Algospongia


Algospongia is a class of small, calcified fossil organisms of uncertain taxonomic position, assigned in a comprehensive 2010 review to "Animalia" incertae sedis, but both prior to and post that to an unnamed phylum of Algae; other workers simply list them as Problematica, a term historically used for a number of other fossil zoological groups of uncertain placement. 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 of the last named genus has been disputed.

Background and possible taxonomic affinity

Algospongia is a taxon of calcified fossil organisms comprising around 90 accepted genera and several hundred species, treated taxonomically as a single class in an unspecified phylum. Originally considered to be fossil sponges or "pseudo algae", an assignment now refuted, their taxonomic position is somewhat unresolved. Of the three most prominent recent researchers, B. Mamet assigns certain families to Chlorophyta as an order of that phylum, at least one other family to Rhodophyta, and some other genera simply to "Microproblematica". Meanwhile, D. Vachard and P. Cózar treated Algospongia as Algae incertae sedis, before deciding against any algal affinity and reassigning them to the protists, as a group possibly paralleling the Foraminifera. 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. A number of the genera allocated to Algospongia in the present treatment were recently treated as either Foraminifera, or as red or green algae, in the World Register of Marine Species, a situation that is currently under revision. For convenience in the present article, the published summary of Vachard & Cózar is presented below, which treats the relevant genera as a single class under zoological nomenclature, however also recognizing that the botanical treatment now preferred by Vachard may in fact be correct, an approach currently adopted in the Interim Register of Marine and Nonmarine Genera.
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. They share a common wall appearance, 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 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 is characterised by Vachard & Cózar as follows :
while the order Moravamminida is described as follows :

