Archaeamphora: Difference between revisions

Archaeamphora; Temporal range: Early Cretaceous PreꞒ Ꞓ O S D C P T J K Pg N
	Fossils of A. longicervia described in Wong et al. (2015)
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Gymnospermae
Division:	Pinophyta
Class:	Pinopsida
Order:	Pinales
Family:	incertae sedis
Genus:	†Archaeamphora; H.Q.Li
Species:	†A. longicervia
Binomial name
	†Archaeamphora longicervia; H.Q.Li (2005)

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Archaeamphora longicervia is a fossil plant species, the only member of the hypothetical genus Archaeamphora. Fossil material assigned to this taxon originates from the Yixian Formation of northeastern China, dated to the Early Cretaceous (around 145 to 101 million years ago).^[1]

The species was originally described as a pitcher plant with close affinities to extant members of the family Sarraceniaceae. This would make it the earliest known carnivorous plant and the only known fossil record of Sarraceniaceae, or the New World pitcher plant family.^[2]^[3]^[4]^[5] Archaeamphora is also one of the three oldest known genera of angiosperms (flowering plants). Li (2005) wrote that "the existence of a so highly derived Angiosperm in the Early Cretaceous suggests that Angiosperms should have originated much earlier, maybe back to 280 mya as the molecular clock studies suggested".^[1]

Subsequent authors have questioned the identification of Archaeamphora as a pitcher plant and a taxon of angiosperm at all.^[6]^[7]^[8] The fossils more probably represent leaves (needles) of the coniferous Liaoningocladus boii deformed by insect galls.^[9]

Etymology

The generic name Archaeamphora is derived from the Greek αρχαίος, archaíos ("ancient"; combining form in Latin: archae-), and ἀμφορεύς, amphoreus ("pitcher"). The specific epithet longicervia is derived from the Latin longus ("long") and cervicarius ("with a neck"), in reference to the characteristic constriction in the pitcher-like structures of this species.^[1]

Fossil material

All known fossil material of A. longicervia originates from the Jianshangou Formation in Beipiao, western Liaoning, China. These Early Cretaceous beds constitute the lower part of the Yixian Formation,^[10]^[11] which is dated at 124.6 million years old.^[12] Nine specimens of A. longicervia have been found, including holotype CBO0220 and paratype CBO0754.^[1]

Description

Archaeamphora longicervia was supposed to be a herbaceous plant growing to around 50 mm (2.0 in) in height. The stem, at least 21 mm (0.83 in) long by 1.2 mm (0.047 in) wide, bore distinctive vertical ridges and grooves. The pitcher-like structures were ascidiate in form and 30 to 40 mm (1.2 to 1.6 in) long. Mature pitchers and underdeveloped pitchers or phyllodia-like leaves were arranged spirally around the stem. Pitchers consisted of a tubular base, expanded middle section, constriction around the mouth, and a vertical, spoon-shaped lid. A single wing ran down the adaxial side of each pitcher. Three to five parallel major veins were present on the pitchers, along with a few intercostal veins and numerous small veinlets.^[1]

Two unusual bag-like structures were present on each pitcher, one on either side of the central wing. Similar but semi-circular structures were found on the margin of the lid. These structures exhibited strong yellow-green intrinsic fluorescence when exposed to visible light with a wavelength of 500 nm (blue-green).^[1]

Tiny glands, approximately 4 μm in diameter, were found on the inner surface of the pitchers and partially embedded in the grooves along the veins. These also showed very strong golden-yellow fluorescence.^[1]

A single seed was found intimately associated with the fossil material of A. longicervia and is presumed to belong to the same species. It is winged and reticulate-tuberculate in morphology, closely resembling the seeds of Sarraceniaceae taxa. The seed is oval-shaped, covered with black-brown warts, and measures 0.9 by 1.25 mm (0.035 by 0.049 in).^[1]

Taxonomy

The fossil material of A. longicervia was subjected to chemical analysis for oleanane, considered a key marker differentiating angiosperms from gymnosperms.^[13] Oleanane was detected in these specimens, suggesting that they belong to the angiosperms.^[1]

Pitcher plant interpretation

According to Li (2005), several morphological features of A. longicervia indicate a close relationship to Sarraceniaceae: both taxa exhibit one or two pitcher wings, a smooth peristome, and pitchers that extend vertically from the top of a short petiole.^[1]

