Algospongia

Last updated

Algospongia
Temporal range: Ordovician–Permian
Scientific classification Red Pencil Icon.png
Informal group: Algae
Phylum: incertae sedis
Class: Algospongia
G. Termier et al., 1977, orth. mut., emend Vachard & Cózar, 2010
Orders
Synonyms
  • Palaeosiphonocladales (for a subset of taxa)

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.

Contents

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. [lower-alpha 1] 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 promininent 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)

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

  1. 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.
  2. 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.

Related Research Articles

<span class="mw-page-title-main">Chlorophyta</span> Phylum of green algae

Chlorophyta or Prasinophyta is a taxon of green algae informally called chlorophytes. The name is used in two very different senses, so care is needed to determine the use by a particular author. In older classification systems, it refers to a highly paraphyletic group of all the green algae within the green plants (Viridiplantae) and thus includes about 7,000 species of mostly aquatic photosynthetic eukaryotic organisms. In newer classifications, it refers to the sister clade of the streptophytes/charophytes. The clade Streptophyta consists of the Charophyta in which the Embryophyta emerged. In this latter sense the Chlorophyta includes only about 4,300 species. About 90% of all known species live in freshwater. Like the land plants, green algae contain chlorophyll a and chlorophyll b and store food as starch in their plastids.

<span class="mw-page-title-main">Russulales</span> Order of fungi

The Russulales are an order of the Agaricomycetes,. According to the Dictionary of the Fungi, the order consists of 12 families, 80 genera, and 1767 species. According to Species Fungorum, the order contains 13 families, 117 genera, and 3,060 species.

<span class="mw-page-title-main">Golden algae</span> Class of algae

The Chrysophyceae, usually called chrysophytes, chrysomonads, golden-brown algae or golden algae are a large group of algae, found mostly in freshwater. Golden algae is also commonly used to refer to a single species, Prymnesium parvum, which causes fish kills.

<span class="mw-page-title-main">Cephalaspidea</span> Order of gastropods

The order Cephalaspidea, also known as the headshield slugs and bubble snails, is a major taxon of sea slugs and bubble snails, marine gastropod mollusks within the larger clade Euopisthobranchia. Bubble shells is another common name for these families of marine gastropods, some of which have thin bubble-like shells. This clade contains more than 600 species.

<i>Incertae sedis</i> Term to indicate an uncertain taxonomic position

Incertae sedis or problematica is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertainty at specific taxonomic levels is indicated by incertae familiae, incerti subordinis, incerti ordinis and similar terms.

<span class="mw-page-title-main">Eurotiomycetes</span> Class of fungi

Eurotiomycetes is a large class of ascomycetes with cleistothecial ascocarps within the subphylum Pezizomycotina, currently containing around 3810 species according to the Catalogue of Life. It is the third largest lichenized class, with more than 1200 lichen species that are mostly bitunicate in the formation of asci. It contains most of the fungi previously known morphologically as "Plectomycetes".

<span class="mw-page-title-main">Andropogoneae</span> Tribe of grasses

The Andropogoneae, sometimes called the sorghum tribe, are a large tribe of grasses (family Poaceae) with roughly 1,200 species in 90 genera, mainly distributed in tropical and subtropical areas. They include such important crops as maize (corn), sugarcane, and sorghum. All species in this tribe use C4 carbon fixation, which makes them competitive under warm, high-light conditions.

<span class="mw-page-title-main">Cirrina</span> Suborder of octopuses

Cirrina or Cirrata is a suborder and one of the two main divisions of octopuses. Cirrate octopuses have a small, internal shell and two fins on their head, while their sister suborder Incirrina has neither. The fins of cirrate octopods are associated with a unique cartilage-like shell in a shell sac. In cross-section, the fins have distinct proximal and distal regions, both of which are covered by a thin surface sheath of muscle.

<span class="mw-page-title-main">Ensifera</span> Suborder of cricket-like animals

Ensifera is a suborder of insects that includes the various types of crickets and their allies including: true crickets, camel crickets, bush crickets or katydids, grigs, weta and Cooloola monsters. This and the suborder Caelifera make up the order Orthoptera. Ensifera is believed to be a more ancient group than Caelifera, with its origins in the Carboniferous period, the split having occurred at the end of the Permian period. Unlike the Caelifera, the Ensifera contain numerous members that are partially carnivorous, feeding on other insects, as well as plants.

