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'''''Schinderhannes bartelsi''''' is an [[anomalocarid]] known from one specimen from the [[lower Devonian]] [[Hunsrück Slate]]s. Its discovery was astonishing because previously, [[anomalocaridid]]s were known only from exceptionally well-preserved fossil beds ([[Lagerstätte]]n) from the Cambrian, 100 million years earlier.<ref name=Kuhl2009>{{cite journal |author=Gabriele Kühl |author2=Derek E. G. Briggs |author3=Jes Rust |name-list-style=amp |year=2009 |title=A great-appendage arthropod with a radial mouth from the Lower Devonian Hunsrück Slate, Germany |journal=[[Science (journal)|Science]] |volume=323 |issue=5915 |pages=771–773 |pmid=19197061 |doi=10.1126/science.1166586 |bibcode=2009Sci...323..771K}}</ref>
'''''Schinderhannes bartelsi''''' is a species of [[Hurdiidae|hurdiid]] [[radiodont]] (anomalocaridid) known from one specimen from the [[lower Devonian]] [[Hunsrück Slate]]s. Its discovery was astonishing because previously, radiodonts were known only from exceptionally well-preserved fossil beds ([[Lagerstätte]]n) from the Cambrian, 100 million years earlier.<ref name=Kuhl2009>{{cite journal |author=Gabriele Kühl |author2=Derek E. G. Briggs |author3=Jes Rust |name-list-style=amp |year=2009 |title=A great-appendage arthropod with a radial mouth from the Lower Devonian Hunsrück Slate, Germany |journal=[[Science (journal)|Science]] |volume=323 |issue=5915 |pages=771–773 |pmid=19197061 |doi=10.1126/science.1166586 |bibcode=2009Sci...323..771K}}</ref>

Anomalocaridids, such as ''[[Anomalocaris]]'', were organisms determined to be basal arthropods. These creatures looked quite unlike any organism living today—they had segmented [[exoskeletons]], with lateral lobes used for swimming, typically large compound eyes, often set on stalks, and most strikingly, a pair of large, claw-like [[great appendage]]s that resembled headless shrimp. These appendages are thought to have passed food to the animal's mouth, which resembled a ring of sliced pineapple.<ref name=WonderfulLife>{{cite book |author=Stephen Jay Gould |author-link=Stephen Jay Gould |year=1989 |title=[[Wonderful Life (book)|Wonderful Life]]: The Burgess Shale and the Nature of History |publisher=W. W. Norton & Company |isbn=978-0-393-02705-1}}</ref>


==Discovery==
==Discovery==
Line 15: Line 13:


==Morphology==
==Morphology==
[[File:Schinderhannes_NT2.jpg|thumb|left|Outdated reconstruction of ''Schinderhannes.'' The reconstruction of putative trunk appandages are most likely an misinterpretation of setal blades and muscles.<ref name=":0">{{Cite journal|last=Ortega-Hernández|first=Javier|date=2016|title=Making sense of ‘lower’ and ‘upper’ stem-group Euarthropoda, with comments on the strict use of the name Arthropoda von Siebold, 1848: Upper and lower stem-Euarthropoda|url=http://doi.wiley.com/10.1111/brv.12168|journal=Biological Reviews|language=en|volume=91|issue=1|pages=255–273|doi=10.1111/brv.12168|via=}}</ref>]]
[[File:Schinderhannes_NT2.jpg|thumb|left|Life restoration]]
''Schinderhannes'' is about {{convert|10|cm}} long.<ref name="Kuhl2009" /> Like other radiodonts, the head bears a pair of spiny [[Radiodonta#Frontal appendage|frontal appendages]], a radially-arranged ventral mouthpart ([[Radiodonta#Oral cone|oral cone]]), and a pair of large lateral [[compound eyes]].<ref name="Kuhl2009" /> The eyes were in a relatively anterior position in contrast to other [[Hurdiidae|hurdiids]].<ref name=":1" /> Detailed morphology of the frontal appendages and oral cone are equivocal due to the limited preservation, but the former represent typical hurdiid features (e.g. subequal blade-like endites).<ref>{{Cite journal|last=Pates|first=Stephen|last2=Daley|first2=Allison C.|last3=Butterfield|first3=Nicholas J.|date=2019-06-11|title=First report of paired ventral endites in a hurdiid radiodont|url=https://doi.org/10.1186/s40851-019-0132-4|journal=Zoological Letters|volume=5|issue=1|pages=18|doi=10.1186/s40851-019-0132-4|issn=2056-306X|pmc=PMC6560863|pmid=31210962}}</ref><ref name=":1">{{Cite journal|last=Moysiuk|first=J.|last2=Caron|first2=J.-B.|date=2019-08-14|title=A new hurdiid radiodont from the Burgess Shale evinces the exploitation of Cambrian infaunal food sources|url=https://royalsocietypublishing.org/doi/10.1098/rspb.2019.1079|journal=Proceedings of the Royal Society B: Biological Sciences|volume=286|issue=1908|pages=20191079|doi=10.1098/rspb.2019.1079|pmc=PMC6710600|pmid=31362637}}</ref> There are traces of lateral structues originally thought to be the shaft regions of frontal appendages, which may rather represent [[Radiodonta#Head sclerites, eyes and trunk|P-elements]] (lateral sclerites) as seen in other radiodonts.<ref name=":1" /> The boundary of head and trunk ('neck') was broad with a pair of long, ventrally-protruded flaps.<ref name="Kuhl2009" /> The trunk compose of 12 body segments indicated by soft dorsal [[cuticle]]<ref name=":0" /> (originally thought to be rigid [[tergites]]<ref name="Kuhl2009" />)<ref name=Kuhl2009/>. The first 10 segments possess pairs of striated structures originally interpreted as biramous (branched) ventral flaps,<ref name="Kuhl2009" /> but later investigations from other radiodonts (e.g. ''[[Lyrarapax]]'') suggest it may rather represent setal blades (dorsal gill-like structures of radiodont) and flap [[muscles]].<ref name=":0" /> The 11th segment bears another pair of shorter, rounded flaps.<ref name="Kuhl2009" /> The final segment lacking appendages and terminated with a long, spine-like [[telson]].<ref name="Kuhl2009" />
''Schinderhannes'' is about {{convert|10|cm}} long; like other anomalocaridids, it bears a pair of great appendages (very similar to those of ''[[Hurdia]]''),<ref name=Daley2009/> a radial ''[[Peytoia]]'' 'pineapple-ring' mouth, and large, stalked, compound eyes. It has 12 body segments; large flap-like structures used for swimming protrude from just behind the head, and the 11th segment bears lateral
flukelike appendage.<ref name=Kuhl2009/>


==Ecology==
==Ecology==
The preserved contents of its digestive tract are typical of those of other predators',<ref name=Butterfield2002>{{cite journal |author=Nicholas J. Butterfield |year=2002 |title=''Leanchoilia'', and the interpretation of three-dimensional structures in Burgess Shale-type fossils |journal=[[Paleobiology (journal)|Paleobiology]] |volume=28 |issue=1 |pages=155–171 |doi=10.1666/0094-8373(2002)028<0155:LGATIO>2.0.CO;2 |jstor=3595514}}</ref> and this lifestyle is supported by the raptorial nature of the spiny great appendages and the size of the eyes.<ref name=Kuhl2009/> The organism was clearly a competent swimmer, propelling itself with the 'flippers' attached to its head, and using its wing-like lobes on the 11th segment to steer.<ref name=Kuhl2009/> These lobes presumably derived from the lateral lobes of Cambrian anomalocaridids, ancestors that used lobes along their sides to swim, and lacked the specializations of ''Schinderhannes''.<ref name=Kuhl2009/>
The preserved contents of its digestive tract are typical of those of other predators',<ref name=Butterfield2002>{{cite journal |author=Nicholas J. Butterfield |year=2002 |title=''Leanchoilia'', and the interpretation of three-dimensional structures in Burgess Shale-type fossils |journal=[[Paleobiology (journal)|Paleobiology]] |volume=28 |issue=1 |pages=155–171 |doi=10.1666/0094-8373(2002)028<0155:LGATIO>2.0.CO;2 |jstor=3595514}}</ref> and this lifestyle is supported by the raptorial nature of the spiny frontal appendages and the size of the eyes.<ref name=Kuhl2009/> ''Schinderhannes'' may had been a swimmer ([[nekton]]), propelling itself with the long flaps attached to its head, and using its shorter flaps on the 11th segment to steer.<ref name=Kuhl2009/> These flaps presumably derived from the lateral flaps of Cambrian radiodonts that used lobes along their sides to swim, but lacked the specializations as seen in ''Schinderhannes''.<ref name=Kuhl2009/>