Li (2005) suggests that A. longicervia is morphologically similar to modern Sarracenia purpurea. It shares with this species the spiral arrangement of its pitchers and phyllodia-like tubular leaves with parallel major veins. Archaeamphora longicervia also shows a resemblance to species of the genus Heliamphora in having pitchers with a long neck and upright lid. Of particular note is the similarity between the thick semi-circular structures on the lid of A. longicervia and the large nectar-secreting "bubble" present on the upper posterior portion of Heliamphora exappendiculata^{[nb 1]} pitchers.^[1]

Li (2005) mentions the discovery of another type of "pitcher plant" from the same formation. This variety differs from the type material of A. longicervia in having pitchers that lack any constriction before the mouth, instead gradually expanding from the petiole into a hollow trumpet-like shape. He suggests that it "should be a different species" from A. longicervia. An intermediate form with a wider neck is also reported, suggesting that these plants were already a diversified group in the Early Cretaceous.^[1]

Current understanding

Heřmanová & Kvaček (2010) opined that the pitcher plant interpretation of Archaeamphora is "problematic and the fossil is in need of revision".^[7]

In their 2011 book, Sarraceniaceae of South America, McPherson et al. summarised current thinking on Archaeamphora as follows:^[6]

Serious doubt is emerging that reduces the likelihood that Archaeamphora longicervia belongs in the Sarraceniaceae lineage, or was even a pitcher plant at all. [...] Although Archaeamphora might well be a representative of the earliest flowering plants on Earth [...] it is very unlikely that it represents an ancestor of Sarraceniaceae since it is much too old to be part of the advanced "crown group" of Ericales to which Sarraceniaceae belong. [...] Another contradiction is that except for Archaeamphora, there is no other evidence to suggest that Sarraceniaceae evolved outside the New World, to which all extant members of the family are endemic.

Wong et al. (2015) put forward a new perspective as follows:^[9]

Archaeamphora longicervia H. Q. Li was described as an herbaceous, Sarraceniaceae-like pitcher plant from the mid Early Cretaceous Yixian Formation of Liaoning Province, northeastern China. Here, a re-investigation of A. longicervia specimens from the Yixian Formation provides new insights into its identity and the morphology of pitcher plants claimed by Li. We demonstrate that putative pitchers of Archaeamphora are insect-induced leaf galls that consist of three components: (1) an innermost larval chamber; (2) an intermediate zone of nutritive tissue; and (3) an outermost wall of sclerenchyma. Archaeamphora is not a carnivorous, Sarraceniaceae-like angiosperm, but represents insect-galled leaves of the previously reported gymnosperm Liaoningocladus boii G. Sun et al. from the Yixian Formation.

Habitat

The area inhabited by A. longicervia is thought to have experienced significant climatic fluctuations during the Early Cretaceous, ranging from arid or semi-arid to more humid conditions.^[15] The substrate in the region was mostly composed of lacustrine sediments and volcanic rocks.^[16]

Notes

^ This taxon was recognised as a variety of Heliamphora heterodoxa when Archaeamphora was described in 2005.^[1] It was raised to species rank the following year.^[14]