Classification

The treatment below is based on that published by Vachard & Cózar in 2010, ; more recent genera and, in some cases, corrections have been added per other available sources. If the group is considered zoological, nomenclature should follow the provisions of the International Code of Zoological Nomenclature ; if treated as Algae incertae sedis as per Vachard et al., 2016 and Vachard, 2021, the botanical Code applies. The possibly analogous Foraminifera are members of the clade SAR, which is presently assigned to the kingdom Chromista in many classifications.
Kingdom incertae sedis
  • Phylum incertae sedis
  • * 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?
  • **** Family Aoujgaliidae Termier, Termier & Vachard, 1975 / bot.: Aoujgaliaceae Termier, Termier & Vachard, 1975
  • ***** Aoujgalia G. Termier & H. Termier, 1950
  • ***** Asteroaoujgalia Brenckle, 2004
  • ***** Costacheoides Vachard & Cózar, 2010
  • ***** Epistacheoides A.A. Petryk & B.L. Mamet, 1972
  • ***** Pseudostacheoides A.A. Petryk & B.L. Mamet, 1972
  • ***** Roquesselsia Termier, Termier & Vachard, 1977
  • ***** Sinustacheoides Termier, Termier & Vachard, 1977
  • ***** Stacheoides R.H. Cummings, 1955
  • ***** Valuzieria Termier, Termier & Vachard, 1977
  • ***** Cuneiphycus J.H. Johnson, 1960
  • ***** Iberiaella L.G. Racz, 1984
  • ***** Masloviporidium J.R. Groves & B.L. Mamet in D.F. Toomey & M.H. Nitecki, 1985
  • ***** Multiporidium Vachard & Cózar, 2010
  • **** Family Stacheiidae Loeblich & Tappan, 1961 / bot.: Stacheiaceae Loeblich & Tappan, 1961
  • ***** Fourstonella R.H. Cummings, 1955
  • ***** Stacheia Brady, 1876
  • **** Family Ungdarellidae Maslov, 1956 / bot.: Ungdarellaceae Maslov, 1954
  • ***** Cheggatella J. Poncet, 1989
  • ***** Cheggatellina B. Mamet & A. Preat, 2013
  • ***** Erevanella V.P. Maslov, 1962
  • ***** Foliophycus J.H. Johnson, 1960
  • ***** Komia K.B. Korde, 1951
  • ***** Petschoria K.B. Korde, 1951
  • ***** Ungdarella V.P. Maslov, 1956
  • ***** Ungdarellita Mamet & Villa, 2004
  • *** Suborder Calcifoliina G. Termier et al., 1977 nomen translat. Vachard & Cózar, 2010 / bot.: -inae?
  • **** Family Calcifoliidae Shuysky in Chuvashov et al., 1987 / bot.: Calcifoliaceae Termier, Termier & Vachard, 1977
  • ***** Calcifolium M.S. Svecov & L.M. Birina, 1935
  • ***** Falsocalcifolium D. Vachard & P. Cózar, 2005
  • ***** Frustulata V.D. Saltovskaja in M.R. Dzhalolov, 1984
  • **** Family Fasciellidae Shuysky, 1999 / bot.: Fasciellaceae Shuysky, 1999
  • ***** Fasciella R.M. Ivanova, 1973
  • ***** Fascifolium D. Vachard, A. Karim & P. Cózar in Vachard & Cózar, 2010
  • ** Order Moravammida Pokorný, 1951 / bot.: Moravamminales Pokorný, 1951 ex Vachard in Termier et al., 1975; syn.: Palaeosiphonocladales Shuysky, 1985
  • *** Suborder Beresellina Vachard, 1994, nomen. translat. Vachard & Cózar, 2010 / bot.: -inae?
  • **** Family Beresellidae Maslov & Kulik, 1956 / bot.: Beresellaceae Deloffre, 1988
  • ***** Ardengostella Vachard in Perret & Vachard, 1977
  • ***** Beresella V.N. Makhaev ex V.P. Maslov & E.L. Kulik, 1956
  • ***** Dvinella I.V. Khvorova, 1949
  • ***** Trinodella Vachard & Cózar, 2010
  • ***** Uraloporella K.B. Korde, 1950
  • *** Suborder Donezellina G. Termier et al., 1975, nomen translat. Vachard & Cózar, 2010 / bot.: -inae?
  • **** Family Claracrustidae Vachard in Vachard et al., 2001 / bot.: Claracrustaceae Vachard in Vachard et al., 2001
  • ***** Asphaltinella B. Mamet & A. Roux, 1978
  • ***** Claracrusta Vachard & Montenat, 1981
  • ***** Denisella Vachard & Cózar, 2010
  • ***** Kleinbergella B. Mamet & F. Boulvain, 1992
  • ***** Peristacheia B. Mamet & A. Roux, 1983
  • ***** Pokorninella Vachard in Perret & Vachard, 1977
  • ***** Precorninella Vachard, 1991
  • ***** Ungdarellina B. Mamet, 2002
  • **** Family Donezellidae Termier, Termier & Vachard, 1975 / bot.: Donezellaceae Termier, Termier & Vachard, 1975
  • ***** Alanyana T. Güvenç, 1967
  • ***** Donezella V.P. Maslov, 1929 ex Vachard in Meissami et al., 1978
  • ***** Kamaenella B.L. Mamet & A. Roux, 1975
  • ***** Praedonezella E.L. Kulik in O.L. Einor, 1973