==Significance==
==Significance==
The organism's discovery was most significant because of the huge range extension of the anomalocaridids it caused: the group was only previously known from lagerstätten of the lower-to-middle Cambrian, 100&nbsp;million years before. This underlined the utility of [[lagerstätte]]n like the [[Hunsrück Slate]]: these exceptionally preserved fossil horizons may be the only available opportunity to observe non-mineralised forms.<ref name=Butterfield1995>{{cite journal |author=Nicholas J. Butterfield |year=1995 |title=Secular distribution of Burgess-Shale-type preservation |journal=[[Lethaia]] |volume=28 |issue=1 |pages=1–13 |doi=10.1111/j.1502-3931.1995.tb01587.x}}</ref>
''Schinderhannes''<nowiki/>'s discovery was most significant because of the huge range extension of the radiodonts it caused: the group was only previously known from lagerstätten of the lower-to-middle [[Cambrian]], 100&nbsp;million years before. This underlined the utility of [[lagerstätte]]n like the [[Hunsrück Slate]]: these exceptionally preserved fossil horizons may be the only available opportunity to observe non-mineralised forms.<ref name=Butterfield1995>{{cite journal |author=Nicholas J. Butterfield |year=1995 |title=Secular distribution of Burgess-Shale-type preservation |journal=[[Lethaia]] |volume=28 |issue=1 |pages=1–13 |doi=10.1111/j.1502-3931.1995.tb01587.x}}</ref>


The discovery of ''Schinderhannes'' has also prompted novel hypothesis about the classification of basal [[arthropods]]. One classification scheme has ''Schinderhannes'' [[Sister group|sister]] to the euarthropods ([[Crown group|crown]] or 'true' arthropods) instead of other radiodonts, based on the characters which interpreted to be euarthropod-like (e.g. [[tergite]], [[Arthropod leg#Biramous and uniramous|biramous appendage]]). This would mean that the euarthropod lineage evolved from a [[paraphyletic]] grade of radiodonts, and that the group of basal arthropods with '[[great appendage|great/frontal appendage]]s' are not a natural grouping, and the biramous appendages of arthropods may then have arisen through fusion of radiodont lateral flaps and gills.<ref name=Kuhl2009/> However, this scenario had been challenged by later investigations, as the putative euarthropod-like features were questioned to be rather radiodont-like characters (e.g. soft trunk [[cuticle]], setal blades and paired flap muscles).<ref name=":0" /> [[Phylogenetic]] analysis with focus on [[Radiodonta]] also repeatly placed ''Schinderhannes'' within the radiodont family [[Hurdiidae]].<ref>{{Cite journal|last=Vinther|first=Jakob|last2=Stein|first2=Martin|last3=Longrich|first3=Nicholas R.|last4=Harper|first4=David A. T.|date=2014|title=A suspension-feeding anomalocarid from the Early Cambrian|url=https://www.nature.com/articles/nature13010|journal=Nature|language=en|volume=507|issue=7493|pages=496–499|doi=10.1038/nature13010|issn=1476-4687|via=}}</ref><ref>{{Cite journal|last=Cong|first=Peiyun|last2=Ma|first2=Xiaoya|last3=Hou|first3=Xianguang|last4=Edgecombe|first4=Gregory D.|last5=Strausfeld|first5=Nicholas J.|date=2014|title=Brain structure resolves the segmental affinity of anomalocaridid appendages|url=https://www.nature.com/articles/nature13486|journal=Nature|language=en|volume=513|issue=7519|pages=538–542|doi=10.1038/nature13486|issn=1476-4687|via=}}</ref><ref>{{Cite journal|last=Van Roy|first=Peter|last2=Daley|first2=Allison C.|last3=Briggs|first3=Derek E. G.|date=2015|title=Anomalocaridid trunk limb homology revealed by a giant filter-feeder with paired flaps|url=https://www.nature.com/articles/nature14256|journal=Nature|language=en|volume=522|issue=7554|pages=77–80|doi=10.1038/nature14256|issn=1476-4687|via=}}</ref><ref>{{Cite journal|last=Lerosey-Aubril|first=Rudy|last2=Pates|first2=Stephen|date=2018-09-14|title=New suspension-feeding radiodont suggests evolution of microplanktivory in Cambrian macronekton|url=https://www.nature.com/articles/s41467-018-06229-7|journal=Nature Communications|language=en|volume=9|issue=1|pages=3774|doi=10.1038/s41467-018-06229-7|issn=2041-1723|pmc=PMC6138677|pmid=30218075}}</ref><ref name=":1" />
The organism has also prompted novel hypotheses about the classification of early arthropods. One classification scheme has ''Schinderhannes'' classified basally to the crown arthropods, but closer to that group than ''Anomalocaris''. This would mean that the crown arthropod lineage evolved from a paraphyletic grade of anomalocaridids, and that the group of early arthropods with short '[[great appendage]]s' are not a natural grouping.<ref name=Kuhl2009/> The [[biramous limb]] of arthropods may then have arisen through fusion of anomalocaridid lateral lobes and gills.<ref name=Daley2009>{{cite journal |author=Allison C. Daley |author2=Graham E. Budd |author3=Jean-Bernard Caron|author4=Gregory D. Edgecombe |author5=Desmond Collins |name-list-style=amp |year=2009 |title=The Burgess Shale anomalocaridid ''Hurdia'' and its significance for early euarthropod evolution |journal=[[Science (journal)|Science]] |volume=323 |issue=5921 |pages=1597–1600 |pmid=19299617 |doi=10.1126/science.1169514 |bibcode=2009Sci...323.1597D}}</ref>


