Rhamphorhynchus: Difference between revisions

{{For|the orchid genus which formerly had the same name|AspidogyneRhamphorhynchus (plant)}}

{{automaticAutomatic taxobox

| fossil_range = [[Late Jurassic]] ([[Tithonian]]), {{fossil range|150.8|148.5}}

| type_species = {{extinct}}'''''PterodactylusRhamphorhynchus longicaudusmuensteri'''''

*{{extinct}}'''''R. muensteri''''' <br /><small>([[Georg August Goldfuss|Goldfuss]], 1831)</small>

*{{extinct}}'''''R. longicaudus''''' <br /><small>(Meyer, 1846)</small>

*{{extinct}}'''''R. etchesi''''' <br /><small>O'Sullivan & Martill, 2015</small>

*{{extinct}}'''''R. jessoni'''''? <br /><small>Lydekker, 1890 (''[[nomen dubium]]'')</small>

*{{extinct}}'''''R. tendagurensis'''''? <br /><small>[[Hans Reck|Reck]], 1931 (''nomen dubium'')</small>

'''''Rhamphorhynchus''''' ({{IPAc-en|ˌ|r|æ|m|f|ə|ˈ|r|ɪ|ŋ|k|ə|s}},{{refn|{{OxfordDictionariesCite encyclopedia |url=http://www.lexico.com/definition/Rhamphorhynchus |accessarchive-url=https://web.archive.org/web/20200322182714/https://www.lexico.com/definition/rhamphorhynchus |url-status=dead |archive-date=20162020-0103-22 |title=Rhamphorhynchus |dictionary=[[Lexico]] UK English Dictionary |publisher=[[Oxford University Press]]}} }} from [[Ancient Greek]] ''rhamphos'' meaning "beak" and ''rhynchus'' meaning "snout") is a [[genus]] of long-tailed [[pterosaur]]s in the [[Jurassic]] period. Less specialized than contemporary, short-tailed [[pterodactyloid]] pterosaurs such as ''[[Pterodactylus]]'', it had a long tail, stiffened with [[ligament]]s, which ended in a characteristic soft-tissue tail vane. The mouth of ''Rhamphorhynchus'' housed needle-like teeth, which were angled forward, with a curved, sharp, beak-like tip lacking teeth, indicating a diet mainly of [[fish]]; indeed, fish and [[cephalopod]] remains are frequently found in ''Rhamphorhynchus'' abdominal contents, as well as in their coprolites[[coprolite]]s.<ref>{{cite journal | last1 = Frey | first1 = E. | last2 = Tischlinger | first2 = H. | year = 2012 | title = The Late Jurassic pterosaur Rhamphorhynchus, a frequent victim of the ganoid fish Aspidorhynchus? | journal = PLOS ONE | volume = 7 | issue = 3| page = e31945 | doi=10.1371/journal.pone.0031945 | pmid=22412850 | pmc=3296705| bibcode = 2012PLoSO...731945F | doi-access = free }}</ref>

Although fragmentary [[fossil]] remains possibly belonging to ''Rhamphorhynchus'' have been found in [[England]], [[Tanzania]], and [[Spain]], the best preserved specimens come from the [[Solnhofen limestone]] of [[Bavaria]], [[Germany]]. Many of these fossils preserve not only the bones but impressions of soft tissues, such as wing membranes. Scattered teeth believed to belong to ''Rhamphorhynchus'' have been found in [[Portugal]] as well.<ref name="cranfield-Rhamphorhynchus">''Rhamphorhynchus.'' In: Cranfield, Ingrid (ed.). ''The Illustrated Directory of Dinosaurs and Other Prehistoric Creatures''. London: Salamander Books, Ltd. Pp. 302-305.</ref>

The original species, ''Pterodactylus münsteri'', remained misclassified until a re-evaluation was published by [[Richard Owen]] in an 1861 book, in which he renamed it as ''Rhamphorhynchus münsteri''.<ref name="owen1861">Owen, R. (1861). ''Palaeontology, or a Systematic Summary of Extinct Animals and their Geological Relations''. Adam and Charles Black, Edinburgh, 1-463.</ref> The type specimen of ''R. muensteri'', described by Münster and Goldfuss, was lost during [[World War II]]. If available, a new specimen, or [[Type (biology)|neotype]], is designated as the type specimen if the original is lost or deemed too poorly preserved. [[Peter Wellnhofer]] declined to designate a neotype in his 1975 review of the genus, because a number of high-quality casts of the original specimen were still available in museum collections.<ref name="bennett1995">{{Cite journal | last1 = Bennett | first1 = S. C. | year = 1995 | title = A statistical study of ''Rhamphorhynchus'' from the Solnhofen Limestone of Germany: Year-classes of a single large species | journal = Journal of Paleontology | volume = 69 | issue = 3| pages = 569–580 | doi = 10.1017/S0022336000034946 | s2cid = 88244184 }}</ref> These can serve as [[Type (biology)|plastotypes]].

