View source for Biology:Macroscaphitidae

{{Short description|Extinct family of ptychoceratoid cephalopods}}
{{Taxobox
| fossil_range = {{fossil range|130|93.5|latest=Cenomanian}}Lower Barremian – Lower Cenomanian
| image = Macroscaphites yvanii.jpg
| image_caption = Specimen of ''Macroscaphites yuani'' Meek 1876, from the Aptian (Early Cretaceous), Malenowit, Friedek, Schlesien.<ref>{{Cite journal|last=Gall|first=Larry|title=Invertebrate Paleontology Division, Yale Peabody Museum|year=2022 |publisher=Yale University Peabody Museum |url=https://www.gbif.org/dataset/854bc2fc-f762-11e1-a439-00145eb45e9a|doi=10.15468/nqheui}}</ref>
| regnum            = Animalia
| phylum            = [[Biology:Mollusca|Mollusca]]
| classis           = [[Biology:Cephalopoda|Cephalopoda]]
| subclassis        = †[[Biology:Ammonoidea|Ammonoidea]]
| ordo              = 
| superfamilia = †Ptychoceratoidea
| familia           = †'''Macroscaphitidae'''
| familia_authority = Hyatt, 1900
| type_genus = †''[[Biology:Macroscaphites|Macroscaphites]] (Macroscaphites)''
| type_genus_authority = Meek, 1876
| subdivision_ranks = Subgenera
| subdivision = 
  {{genus list
  |†''Macroscaphites (Macroscaphites)''|Meek, 1876
  |†''Macroscaphites (Costidiscus)''|Uhlig, 1883
  }}
}}

'''Macroscaphitidae'''<ref>{{Cite web |title=Macroscaphitidae |url=https://www.gbif.org/species/4621526 |access-date=2022-04-30 |publisher=GBIF |work=GBIF Backbone Taxonomy | type= Checklist dataset |doi= 10.15468/39omei}}</ref> ([[Social:Greek language|Greek]] μακρός, long and σκάφος, boat) is an extinct family of ptychoceratoid cephalopods from the subclass [[Biology:Ammonoidea|Ammonoidea]] that lived from the Lower Barremian to the Lower [[Earth:Cenomanian|Cenomanian]] stages of the [[Earth:Cretaceous|Cretaceous]]. Fossils of Macroscaphitidae were found all around the world although the abundance of found fossils is rather limited. Known fossils from collections were found largely in Europe (e.g. Italy, France, Czech Republic), South America (Mexico, Venezuela) and Africa (e.g. Mozambique).<ref>{{Cite web|title=''Macroscaphitidae'' in The Paleobiology Database|year=2022 |url=https://www.gbif.org/occurrence/search?dataset_key=bb5b30b4-827e-4d5e-a86a-825d65cb6583&taxon_key=4621526| work= Global Biodiversity Information Facility | type= database query result}}</ref> It is known for some species of which complete specimens were found that these animals developed a hetermorphic shell, i.e. the coiling of the shell was not regular, such that the first whirls formed a planispirally coiled evolute section as seen in homomorphic ammonites, but had an additional straight middle part and a presumably upwards facing aperture. Due to their odd morphology the taxonomic classification of Macroscaphitidae changed often over time since their discovery and may not be finally settled even now.

== Classification ==
The Macroscaphitidae comprise a family in the subclass of [[Biology:Ammonoidea|Ammonoidea]] in the class of [[Biology:Cephalopoda|Cephalopoda]] in the [[Biology:Phylum|phylum]] [[Biology:Mollusca|Mollusca]]. Due to their curious morphology multiple attempts to classify them into a superfamily were made. While the Macroscaphitidae were at first assigned to Hamitaceae in the 1950s,<ref>Wright, C. W. (1952). A classification of the Cretaceous ammonites. Journal of Paleontology, 213-222.</ref> this was revised in 2000 when they were classified as belonging to the superfamily [[Biology:Ancyloceratoidea|Ancyloceratoidea]].<ref>G. Landra, F. Cecca, and Z. Vasicek. 2000. Early Aptian ammonites from the top of the Maiolica and the anoxic "Selli level" (Lombardy, Southern Alps). Bollettino della Societa Paleontologica Italiana 39(1):29-45</ref> However, in 2007 it was proposed to assign Macroscaphitidae to the superfamily Ptychoceratiodea,<ref>J. Klein, R. Busnardo, M. Company, G. Delanoy, M. Kakbadze, S. Reboulet, P. Ropol, Z. Vasicek, and J. Vermeulen. 2007. Fossilium Catalogus. I: Animalia. Pars 144: Lower Cretaceous Ammonites III. Bochianitoidea, Protancyloceratoidea, Ancyloceratoidea, Ptychoceratoidea 1-381</ref> where Macroscaphitidae still belong today.
[[File:Phylogeny Macroscaphitidae.png|thumb|left|upright=1.8|Phylogeny of Ammonitida with emphasis on Macroscaphitidae (own figure based on a dataset by Raia et al. (2015)<ref>Raia, P., Passaro, F., Carotenuto, F., Maiorino, L., Piras, P., Teresi, L., ... & Fortelius, M. 2015. Cope’s rule and the universal scaling law of ornament complexity. The American Naturalist, 186(2), 165-175.</ref> and created with the R ''paleotree'' package<ref>Bapst, David W. (2012).  paleotree: an R package for paleontological and phylogenetic analyses of evolution.
  Methods Ecol. Evol. 3:803-807.</ref>)]]