==References==
==References==

Revision as of 15:08, 7 February 2021

Schinderhannes bartelsi
Temporal range: Early Devonian 408–400 Ma
The holotype of Schinderhannes. Credit: Steinmann Institute/University of Bonn
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: †Dinocaridida
Order: †Radiodonta
Family: †Hurdiidae
Genus: †Schinderhannes
Species:
†S. bartelsi
Binomial name
†Schinderhannes bartelsi
Kühl, Briggs & Rust, 2009

Schinderhannes bartelsi is a species of hurdiid radiodont (anomalocaridid) known from one specimen from the lower Devonian Hunsrück Slates. Its discovery was astonishing because previously, radiodonts were known only from exceptionally well-preserved fossil beds (Lagerstätten) from the Cambrian, 100 million years earlier.[1]

Discovery

The single specimen was discovered in the Eschenbach-Bocksberg Quarry in Bundenbach, and is named after the outlaw Schinderhannes who frequented the area. Its specific epithet bartelsi honours Christoph Bartels, a Hunsrück Slate expert. The specimen is now housed in the Naturhistorisches Museum, Mainz.[1]

Morphology

Outdated reconstruction of Schinderhannes. The reconstruction of putative trunk appandages are most likely an misinterpretation of setal blades and muscles.[2]

Schinderhannes is about 10 centimetres (3.9 in) long.[1] Like other radiodonts, the head bears a pair of spiny frontal appendages, a radially-arranged ventral mouthpart (oral cone), and a pair of large lateral compound eyes.[1] The eyes were in a relatively anterior position in contrast to other hurdiids.[3] Detailed morphology of the frontal appendages and oral cone are equivocal due to the limited preservation, but the former represent typical hurdiid features (e.g. subequal blade-like endites).[4][3] There are traces of lateral structues originally thought to be the shaft regions of frontal appendages, which may rather represent P-elements (lateral sclerites) as seen in other radiodonts.[3] The boundary of head and trunk ('neck') was broad with a pair of long, ventrally-protruded flaps.[1] The trunk compose of 12 body segments indicated by soft dorsal cuticle[2] (originally thought to be rigid tergites[1])[1]. The first 10 segments possess pairs of striated structures originally interpreted as biramous (branched) ventral flaps,[1] but later investigations from other radiodonts (e.g. Lyrarapax) suggest it may rather represent setal blades (dorsal gill-like structures of radiodont) and flap muscles.[2] The 11th segment bears another pair of shorter, rounded flaps.[1] The final segment lacking appendages and terminated with a long, spine-like telson.[1]

Ecology

The preserved contents of its digestive tract are typical of those of other predators',[5] and this lifestyle is supported by the raptorial nature of the spiny frontal appendages and the size of the eyes.[1] Schinderhannes may had been a swimmer (nekton), propelling itself with the long flaps attached to its head, and using its shorter flaps on the 11th segment to steer.[1] These flaps presumably derived from the lateral flaps of Cambrian radiodonts that used lobes along their sides to swim, but lacked the specializations as seen in Schinderhannes.[1]

Significance

Schinderhannes's discovery was most significant because of the huge range extension of the radiodonts it caused: the group was only previously known from lagerstätten of the lower-to-middle Cambrian, 100 million years before. This underlined the utility of lagerstätten like the Hunsrück Slate: these exceptionally preserved fossil horizons may be the only available opportunity to observe non-mineralised forms.[6]