<span class="mw-page-title-main">Pygidicranidae</span> Family of earwigs

Pygidicranidae is a family of earwigs, formerly placed in the suborder Forficulina, now in the suborder Neodermaptera. The family currently contains twelve subfamilies and twenty six genera. Eight of the subfamilies are monotypic, each containing a single genus. Of the subfamilies, both Astreptolabidinae and Burmapygiinae are extinct and known solely from fossils found in Burmese amber. Similarly Archaeosoma, Gallinympha, and Geosoma, which have not been placed into any of the subfamilies, are also known only from fossils. Living members of the family are found in Australia, South Africa, North America, and Asia. The monotypic genus Anataelia, described by Ignacio Bolivar in 1899, is found only on the Canary Islands. As with all members of Neodermaptera, pygidicranids do not have any ocelli. The typical pygidicranid bodyplan includes a small, flattened-looking body, which has a dense covering of bristly hairs (setae). The pair of cerci at the end of the abdomen are symmetrical in structure. The head is broad, with the fourth, fifth and sixth antenna segments (antennomeres) that are not transverse. In general Pygidicranids also have equally sized ventral cervical sclerites, and in having the rearmost sclerite separated from, or only touching the center of the prosternum. Cannibalism of young has been observed in at least one species in the family, Challia hongkongensis, in which an adult female was found eating a still-living nymph of the same species. The same species in a different area has been observed possibly eating fruits or seeds, making the species an omnivore.

<span class="mw-page-title-main">Stylonurina</span> Extinct suborder of arthropods

Stylonurina is one of two suborders of eurypterids, a group of extinct arthropods commonly known as "sea scorpions". Members of the suborder are collectively and informally known as "stylonurine eurypterids" or "stylonurines". They are known from deposits primarily in Europe and North America, but also in Siberia.

<span class="mw-page-title-main">Merycoidodontoidea</span> Extinct superfamily of mammals

Merycoidodontoidea, sometimes called "oreodonts" or "ruminating hogs", is an extinct superfamily of prehistoric cud-chewing artiodactyls with short faces and fang-like canine teeth. As their name implies, some of the better known forms were generally hog-like, and the group has traditionally been placed within the Suina, though some recent work suggests they may have been more closely related to camels. "Oreodont" means "mountain teeth", referring to the appearance of the molars. Most oreodonts were sheep-sized, though some genera grew to the size of cattle. They were heavy-bodied, with short four-toed hooves and comparatively long tails.

<span class="mw-page-title-main">Incirrata</span> Suborder of octopuses

Incirrata is a suborder of the order Octopoda. The suborder contains the classic "benthic octopuses," as well as many pelagic octopus families, including the paper nautiluses. The incirrate octopuses are distinguished from the cirrate octopuses by the absence in the former of the "cirri" filaments for which the cirrates are named, as well as by the lack of paired swimming fins on the head, and lack of a small internal shell.

<span class="mw-page-title-main">Phasmatidae</span> Family of stick insects

The Phasmatidae are a family of the stick insects. They belong to the superfamily Anareolatae of suborder Verophasmatodea.

<span class="mw-page-title-main">Pleosporales</span> Order of fungi

The Pleosporales is the largest order in the fungal class Dothideomycetes. By a 2008 estimate it contains 23 families, 332 genera and more than 4700 species. The majority of species are saprobes on decaying plant material in fresh water, marine, or terrestrial environments, but several species are also associated with living plants as parasites, epiphytes or endophytes. The best studied species cause plant diseases on important agricultural crops e.g. Cochliobolus heterostrophus, causing southern corn leaf blight on maize, Phaeosphaeria nodorum causing glume blotch on wheat and Leptosphaeria maculans causing a stem canker on cabbage crops (Brassica). Some species of Pleosporales occur on animal dung and a small number occur as lichens and rock-inhabiting fungi.