=== Taxa ===
The family includes merely one genus (''[[Biology:Macroscaphites|Macroscaphites]]'' Meek 1876) with two subgenera, the nominative (or type) genus ''Macroscaphites (Macroscaphites)'' Meek 1876 and the other subgenus ''Macroscaphites (Costidscus)'' Uhlig 1883.<ref>{{Cite web | title=Macroscaphitidae |url=https://paleobiodb.org/classic/checkTaxonInfo?taxon_no=130400&is_real_user=1| access-date=2022-09-13|website=Paleobiology Database}}</ref> Some sources suggest more genera, such as ''Acantholytoceras'' Spath 1932, ''Ivaldiceras'' Delanoy 2010 or ''Rugacrioceras'' Vermeulen 2007, but the viability of such claims remains questionable.<ref>{{cite web |editor= Bagni Liggia |url=https://www.bagniliggia.it/WMSD/HtmFamily/MACROSCAPHITIDAER.htm |work=Worldwide Mollusc Species Database |title= Family: Macroscaphitidae |access-date=2022-02-10}}</ref> The type genus ''Macroscaphites (Macroscaphites)'' Meek 1876 contains four species, whereof ''Macroscaphites yuani'' Puzos 1831 constitutes the type species. The subgenus ''Macroscaphites (Costidiscus)'' Uhlig 1883 solely contains one species, ''Macroscaphites (Costidscus) reticostatus'' d’Orbigny 1840.

{{flowlist}}
Fm. †Macroscaphitidae Hyatt 1900
:G. †''Macroscaphites'' Meek 1876
::Subg. †''Macroscaphites (Costidiscus)'' Uhlig 1883
:::†''Macroscaphites (Costidiscus) recticostatus'' d'Orbigny 1840
::Subg. †''Macroscaphites (Macroscaphites)'' Meek 1876
:::†''Macroscaphites perforatus'' Avram 1984
:::†''Macroscaphites soaresi'' Da Silva 1962
:::†''Macroscaphites striatisulcatus'' d'Orbigny 1841
:::†''Macroscaphites yuani'' Puzos 1831
{{endflowlist}}

== Morphology ==
[[File:Costidiscus recticostatus (d'Orbigny, 1840).png|thumb|upright=1.4|''Costidiscus recticostatus'' (d'Orbigny, 1840)<ref>{{Cite journal|title=The fossil collection (F) of the Muséum national d'Histoire naturelle (MNHN – Paris)|year=2023 |url=https://www.gbif.org/dataset/b275a4c1-9859-4f3c-8ead-d86dde820fbc|doi=10.15468/fjjmkb |author1=Chagnoux S}}</ref>]]
Due to the scarce amount of fossils found of members of Macroscaphitidae, final remarks about their morphology are difficult to infer. Additionally, just few fossils have been preserved entirely. Nevertheless, a couple of specimens suggest that a wide range of sizes, up to tens of centimeters (e.g. ''Macroscaphites yuani'', see picture at the top). While the general anatomy of members of Macroscaphitidae is supposed to largely follow the one of homomorphic ammonites (i.e. aragonitic shell, supposedly complex suture lines etc.){{or | date= July 2023 | reason= previous source here, https://paleobiodb.org/classic/displayReference?reference_no=9159, says "unpublished; stated without evidence". citation template prior to script-assisted damage stated "personal information" }} much remains yet unknown, due to the small number of specimens. However, some complete specimens infer that the shells of ''Macroscaphites'' can be more or less divided into three parts: the planispirally coiled phragmocone (as in homomorphic ammonites), a straight part of the living chamber and a hook with an almnost 180 degree bend at the end where the aperture is located. Apart from that little is known about further morphological features.