References

^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o Li, H. 2005. "Early Cretaceous sarraceniacean-like pitcher plants from China" (PDF). Archived from the original (PDF) on 2006-09-02. Retrieved 2007-01-27. Acta Botanica Gallica 152(2): 227-234. (Supplementary Information 1 Archived 2016-02-22 at the Wayback Machine, Supplementary Information 2 Archived 2016-06-30 at the Wayback Machine)
^ Krutzsch, W (1985). "Über Nepenthes-Pollen im europäischen Tertiär". Gleditschia. 13: 89–93.
^ Krutzsch, W (1989). "Paleogeography and historical phytogeography (paleochorology) in the Neophyticum". Plant Systematics and Evolution. 162 (1–4): 5–61. Bibcode:1989PSyEv.162....5K. doi:10.1007/BF00936909. S2CID 10454432.
^ Anderson, J.A.R.; Muller, J. (1975). "Palynological study of a Holocene peat and a Miocene coal deposit from NW Borneo". Review of Palaeobotany and Palynology. 19 (4): 291–351. Bibcode:1975RPaPa..19..291A. doi:10.1016/0034-6667(75)90049-4.
^ Kumar, M (1995). "Pollen tetrads from Palaeocene sediments of Meghalaya, India: comments on their morphology, botanical affinity and geological records". Palaeobotanist. 43 (1): 68–81.
^ ^a ^b McPherson, S., A. Wistuba, A. Fleischmann & J. Nerz 2011. Sarraceniaceae of South America. Redfern Natural History Productions, Poole.
^ ^a ^b Heřmanová, Z.; Kvaček, J. (2010). "Late Cretaceous Palaeoaldrovanda, not seeds of a carnivorous plant, but eggs of an insect". Journal of the National Museum (Prague), Natural History Series. 179 (9): 105–118. Archived from the original on 2015-10-09. Retrieved 2012-10-26.
^ Brittnacher, J (2013). "Phylogeny and biogeography of the Sarraceniaceae". Carnivorous Plant Newsletter. 42 (3): 99–106. doi:10.55360/cpn423.jb578. S2CID 247114275.
^ ^a ^b Wong, WO; Dilcher, DL; Labandeira, CC; Sun, G; Fleischmann, A (2015). "Early Cretaceous Archaeamphora is not a carnivorous angiosperm". Frontiers in Plant Science. 6: 326. doi:10.3389/fpls.2015.00326. PMC 4423337. PMID 25999978.
^ Sun, G., S.-L. Zheng, D.L. Dilcher, Y.D. Wang & S.W Mei 2001. Early Angiosperms and their Associated Plants from Western Liaoning, China. Shanghai Scientific and Technological Education Publishing House, 227 pp.
^ Sun, G.; Dilcher, D.L.; Zheng, S.-L.; Zhou, Z.-K. (1998). "In Search of the First Flower: A Jurassic Angiosperm, Archaefructus, from Northeast China" (PDF). Science. 282 (5394): 1692–1695. doi:10.1126/science.282.5394.1692. PMID 9831557. Archived from the original (PDF) on 2011-07-19.
^ Swisher, III; Wang, Y-Q.; Wang, X.-L.; Xu, X.; Wang, Y. (1999). "Cretaceous age for the feathered dinosaurs of Liaoning, China" (PDF). Nature. 400 (6739): 58–61. Bibcode:1999Natur.400...58S. doi:10.1038/21872. S2CID 4424629.
^ Moldowan, J.M.; Dahl, J.; Huizinga, B.J.; Fago, F.J.; Hickey, L.J.; Peakman, T.M.; Taylor, O.W. (1994). "The molecular fossil record of oleanane and its relationship to Angiosperms". Science. 265 (5173): 768–771. doi:10.1126/science.265.5173.768. PMID 17736275. S2CID 9389874.
^ Nerz, J.; Wistuba, A. (2006). "Heliamphora exappendiculata, a clearly distinct species with unique characteristics". Carnivorous Plant Newsletter. 35 (2): 43–51. doi:10.55360/cpn352.jn909. S2CID 247149195.
^ Barrett, P.M.; Hilton, J. M. (2006). "The Jehol Biota (Lower Cretaceous, China): new discoveries and future prospects". Integrative Zoology. 1 (1): 15–17. doi:10.1111/j.1749-4877.2006.00006.x. PMID 21395985.
^ Tan, J.-J.; Ren, D. (2006). "Ovatucupes: A New Cupedid Genus (Coleoptera: Archostemata: Cupedidae) From The Jehol Biota (Late Jurassic) Of Western Liaoning, China" (PDF). Entomological News. 117 (2): 223–232. doi:10.3157/0013-872x(2006)117[223:oancgc]2.0.co;2. S2CID 84331871.

[exa-15] This taxon was recognised as a variety of Heliamphora heterodoxa when Archaeamphora was described in 2005.^[1] It was raised to species rank the following year.^[14]

[Li-1] ^ ^a ^b ^c ^d ^e ^f ^g ^h ⁱ ^j ^k ^l ^m ⁿ ^o Li, H. 2005. "Early Cretaceous sarraceniacean-like pitcher plants from China" (PDF). Archived from the original (PDF) on 2006-09-02. Retrieved 2007-01-27. Acta Botanica Gallica 152(2): 227-234. (Supplementary Information 1 Archived 2016-02-22 at the Wayback Machine, Supplementary Information 2 Archived 2016-06-30 at the Wayback Machine)

[Krutzsch1985-2] Krutzsch, W (1985). "Über Nepenthes-Pollen im europäischen Tertiär". Gleditschia. 13: 89–93.

[Krutzsch1989-3] Krutzsch, W (1989). "Paleogeography and historical phytogeography (paleochorology) in the Neophyticum". Plant Systematics and Evolution. 162 (1–4): 5–61. Bibcode:1989PSyEv.162....5K. doi:10.1007/BF00936909. S2CID 10454432.

