a proposal that has been admitted as "possible" by subsequent workers e.g. Liu et al., 2016.[11]
Notes
^As at 2021 (although the information cited may date from earlier compilation), the World Foraminifera Database (which forms part of the World Register of Marine Species) lists 18 of the present "algosponge" genera as Foraminifera (Alanyana, Aoujgalia, Baculella, Disonella, Evlania, Fourstonella, Kettnerammina, Litya, Moravammina, Palachemonella/Palaschemonella, Proninella, Saccorhina, Septammina, Stacheia, Stacheoides, Triplosphaerina, Vasicekia and Wetheredella), while the algal portion of WoRMS (imported from AlgaeBase) lists 14 algosponge genera as green algae (Anthracoporellopsis, Asphaltinella, Beresella, Crassikamaena, Cribrokamaena, Dvinella, Eomizzia, Issinella, Jansaella, Kamaena, Kamaenella, Parakamaena, Trinodella and Uraloporella) and 7 as red algae (Amorfia, Cuneiphycus, Donezella, Epistacheoides, Komia, Masloviporidium and Ungdarella), with Metakamaena assigned to "Protozoa", Asphaltina to "Biota incertae sedis", and the remaining genera not yet mentioned.[7] However it appears (March 2023) that at least some of these names have more recently been reallocated in AlgaeBase, information that is anticipated to be reflected in WoRMS in due course.
^Both the validity, and the taxonomic assignment, of Wetheredella, the type genus of this suborder and of the family Wetheredellidae, have been challenged by Jarochowska & Munnecke (2014) (for reference refer text), who contend that it is in fact a junior synonym of the genus Allonema Ulrich & Bassler, 1904, presently classified as [Animalia] incertae sedis. If this proposition is accepted by subsequent workers, the naming and validity of this suborder and its constituents, plus their present assignment to Algospongia, would require re-assessment.
References
^ abcdeVachard, D.; Cózar, P. (2010). "An attempt of classification of the Palaeozoic "incertae sedis" Algospongia". Revista Española de Micropaleontología. 42 (2): 129–241.
^Termier, H.; Termier, G.; Vachard, D. (1977). "On Moravamminida and Aoujgaliida (Porifera, Ischyrospongia): Upper Paleozoic "Pseudo Algae"". In Flügel, E. (ed.). Fossil Algae. Springer, Berlin, Heidelberg. pp. 215–219. ISBN9783642665189.
^Mamet, B.; Preat, A. (2013). "Essai de description d'algues nouvelles paléozoïques". Geologica Belgica. 16 (1–2): 35–48.
^Mamet, B. (1991). "Carboniferous calcareous algae". In Riding, R. (ed.). Calcareous Algae and Stromatolites. Springer-Verlag. pp. 370–451. ISBN9783642523373.
^Vachard, D. (2021). "Calcareous Algae (Rhodophyta and Chlorophyta)". In Elias, S.; Alderton, D. (eds.). Encyclopedia of Geology, Second Edition, Volume 3. Academic Press. pp. 389–406. ISBN9780081029091. In this work, the author states: "The Algospongia Termier et al. are subdivided into two orders: Moravamminales and Aougaliiales, six suborders (including Beresellina, Donezellina and Calcifoliina) and 17 families. However, some authors still consider that the Aougaliida are red algae, and Moravamminales green algae."
^"Algospongia". www.irmng.org. IRMNG. 2021. Retrieved 1 September 2021.
^Vachard, D.; Dreesen, R.; Marion, J.-M.; Mottequin, B. (2016). "New data on the incertae sedis biota and foraminifera of the mid-Famennian Baelen Member (Late Devonian, eastern Belgium)". Palaeobiodiversity and Palaeoenvironments. 97 (3): 565–584. doi:10.1007/s12549-016-0263-y. hdl:2268/130014. S2CID132979572.
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