== Life and ecology ==

It is likely that ''Macroscaphites'' lived similar to other, better known, heteromorphic ammonites. One assumption is that the whole living chamber was filled with the softbody to deduce the center of mass and buoyancy of the ammonite.<ref>Trueman A. E. 1941. The ammonite body-chamber, with special reference to the buoyancy and mode of life of the living ammonite. Quarterly Journal of the Geological Society of London, 384: 339-383.</ref> This commonly used model suggests for hetermorphic ammonites that the center of mass lies vertically below the center of buoyancy (to keep the animal stable in the water column) and thus predicts that the aperture of ''Macroscaphites'' was probably facing upwards.<ref>Kakabadzé M. V, and Sharikadzé M. Z. 1993. On the mode of life of heteromorph ammonites heterocone, ancyclocone, ptychocone), p. 209-215. In S. Elmi, C. Mangold and Y. Almeras (eds.), 3-eme Symposium international sur le Cephalopodes actuels et fossiles. Geobios Memoire Special 15.</ref> In addition to that it has been proposed for other heteromorphic ammonites that there was a second stable position with the aperture tilted toward the sea floor to prey on bottom dwellers. However, this remains controversial for Macroscaphitidae since e.g. specimens of ''Macroscaphites yuani'' suggest an aperture facing towards the phragmocone, a so-called ancyclocone, and thus preventing it from being tilted towards the sea floor.<ref>Monks, N., & Young, J. R. (1998). Body position and the functional morphology of Cretaceous heteromorph ammonites. Palaeontologia Electronica, 1(1), 15.</ref> Thus the exact feeding behavior of ''Macroscaphites'' remains unknown. While it is proposed that members of Macroscaphitidae were fast-moving nektonic carnivores due to there hydrodynamic anatomy, studies of other heteromorphic ammonites suggest a rather nektobenthonic lifestyle.<ref>Ebel K. 1992. Mode of life and soft body shape of heteromorph ammonites. Lethaia, 25: 179-193.</ref><ref>Klinger, H. C. 1980. Speculations on Buoyancy Control and Ecology in some Heteromorph Ammonites, p. 337-355. In House, M. R. and Senior, J. R. (eds.), The Ammonoidea. Systematics Association Special Volume 18. Academic Press, London, UK.</ref> Foraminifera, crustaceans and ostracods found in the guts of some species provide evidence that some heteromorphic ammonites fed on benthic animals,<ref>Lehmann U. & Weitschat W, 1973. Zur Anatomie und Okologie von Ammoniten: Funde von Kropf und Kiemen. Paläontologische Zeitschrift, 47: 69-76.</ref> making a epifaunal lifestyle rather likely. Furthermore, all heteromorphic ammonite lack the hyponomic sinus e.g. found in recent ''Nautilius'' that is used for manoeuvreable jetting, thus making a comparable lifestyle to ''Nautilus'' unlikely. In addition most fossil remnants of heteromorphic ammonites, including those of the Macroscaphitidae, were found in clay or marl rather than sandy facies suggesting a substrate dependend lifestyle.<ref>Marcinowski R. & Wiedmann J. 1990. The Albian ammonites of Poland. Palaeontologica Polonica, 50: 1-94.</ref> It was thus concluded that probably most heteromorphic ammonites, including Macroscaphitidae were bottom dwelling, like a small octopus with a mobile burrow.<ref>Monks, Neale and Young, Jeremy R. 1998. Body position and the functional morphology of Cretaceous Heteromorph ammonites, Palaeontologia Electronica Vol. 1, Issue 1; 1A; 15p. https://doi.org/10.26879/98001
palaeo-electronica.org/content/1-1-hetemorph-ammonites</ref>