In 2015, a new species of ''Rhamphorhynchus, R. etchesi'' was named for associated remains of a left and right wing from the [[Kimmeridge Clay]] in the United Kingdom, the name commemorates the discoverer, [[Steve Etches]], a local collector of the fossils of the KimmerageKimmeridge Clay. It is distinguished from other species of ''Rhamphorhynchus'' by "the unique length ratio between wing phalanx 1 and wing phalanx 2"<ref>{{Cite journal|last1=O'Sullivan|first1=Michael|last2=Martill|first2=David M.|date=June 2015|title=Evidence for the presence of Rhamphorhynchus (Pterosauria: Rhamphorhynchinae) in the Kimmeridge Clay of the UK|journal=Proceedings of the Geologists' Association|language=en|volume=126|issue=3|pages=390–401|doi=10.1016/j.pgeola.2015.03.003|url=https://researchportal.port.ac.uk/portal/files/8204720/Evidence_for_the_presence_of_Rhamphorhynchus.pdf}}</ref>

The largest known specimen of ''Rhamphorhynchus muensteri'' (catalog number BMNH 37002) measures {{Convert|1.26|m|ft|sp=us}} long with a wingspan of {{Convert|1.81|m|ft|sp=us}}. A very large, fragmentary rhamphorhynchid specimen from Ettling in Germany may also belong to the genus, in which case ''Rhamphorhynchus'' would be the largest known non-pterodactyloid pterosaur and one of the largest pterosaurs known from the Jurassic. This specimen represents an individual around 180% the size of the next largest specimen of the genus, with an estimated wingspan of over 3 metres. <ref>{{cite journal |last1=Spindler |first1=Frederik |last2=Ifrim |first2=Christina |title=Die Spur einer Spur – ein möglicher erster Flugsaurier aus Ettling Trace of a trace – a putative first pterosaur from the Ettling locality |journal=Archaeopteryx|volume=37|pagepages= 75–83|date=2021 |url=https://www.researchgate.net/publication/357673254_Spindler_Ifrim_2021_Ettling_pterosaur_transitional_or_largest_rhamphorhynchid357673254}}</ref>

Contrary to a 1927 report by pterosaur researcher [[Ferdinand Broili]], ''Rhamphorhynchus'' lacked any bony or soft tissue crest, as seen in several species of contemporary small pterodactyloid pterosaurs. Broili claimed to have found a two-millimeter-tall crest made of thin bone that ran much of the skull's length in one ''Rhamphorhynchus'' specimen, evidenced by an impression in the surrounding rock and a few small fragments of the crest itself.<ref name="broili1927">{{Cite journal | last1 = Broili | first1 = F. | year = 1927 | title = Ein Exemplar von ''Rhamphorhynchus'' mit Resten von Schwimmhaut | journal = Sitzungs-Berichte der Bayerischen Akademie der Wissenschaften Mathematisch Naturwissenschaftlichen Abteilung | volume = 1927 | pages = 29–48 }}</ref> However, subsequent examination of this specimen by Wellnhofer in 1975 and Bennett in 2002 using both visible and ultraviolet light found no trace of a crest; both concluded that Broili was mistaken. The supposed crest, they concluded, was simply an artifact of preservation.<ref name="wellnhofer1975">{{cite journal | last1 = Wellnhofer | first1 = P. | year = 1975 | title = Die Rhamphorhynchoidea (Pterosauria) der Oberjura-Plattenkalke Süddeutschlands | journal = Palaeontographica A | volume = 148 | pages = 1–33 }}, '''148''': 132-186, '''149''': 1-30.</ref><ref name="bennett2002">{{Cite journal | doi = 10.1671/0272-4634(2002)022[0043:STPOTC]2.0.CO;2 | last1 = Bennett | first1 = S.C. | year = 2002 | title = Soft tissue preservation of the cranial crest of the pterosaur ''Germanodactylus'' from Solnhofen | journal = Journal of Vertebrate Paleontology | volume = 22 | issue = 1| pages = 43–48 | s2cid = 86308635 }}</ref> The teeth of ''Rhamphorhynchus'' intermesh when the jaw is closed and are suggestive of a [[Piscivore|piscivorous]] diet.<ref name="cranfield-Rhamphorhynchus" /> There are twenty teeth in the upper jaws and fourteen in the lower jaws.<ref name="cranfield-Rhamphorhynchus" />