The Rhodogorgonales are an order of red algae, a sister group to the corallines. They are always thalloid and calcified; their calcification is very different from the corallines, as individual calcite crystals are deposited in the cell wall of specialised cells; this suggests that the evolution of calcification may have been independent from the corallines. They have no fossil record.

<i>Solenopora</i> Extinct genus of algae

The extinct Solenoporaceae have traditionally been interpreted as a group of red algae ancestral to the Corallinales.

Wetheredella is a genus of calcimicrobes initially described from the Silurian of England, and subsequently reported from the Upper Ordovician to the end of the Carboniferous periods; its reefs are stated as being characteristic of the Ordovician-Silurian periods. Its taxonomic position is uncertain; it has been suggested to be a foraminiferan, a cyanobacterium or simply treated as a microproblematicum; Vachard & Cózar (2010) refer it to the Algospongia, a similarly controversial group that they assigned to the Protista but later, per Vachard, 2021, to Algae incertae sedis, in its own family and suborder (Wetheredellina) in the order Moravamminida. The genus is named in honor of the geologist Edward Wethered.

<span class="mw-page-title-main">Errantia</span> Subclass of annelid worms

Errantia is a diverse group of marine polychaete worms in the phylum Annelida. Traditionally a subclass of the paraphyletic class Polychaeta, it is currently regarded as a monophyletic group within the larger Pleistoannelida, composed of Errantia and Sedentaria. These worms are found worldwide in marine environments and brackish water.

<span class="mw-page-title-main">Ovalentaria</span> Clade of fishes

Ovalentaria is a clade of ray-finned fishes within the Percomorpha, referred to as a subseries. It is made up of a group of fish families which are referred to in Fishes of the World's fifth edition as incertae sedis, as well as the orders Mugiliformes, Cichliformes, and Blenniiformes. It was named by W. L. Smith and T. J. Near in Wainwright et al. (2012) based on a molecular phylogeny, but the authors suggested that the group was united by the presence of demersal eggs that are attached to a substrate. Some authors have used the ordinal name Stiassnyiformes for a clade including Mugiloidei, Plesiopidae, Blenniiformes, Atherinomorpha, and Cichlidae, and this grouping does appear to be monophyletic.

References

  1. 1 2 3 4 5 Vachard, D.; Cózar, P. (2010). "An attempt of classification of the Palaeozoic "incertae sedis" Algospongia". Revista Española de Micropaleontología. 42 (2): 129–241.
  2. 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. ISBN   9783642665189.
  3. Mamet, B.; Preat, A. (2013). "Essai de description d'algues nouvelles paléozoïques". Geologica Belgica. 16 (1–2): 35–48.
  4. Mamet, B. (1991). "Carboniferous calcareous algae". In Riding, R. (ed.). Calcareous Algae and Stromatolites. Springer-Verlag. pp. 370–451. ISBN   9783642523373.
  5. 1 2 Vachard, D.; Cózar, P.; Aretz, M.; Izart, A. (2016). "Late Viséan-early Serpukhovian cyanobacteria and algae from the Montagne Noire (France); taxonomy and biostratigraphy". Bulletin of Geosciences. 91 (3): 433–466. doi: 10.3140/bull.geosci.1613 .
  6. 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. ISBN   9780081029091. 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."
  7. Online query to WoRMS, 1 September 2021, using the "WoRMS Taxon Match" facility, https://www.marinespecies.org/aphia.php?p=match
  8. "Algospongia". www.irmng.org. IRMNG. 2021. Retrieved 1 September 2021.
  9. 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 . S2CID   132979572.
  10. Jarochowska, E.; Munnecke, A. (2014). "The Paleozoic problematica Wetheredella and Allonema are two aspects of the same organism". Facies. 60 (2): 651–662. doi:10.1007/s10347-014-0399-z. S2CID   129772080.
  11. Liu, L.; Wu, Y.; Yang, H.; Riding, R. (2016). "Ordovician calcified cyanobacteria and associated microfossils from the Tarim Basin, Northwest China: systematics and significance". Journal of Systematic Palaeontology. 14 (3): 183–210. doi:10.1080/14772019.2015.1030128. S2CID   129684955.