[4] Anderson, J.A.R.; Muller, J. (1975). "Palynological study of a Holocene peat and a Miocene coal deposit from NW Borneo". Review of Palaeobotany and Palynology. 19 (4): 291–351. Bibcode:1975RPaPa..19..291A. doi:10.1016/0034-6667(75)90049-4.

[5] Kumar, M (1995). "Pollen tetrads from Palaeocene sediments of Meghalaya, India: comments on their morphology, botanical affinity and geological records". Palaeobotanist. 43 (1): 68–81.

[McPherson2011-6] McPherson, S., A. Wistuba, A. Fleischmann & J. Nerz 2011. Sarraceniaceae of South America. Redfern Natural History Productions, Poole.

[eggs-7] Heřmanová, Z.; Kvaček, J. (2010). "Late Cretaceous Palaeoaldrovanda, not seeds of a carnivorous plant, but eggs of an insect". Journal of the National Museum (Prague), Natural History Series. 179 (9): 105–118. Archived from the original on 2015-10-09. Retrieved 2012-10-26.

[8] Brittnacher, J (2013). "Phylogeny and biogeography of the Sarraceniaceae". Carnivorous Plant Newsletter. 42 (3): 99–106. doi:10.55360/cpn423.jb578. S2CID 247114275.

[Wong-2015-9] Wong, WO; Dilcher, DL; Labandeira, CC; Sun, G; Fleischmann, A (2015). "Early Cretaceous Archaeamphora is not a carnivorous angiosperm". Frontiers in Plant Science. 6: 326. doi:10.3389/fpls.2015.00326. PMC 4423337. PMID 25999978.

[10] Sun, G., S.-L. Zheng, D.L. Dilcher, Y.D. Wang & S.W Mei 2001. Early Angiosperms and their Associated Plants from Western Liaoning, China. Shanghai Scientific and Technological Education Publishing House, 227 pp.

[11] Sun, G.; Dilcher, D.L.; Zheng, S.-L.; Zhou, Z.-K. (1998). "In Search of the First Flower: A Jurassic Angiosperm, Archaefructus, from Northeast China" (PDF). Science. 282 (5394): 1692–1695. doi:10.1126/science.282.5394.1692. PMID 9831557. Archived from the original (PDF) on 2011-07-19.

[12] Swisher, III; Wang, Y-Q.; Wang, X.-L.; Xu, X.; Wang, Y. (1999). "Cretaceous age for the feathered dinosaurs of Liaoning, China" (PDF). Nature. 400 (6739): 58–61. Bibcode:1999Natur.400...58S. doi:10.1038/21872. S2CID 4424629.

[13] Moldowan, J.M.; Dahl, J.; Huizinga, B.J.; Fago, F.J.; Hickey, L.J.; Peakman, T.M.; Taylor, O.W. (1994). "The molecular fossil record of oleanane and its relationship to Angiosperms". Science. 265 (5173): 768–771. doi:10.1126/science.265.5173.768. PMID 17736275. S2CID 9389874.

[14] Nerz, J.; Wistuba, A. (2006). "Heliamphora exappendiculata, a clearly distinct species with unique characteristics". Carnivorous Plant Newsletter. 35 (2): 43–51. doi:10.55360/cpn352.jn909. S2CID 247149195.

[16] Barrett, P.M.; Hilton, J. M. (2006). "The Jehol Biota (Lower Cretaceous, China): new discoveries and future prospects". Integrative Zoology. 1 (1): 15–17. doi:10.1111/j.1749-4877.2006.00006.x. PMID 21395985.

[17] Tan, J.-J.; Ren, D. (2006). "Ovatucupes: A New Cupedid Genus (Coleoptera: Archostemata: Cupedidae) From The Jehol Biota (Late Jurassic) Of Western Liaoning, China" (PDF). Entomological News. 117 (2): 223–232. doi:10.3157/0013-872x(2006)117[223:oancgc]2.0.co;2. S2CID 84331871.