The smallest known ''Rhamphorhynchus'' specimen has a wingspan of only {{convert|290 millimeters|mm|1|sp=us}}; however, it is likely that even such a small individual was capable of flight. Bennett examined two possibilities for hatchlings: that they were [[altricial]], requiring some period of parental care before leaving the nest, or that they were [[precocial]], hatching with sufficient size and ability for flight. If precocious, Bennett suggested that clutches would be small, with only one or two eggs laid per clutch, to compensate for the relatively large size of the hatchings. Bennett did not speculate on which possibility was more likely, though the discovery of a pterosaur embryo (''[[Avgodectes]]'') with strongly ossified bones suggests that pterosaurs in general were precocial, able to fly soon after hatching with minimal parental care.<ref name=wz05>{{cite journal |last=Wang |first=X. |author2=Zhou, Z. |year=2004 |title=Pterosaur embryo from the Early Cretaceous |journal=Nature |volume=429 |pages=623 |doi=10.1038/429621a |pmid=15190343 |issue=6992|s2cid=4428545 |doi-access=free }}</ref> This theory was contested by a [[Histology|histological]] study of ''Rhamphorhynchus'' that showed the initial rapid growth was followed by a prolonged period of slow growth.<ref>{{Cite journal | last1 = Prondvai | first1 = E. | last2 = Stein | first2 = K. | last3 = Ősi | first3 = A. | last4 = Sander | first4 = M. P. | editor1-last = Soares | editor1-first = Daphne | title = Life history of ''Rhamphorhynchus'' inferred from bone histology and the diversity of pterosaurian growth strategies | doi = 10.1371/journal.pone.0031392 | journal = PLOS ONE | volume = 7 | issue = 2 | pages = e31392 | year = 2012 | pmid = 22355361| pmc =3280310 | bibcode = 2012PLoSO...731392P | doi-access = free }}</ref>

In 2020, published ontogenetic analyses indicated that ''RhamphorynchusRhamphorhynchus'' could fly soon after hatching, supporting the theory of precociality in the species. It has also been suggested that juveniles may have occupied different sequential niches throughout their growth as they matured.<ref>{{cite journal|authorsauthor1=Hone, D.W.E.; |author2=Ratcliffe, J.M.; |author3=Riskin, D.K.; |author4=Hermanson, J.W.; |author5=Reisz, R.R. |year=2020|title=Unique near isometric ontogeny in the pterosaur ''Rhamphorhynchus'' suggests hatchlings could fly|journal=Lethaia|volume=54|pages=106–112|doi=10.1111/let.12391|doi-access=free}}</ref>

Though ''Rhamphorhynchus'' is often depicted as an aerial piscivore, recent evidence suggests that, much like most modern aquatic birds, it probably foraged while swimming. Like several [[pteranodontia]]ns it has hatchet-shaped deltopectoral crests, a short torso and short legs, all features associated with water based launching in pterosaurs. Its feet are broad and large, being useful for propulsion, and the predicted floating position is adequate by pterosaur standards.<ref>{{cite journal | last1 = Witton | first1 = M. P. | year = 2015 | title = Were early pterosaurs inept terrestrial locomotors? | journal = PeerJ | volume = 3 | page = e1018 | doi=10.7717/peerj.1018 | pmid=26157605 | pmc=4476129 | doi-access = free }}</ref> The animal's ability to swim may account for the genus' generally excellent fossil record, being in a position where preservation would be much easier.

Both [[Koh Ting-Pong]] and [[Peter Wellnhofer]] recognized two distinct groups among adult ''Rhamphorhynchus muensteri'', differentiated by the proportions of the neck, wing, and hind limbs, but particularly in the ratio of skull to [[humerus]] length. Both researchers noted that these two groups of specimens were found in roughly a 1:1 ratio, and interpreted them as different sexes.<ref name="wellnhofer1975"/><ref name="koh1937"/> Bennett tested for sexual dimorphism in ''Rhamphorhynchus'' by using a statistical analysis, and found that the specimens did indeed group together into small-headed and large-headed sets. However, without any known variation in the actual form of the bones or soft tissue (morphological differences), he found the case for sexual dimorphism inconclusive.<ref name="bennett1995"/> A 2024 study by Habib and Hone ''et al''., suggests that the high degree of tail variation in mature specimens may represent increased sexual selection in ''Rhamphorhynchus'', though it is equally likely a result of reduced flight constraint.<ref>{{cite journal | url=https://peerj.com/articles/17524/?fbclid=IwZXh0bgNhZW0CMTEAAR3rNFoZH8xTkPXTt8-_P9FszX_vrYYHsVCv7_BS4hkZ7rl0rgGTGlW_RyI_aem_dfglyiWFwgg9lLORZi3jiQ | doi=10.7717/peerj.17524 | doi-access=free | title=Intraspecific variation in the pterosaur ''Rhamphorhynchus muensteri—'' implications for flight and socio-sexual signaling | date=2024 | last1=Habib | first1=Michael B. | last2=Hone | first2=David WE. | journal=PeerJ | volume=12 | pages=e17524 | pmid=39035160 | pmc=11260407 }}</ref>

[[File:Rhamphorhynchus DBFlock.jpg|thumb|[[Paleoart|Life restoration]]reconstruction of a flock of ''R. muensteriRhamphorhynchus'' flying over the sea in Solnhofen area]]