== Fossils and distribution ==
[[File:Fossil distribution of Macroscaphitidae.png|thumb|left|Fossil distribution of Macroscaphitidae|upright=1.7]]
Compared to other ammonoid families just very few fossil specimens of Macroscaphitidae are known from collections. While some specimens are well preserved and provide information about the morphology and potentially about the way of living of ''Macroscaphites'', others are merely partly preserved making a comprehensive analysis of the morphology difficult. Fossils of members of Macroscaphitidae have been found in Europe (France,<ref>A. Kroh, A. Lukeneder, and G. Moosleitner. 2010. The invertebrate fauna from the Barremian of Serre de Bleyton (Drôme, SE France). Annalen des Naturhistorischen Museums in Wien, Serie A 112:569-574</ref> Italy,<ref>F. Cecca, G. Pallini, E. Erba, I. Permoli-Silva, and R. Coccioni. 1994. Hauterivian-Barremian chronostratigraphy based on ammonites, nannofossils, planktonic foraminifera and magnetic chrons from the Mediterranean domain. Cretaceous Research 15:457-467</ref> Serbia, Bulgaria,<ref>J. R. Manolov. 1962. New ammonites from the Barremian of north Bulgaria. Palaeontology 5:527-539</ref> the Czech Republic,<ref>J. Michalik and Z. Vasicek. 1989. Lower Cretaceous Stratigraphy and Paleogeography of the Czechoslovakian Western Carpathians. In J. Wiedmann (ed.), Cretaceous of the Western Tethys. Proceedings 3rd International Cretaceous Symposium, Tuebingen 1987:505-523</ref> Spain,<ref>M. Company, J. Sandoval, and J. M. Tavera. 1995. Lower Barremian ammonite biostratigraphy in the Subbetic Domain (Betic Cordillera, southern Spain). Cretaceous Research 16(2-3):243-256</ref> Georgia<ref>E. V. Kotetishvili. 1989. Biofacies Characteristics of Lower Cretaceous Deposits of Georgia. In J. Wiedmann (ed.), Cretaceous of the Western Tethys. Proceedings 3rd International Cretaceous Symposium, Tuebingen 1987:543-550</ref>), Africa (Egypt,<ref>H. Douvillé. 1916. Les terrains secondaires dans le massif du Moghara à l’Est de l’isthme de Suez. Mémoire de l’Académie des Sciences, Paris 54(2):1-184</ref> South Africa<ref name="one">W. J. Kennedy, H. C. Klinger, and M. V. Kakabadze. 2009. Macroscaphites Meek, 1876, a heteromorph ammonite from the Lower Aptian of southern Mozambique and northern KwaZulu-Natal. African Natural History 5:37-41</ref> and Mozambique<ref name="one" />) and South America (Mexico<ref>R. Barragán, J. A. Moreno-Bedmar, F. Núñez-Useche, L. F. Álvarez-Sánchez, and G. Delanoy. 2021. Ammonite biostratigraphy of two stratigraphic sections of the La Peña Formation (Aptian, Lower Cretaceous) in Nuevo León State, northeast Mexico. Cretaceous Research 125(104862)</ref> and Venezuela<ref>O. Renz. 1982. The Cretaceous ammonites of Venezuela
 1-132</ref>). The fossils have been found in the form of [[Biology:Fossil#Casts and molds|casts, molds]], [[Biology:Fossil#Adpression (compression-impression)|impressions]] and [[Biology:Fossil#Replacement and recrystallization|replacement]]<ref>E. Böse. 1923. Algunas faunas cretácias de Zacatecas, Durango y Guerrero. Boletín del Instituto Géologica de México 42:1-219</ref><ref>Z. Vasicek, D. Rabrenovic, V. J. Radulovic, B. V. Radulovic, and I. Mojsic. 2013. Ammonoids (Desmoceratoidea and Silesitoidea) from the Late Barremian of Boljetin, eastern Serbia. Cretaceous Research 41:39-54</ref><ref>Z. Vasicek, D. Rabrenovic, P. Skupien, V. J. Radulovic, B. V. Radulovic and I. Mojsic. 2014. Ammonites (Phylloceratina, Lytoceratina and Ancyloceratina) and organic-walled dinoflagellate cysts from the Late Barremian in Boljetin, eastern Serbia. Cretaceous Research 47:140-159</ref> and occurred mainly in limestone<ref>G. Delanoy, J. A. Moreno-Bedmar, J.J. Ruiz and D. Tolós Lládser. 2013. Xerticeras gen. nov., a new genus of micromorphic heteromorph ammonite (Ancyloceratina, Ancyloceratidae)from the lower Aptian of Spain. Carnets de Géologie [Notebooks on Geology] CG2013_A02:89-103</ref>

== See also ==
* [[Biology:Ammonoidea|Ammonoidea]]
* [[Biology:Ancyloceratoidea|Ancyloceratoidea]]

== References ==
{{reflist|2}}

{{Taxonbar|from=Q23758312}}

[[Category:Cephalopods]]
[[Category:Prehistoric cephalopod families]]

{{Sourceattribution|Macroscaphitidae}}