  • *** Suborder Moravamminina Pokorný, 1951, nomen translat. Vachard & Cózar, 2010 / bot.: -inae?
  • **** Family Anthracoporellopsidae Shuysky, 1985 / bot.: Anthracoporellopsidacae Shuysky, 1985 ex Vachard in Vachard et al., 1989
  • ***** Anthracoporellopsis V.P. Maslov, 1956
  • ***** Brazhnikovia O.I. Berchenko, 1981
  • ***** Catenaella V.P. Shuysky, 1987
  • ***** Crassikamaena P.L. Brenckle, 1985
  • ***** Culmiella V.P. Shuysky in V.P. Shuysky & D.I. Schirschova in V.N. Dubatolov & T.A. Moskalenko, 1988
  • ***** Dokutchaevskella O.I. Berchenko, 1981
  • ***** Einoriella V.D. Saltovskaja in M.R. Dzhalolov, 1984
  • ***** Evlania Bykova, 1952
  • ***** Evlaniopsis Vachard & Montenat, 1981
  • ***** Groenlandella B.L. Mamet & L. Stemmerik, 2000
  • ***** Pseudonanopora B.L. Mamet & A. Roux, 1975
  • **** Family Issinellidae Deloffre, 1987 / bot.: Issinellaceae Deloffre, 1987
  • ***** Amarellina B. Mamet, 1995
  • ***** Calcicaulis V.P. Shuysky & D.I. Schirschova in V.P. Shuysky, 1987
  • ***** Dreesenulella Vachard, 1991 note: synonymy reversed from that given in Vachard & Cózar, 2010, per Vachard et al., 2016
  • ***** Eouraloporella O.I. Berchenko, 1981
  • ***** Issinella E.A. Reitlinger, 1954
  • ***** Issinellina V.P. Shuysky in V.P. Shuysky & D.I. Schirschova in V.N. Dubatolov & T.A. Moskalenko, 1988
  • ***** Jansaella B.L. Mamet & A. Roux, 1975
  • ***** Lemosquetella B. Mamet & A. Sebbar, 1998
  • ***** Luteotubulus Vachard in Vachard et al., 1977
  • ***** Serrisinella Vachard, 1991
  • ***** Tubus B.I. Tchuvashov, 1985
  • ***** Zidella V.D. Saltovskaja in M.R. Dzhalolov, 1984
  • **** Family Labyrinthoconidae Langer, 1979 / bot.: Labyrinthoconaceae Langer, 1979
  • ***** Labyrinthoconus W. Langer, 1979
  • ***** Proninella Reitlinger in Menner & Reitlinger, 1971
  • **** Family Moravamminidae Pokorný, 1951 / bot.: Moravamminaceae Pokorný, 1951
  • ***** Kettnerammina Pokorný, 1951
  • ***** Moravammina Pokorný, 1951
  • ***** Palachemonella H. Beckmann, 1953
  • ***** Pseudoissinella B.L. Mamet & B. Rudloff, 1972
  • ***** Triangulinella B. Mamet & A. Préat, 1985
  • ***** Vasicekia Pokorný, 1951
  • **** Family Palaeoberesellidae Mamet & Roux, 1974 / bot.: Palaeoberesellaceae Mamet & Roux, 1974
  • ***** Devonoscalae Langer, 1979
  • ***** Exvotarisella G.F. Elliott, 1970
  • ***** Kamaena I.A. Antropov in A.P. Jousé, 1967
  • ***** Kamaenina B. Mamet & A. Preat, 2013
  • ***** Kulikaella O.I. Berchenko, 1981
  • ***** Metakamaena R. Endô in Kobayashi & Toriyama, 1969
  • ***** Palaeoberesella B.L. Mamet & A. Roux, 1975
  • ***** Parakamaena B.L. Mamet & A. Roux, 1975
  • ***** Pseudokamaena B.L. Mamet in A.A. Petryk & B.L. Mamet, 1972
  • ***** Turgajella R.M. Ivanova in Bogush et al., 1990
  • ***** Wapitella B. Mamet & A. Preat, 2013
  • **** Family Uralitidae Vachard, 1991 / bot.: Uralitaceae D. Vachard, 1991
  • ***** Cribrokamaena P.L. Brenckle, 1985
  • ***** Uralites B.I. Tchuvashov, 1973
  • *** Suborder Wetheredellina Vachard in Dil et al., 1977, nomen translat. Vachard & Cózar, 2010 / bot.: -inae?
  • **** Family Asphaltinidae Vachard & Cózar, 2010 / bot.: Asphaltinaceae D. Vachard & P. Cózar, 2010
  • ***** Asphaltina B.L. Mamet in A.A. Petryk & B.L. Mamet, 1972
  • ***** Asphaltinoides J.A. Devera, 1987
  • ***** Permocatena D. Vachard & P. Miconnet, 1990
  • **** Family Wetheredellidae Vachard in Dil et al., 1976 / bot.: Wetheredellaceae Berchenko in Chuvashov et al., 1987
  • ***** Disonella Conil & Lys, 1964
  • ***** Sphaeroporella I.A. Antropov in A.P. Jousé, 1967
  • ***** Wetheredella Wood, 1948

Geological occurrence and biostratigraphic value

The stratigraphic range of selected algosponge genera is summarised in Figure 15 of Vachard & Cózar. 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 persisting through the Permian before all becoming extinct by the end of that period. The same authors also comment 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 and Gondwana."

Phylogeny

Vachard & Cózar 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 of the incertae sedis taxon Allonema by Jarochowska & Munnecke, a proposal that has been admitted as "possible" by subsequent workers e.g. Liu et al., 2016.