Editing Onychophora (section)

=== Velvet worms and their diversification ===
[[File:Helenodora.jpg|thumb|Reconstruction of the Carboniferous possible onychophoran ''[[Helenodora]]''|350x350px]]

The earliest potential onychophoran, ''[[Helenodora]]'', originates from the [[Middle Pennsylvanian|Mid Pennsylvanian]] [[Mazon Creek fossil beds|Mazon Creek lagerstätte]] in the [[United States]]. Originally described in 1980,<ref>{{Cite journal |last=Thompson |first=Ida |last2=Jones |first2=Douglas S. |date=1980 |title=A Possible Onychophoran from the Middle Pennsylvanian Mazon Creek Beds of Northern Illinois |url=https://www.jstor.org/stable/1304204 |journal=Journal of Paleontology |volume=54 |issue=3 |pages=588–596 |issn=0022-3360}}</ref> Helenodora was long regarded as the earliest known onychophoran.<ref name="Jr2000">{{cite journal |last=Poinar |first=George |date=Winter 2000 |title=Fossil onychophorans from Dominican and Baltic amber: ''Tertiapatus dominicanus'' n.g., n.sp. (Tertiapatidae n. fam.) and ''Succinipatopsis balticus'' n.g., n.sp. (Succinipatopsidae n. fam.) with a Proposed Classification of the Subphylum Onychophora |journal=Invertebrate Biology |volume=119 |issue=1 |pages=104–109 |bibcode=2000InvBi.119..104P |doi=10.1111/j.1744-7410.2000.tb00178.x |jstor=3227105 |doi-access=free}}</ref>'''<ref name="G02" />'''<ref name="Smith-OrtegaHz-2014" /><ref name=":8" /> This changed in 2016 when a paper by Murdock et al. doubted its affinity. Based on the absence of many [[Decomposition|decay]]-resistant features ([[Claw|claws]], jaws, slime papillae) and unclear [[ecology]] (''[[Carbotubulus]]'', a late-surviving [[Aquatic animal|aquatic]] [[Lobopodia|lobopodian]], was found in the same [[Geological formation|formation]]), the authors of this study placed ''Helenodora'' as a basal lobopodian most closely related to ''[[Paucipodia]]''.<ref name=":07">{{cite journal |last1=Murdock |first1=D. J. E. |last2=Gabbott |first2=S. E. |last3=Purnell |first3=M. A. |date=2016 |title=The impact of taphonomic data on phylogenetic resolution: ''Helenodora inopinata'' (Carboniferous, Mazon Creek Lagerstätte) and the onychophoran stem lineage |journal=BMC Evolutionary Biology |volume=16 |issue=19 |pages=19 |bibcode=2016BMCEE..16...19M |doi=10.1186/s12862-016-0582-7 |pmc=4722706 |pmid=26801389 |doi-access=free}}</ref> 