The discovery of Schinderhannes has also prompted novel hypothesis about the classification of basal arthropods. One classification scheme has Schinderhannes sister to the euarthropods (crown or 'true' arthropods) instead of other radiodonts, based on the characters which interpreted to be euarthropod-like (e.g. tergite, biramous appendage). This would mean that the euarthropod lineage evolved from a paraphyletic grade of radiodonts, and that the group of basal arthropods with 'great/frontal appendages' are not a natural grouping, and the biramous appendages of arthropods may then have arisen through fusion of radiodont lateral flaps and gills.[1] However, this scenario had been challenged by later investigations, as the putative euarthropod-like features were questioned to be rather radiodont-like characters (e.g. soft trunk cuticle, setal blades and paired flap muscles).[2] Phylogenetic analysis with focus on Radiodonta also repeatly placed Schinderhannes within the radiodont family Hurdiidae.[7][8][9][10][3]

References

  1. ^ a b c d e f g h i j k l m n Gabriele Kühl; Derek E. G. Briggs & Jes Rust (2009). "A great-appendage arthropod with a radial mouth from the Lower Devonian Hunsrück Slate, Germany". Science. 323 (5915): 771–773. Bibcode:2009Sci...323..771K. doi:10.1126/science.1166586. PMID 19197061.
  2. ^ a b c d Ortega-Hernández, Javier (2016). "Making sense of 'lower' and 'upper' stem-group Euarthropoda, with comments on the strict use of the name Arthropoda von Siebold, 1848: Upper and lower stem-Euarthropoda". Biological Reviews. 91 (1): 255–273. doi:10.1111/brv.12168.
  3. ^ a b c d Moysiuk, J.; Caron, J.-B. (2019-08-14). "A new hurdiid radiodont from the Burgess Shale evinces the exploitation of Cambrian infaunal food sources". Proceedings of the Royal Society B: Biological Sciences. 286 (1908): 20191079. doi:10.1098/rspb.2019.1079. PMC 6710600. PMID 31362637.{{cite journal}}: CS1 maint: PMC format (link)
  4. ^ Pates, Stephen; Daley, Allison C.; Butterfield, Nicholas J. (2019-06-11). "First report of paired ventral endites in a hurdiid radiodont". Zoological Letters. 5 (1): 18. doi:10.1186/s40851-019-0132-4. ISSN 2056-306X. PMC 6560863. PMID 31210962.{{cite journal}}: CS1 maint: PMC format (link) CS1 maint: unflagged free DOI (link)
  5. ^ Nicholas J. Butterfield (2002). "Leanchoilia, and the interpretation of three-dimensional structures in Burgess Shale-type fossils". Paleobiology. 28 (1): 155–171. doi:10.1666/0094-8373(2002)028<0155:LGATIO>2.0.CO;2. JSTOR 3595514.
  6. ^ Nicholas J. Butterfield (1995). "Secular distribution of Burgess-Shale-type preservation". Lethaia. 28 (1): 1–13. doi:10.1111/j.1502-3931.1995.tb01587.x.
  7. ^ Vinther, Jakob; Stein, Martin; Longrich, Nicholas R.; Harper, David A. T. (2014). "A suspension-feeding anomalocarid from the Early Cambrian". Nature. 507 (7493): 496–499. doi:10.1038/nature13010. ISSN 1476-4687.
  8. ^ Cong, Peiyun; Ma, Xiaoya; Hou, Xianguang; Edgecombe, Gregory D.; Strausfeld, Nicholas J. (2014). "Brain structure resolves the segmental affinity of anomalocaridid appendages". Nature. 513 (7519): 538–542. doi:10.1038/nature13486. ISSN 1476-4687.
  9. ^ Van Roy, Peter; Daley, Allison C.; Briggs, Derek E. G. (2015). "Anomalocaridid trunk limb homology revealed by a giant filter-feeder with paired flaps". Nature. 522 (7554): 77–80. doi:10.1038/nature14256. ISSN 1476-4687.
  10. ^ Lerosey-Aubril, Rudy; Pates, Stephen (2018-09-14). "New suspension-feeding radiodont suggests evolution of microplanktivory in Cambrian macronekton". Nature Communications. 9 (1): 3774. doi:10.1038/s41467-018-06229-7. ISSN 2041-1723. PMC 6138677. PMID 30218075.{{cite journal}}: CS1 maint: PMC format (link)

External links

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