The earliest definite onychophoran is the [[France|French]] ''[[Antennipatus]]'' from a [[Stephanian (stage)|Stephanian]] ([[Late Pennsylvanian]]<ref name=":06">{{Cite journal |last=Knight |first=John A. |last2=Cleal |first2=Christopher J. |last3=Álvarez-Vázquez |first3=Carmen |date=2023-06-14 |title=The challenge of relating the Kasimovian to west European chronostratigraphy: a critical review of the Cantabrian and Barruelian substages of the Stephanian Stage |url=https://www.lyellcollection.org/doi/10.1144/SP535-2022-189 |journal=Geological Society, London, Special Publications |language=en |volume=535 |issue=1 |pages=31–71 |doi=10.1144/SP535-2022-189 |issn=0305-8719 |doi-access=free}}</ref>) [[Lagerstätte]] in [[Montceau-les-Mines]].<ref name="GarwoodEdgecombe2016" /> This animal had a [[ventral]] (bottom facing) [[mouth]], rings of [[Dermal papillae (Onychophora)|dermal papillae]] on its trunk and [[Appendage|limbs]], and was at least somewhat [[Terrestrial animal|terrestrial]] because of its slime papillae.<ref name="GarwoodEdgecombe2016" /> However, due to the way it was preserved, it is unknown if ''Antennipatus'' belonged to the [[Stem-group|stem]] or [[crown group]] of the two living [[Family (biology)|families]] of velvet worm ([[Peripatidae]] and [[Peripatopsidae]]).<ref name="GarwoodEdgecombe2016">{{cite journal |last1=Garwood |first1=Russell J. |last2=Edgecombe |first2=Gregory D. |last3=Charbonnier |first3=Sylvain |last4=Chabard |first4=Dominique |last5=Sotty |first5=Daniel |last6=Giribet |first6=Gonzalo |year=2016 |title=Carboniferous Onychophora from Montceau-les-Mines, France, and onychophoran terrestrialization |journal=Invertebrate Biology |volume=135 |issue=3 |pages=179–190 |issn=1077-8306 |pmid=27708504 |pmc=5042098 |doi=10.1111/ivb.12130 |doi-access=free }}</ref><ref name=":033">{{Cite journal |last=Giribet |first=Gonzalo |last2=Buckman-Young |first2=Rebecca S. |last3=Costa |first3=Cristiano Sampaio |last4=Baker |first4=Caitlin M. |last5=Benavides |first5=Ligia R. |last6=Branstetter |first6=Michael G. |last7=Daniels |first7=Savel R. |last8=Pinto-da-Rocha |first8=Ricardo |date=2018 |title=The ‘Peripatos’ in Eurogondwana? – Lack of evidence that south-east Asian onychophorans walked through Europe |url=https://bioone.org/journals/Invertebrate-Systematics/volume-32/issue-4/IS18007/The-iPeripatos-i-in-Eurogondwana--Lack-of-evidence-that/10.1071/IS18007.full |journal=Invertebrate Systematics |volume=32 |issue=4 |pages=840–863 |doi=10.1071/IS18007}}</ref><ref name=":7">{{Cite journal |last=Baker |first=Caitlin M |last2=Buckman-Young |first2=Rebecca S |last3=Costa |first3=Cristiano S |last4=Giribet |first4=Gonzalo |date=2021-12-01 |title=Phylogenomic Analysis of Velvet Worms (Onychophora) Uncovers an Evolutionary Radiation in the Neotropics |url=https://academic.oup.com/mbe/article/38/12/5391/6357048?login=true |journal=Molecular Biology and Evolution |volume=38 |issue=12 |pages=5391–5404 |doi=10.1093/molbev/msab251 |issn=1537-1719|pmc=8662635 }}</ref>

Based on [[molecular dating]], crown-group onychophorans diverged around 376 million years ago in the [[Late Devonian]]. This estimation happens regardless if ''[[Antennipatus]]'' is used to constrain the divergence date, or if no calibration is done at all.<ref name=":7" /> The earliest known [[Crown group|crown-group]] onychophoran, ''[[Cretoperipatus]]'', is known from multiple specimens in [[Burmese amber]].'''<ref name="G02">{{cite journal |last1=Grimaldi |first1=David A. |last2=Engel |first2=Michael S. |last3=Nascimbene |first3=Paul C. |date=March 2002 |title=Fossiliferous Cretaceous Amber from Myanmar (Burma): Its Rediscovery, Biotic Diversity, and Paleontological Significance |url=https://www.biodiversitylibrary.org/bibliography/178519 |journal=American Museum Novitates |issue=3361 |pages=1–71 |doi=10.1206/0003-0082(2002)361<0001:FCAFMB>2.0.CO;2 |s2cid=53645124 |hdl=2246/2914}}</ref><ref name="Oliveira">{{cite journal |last1=Oliveira |first1=I. S. |last2=Bai |first2=M |last3=Jahn |first3=H |last4=Gross |first4=V |last5=Martin |first5=C |last6=Hammel |first6=J. U. |last7=Zhang |first7=W |last8=Mayer |first8=G |date=2016 |title=Earliest Onychophoran in Amber Reveals Gondwanan Migration Patterns |journal=Current Biology |volume=26 |issue=19 |pages=2594–2601 |bibcode=2016CBio...26.2594O |doi=10.1016/j.cub.2016.07.023 |pmid=27693140 |doi-access=free}}</ref>'''<ref name=":033" /> All are from the same general location and date to a maximum age of around 98.79 million years ago (the earliest [[Cenomanian]] of the [[Late Cretaceous]]).<ref name=":033" /> Based on its [[Morphology (biology)|morphology]], ''Cretoperipatus'' was early-diverging member of [[Peripatidae]], most closely related to the [[Asia|Asian]] genera ''[[Eoperipatus]]'' and ''[[Typhloperipatus]]''.'''<ref name="Oliveira" />''' 

Despite being preserved in [[amber]], the affinities of the [[Cenozoic]] ''[[Tertiapatus]]'' and ''[[Succinipatopsis]]'' are surrounded in controversy. Some sources consider them to be onychophorans,<ref name="Jr2000" />'''<ref name="Oliveira" />''' while others dismiss this.<ref name="GarwoodEdgecombe2016" /> Ultimately, these animals should be reanalyzed to better grasp their position on the [[Tree of life (biology)|tree of life]].'''<ref name="Oliveira" />'''