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@@ Line 1: / Line 1: @@
-{{short description|Fossil species of Cretaceous-aged flowering plant}}
+{{Short description|Fossil species of Cretaceous-aged flowering plant}}
-{{good article}}
+{{Good article}}
+{{Speciesbox
-{{taxobox
 | fossil_range = {{Geological range/linked|Early Cretaceous}}
-| image = Archaeamphora longicervia.png
+| image = Archaeamphora_fossils.jpg
+| image_alt =
-| image_caption = Artist's reconstruction of ''Archaeamphora longicervia'' after Li (2005)<ref name=Li />
+| image_caption = Fossils of ''A. longicervia'' described in Wong ''et al.'' (2015)
-| regnum = [[Plantae]]
+| genus = Archaeamphora
-| unranked_divisio = [[Angiosperms]]
+| parent_authority = H.Q.Li
-| unranked_classis = ?[[Eudicots]]
+| species = longicervia
-| unranked_ordo = ?[[Asterids]]
+| authority = [[Hongqi Li|H.Q.Li]] (2005)<ref name=Li />
-| ordo = ?[[Ericales]]
+| synonyms =
-| familia = ?[[Sarraceniaceae]]
+| synonyms_ref =
-| genus = '''''Archaeamphora'''''
-| species = '''''A. longicervia'''''
-| binomial = ''Archaeamphora longicervia''
-| binomial_authority = [[Hongqi Li|Li]] (2005)<ref name=Li />
 }}
-'''''Archaeamphora longicervia''''' is a disputed fossil plant species attributed [[extinction|extinct]] species of [[flowering plant]]s and the [[Monotypic taxon|only member]] of the genus '''''Archaeamphora'''''. Fossil material assigned to this [[taxon]] originates from the [[Yixian Formation]] of northeastern [[China]], dated to the [[Early Cretaceous]] (around {{Period span/brief|Early Cretaceous}}).<ref name=Li>Li, H. 2005. {{cite web |url= http://faculty.frostburg.edu/biol/hli/research/Archaeamphora.pdf |title= Early Cretaceous sarraceniacean-like pitcher plants from China. |access-date= 2007-01-27 |archive-url= https://web.archive.org/web/20060902062633/http://faculty.frostburg.edu/biol/hli/research/Archaeamphora.pdf |archive-date= 2006-09-02 |url-status= dead }} ''Acta Botanica Gallica'' '''152'''(2): 227-234. ([http://faculty.frostburg.edu/biol/hli/research/Pitcher/SIGuide.htm Supplementary Information 1], [http://faculty.frostburg.edu/biol/hli/research/Pitcher/Fig-SupplementaryMateral.htm Supplementary Information 2])</ref>
+'''''Archaeamphora longicervia''''' is a fossil plant species, the [[Monotypic taxon|only member]] of the hypothetical genus '''''Archaeamphora'''''. Fossil material assigned to this [[taxon]] originates from the [[Yixian Formation]] of northeastern [[China]], dated to the [[Early Cretaceous]] (around {{Period span/brief|Early Cretaceous}}).<ref name=Li>Li, H. 2005. {{cite web |url= http://faculty.frostburg.edu/biol/hli/research/Archaeamphora.pdf |title= Early Cretaceous sarraceniacean-like pitcher plants from China. |access-date= 2007-01-27 |archive-url= https://web.archive.org/web/20060902062633/http://faculty.frostburg.edu/biol/hli/research/Archaeamphora.pdf |archive-date= 2006-09-02 |url-status= dead }} ''Acta Botanica Gallica'' '''152'''(2): 227-234. ([http://faculty.frostburg.edu/biol/hli/research/Pitcher/SIGuide.htm Supplementary Information 1] {{Webarchive|url=https://web.archive.org/web/20160222131935/http://faculty.frostburg.edu/biol/hli/research/Pitcher/SIGuide.htm |date=2016-02-22 }}, [http://faculty.frostburg.edu/biol/hli/research/Pitcher/Fig-SupplementaryMateral.htm Supplementary Information 2] {{Webarchive|url=https://web.archive.org/web/20160630073138/http://faculty.frostburg.edu/biol/hli/research/Archaeamphora.pdf |date=2016-06-30 }})</ref>
-The species was originally described as a [[pitcher plant]] with close affinities to [[Extant taxon|extant]] members of the [[family (biology)|family]] [[Sarraceniaceae]]. This would make it the earliest known [[carnivorous plant]] and the only known fossil record of Sarraceniaceae, or the New World pitcher plant family.<ref name=Krutzsch1985>{{cite journal | last1 = Krutzsch | first1 = W | year = 1985 | title = Über ''Nepenthes''-Pollen im europäischen Tertiär | url = | journal = Gleditschia | volume = 13 | issue = | pages = 89–93 }}</ref><ref name=Krutzsch1989>{{cite journal | last1 = Krutzsch | first1 = W | year = 1989 | title = Paleogeography and historical phytogeography (paleochorology) in the Neophyticum | url = | journal = Plant Systematics and Evolution | volume = 162 | issue = 1–4| pages = 5–61 | doi = 10.1007/BF00936909 }}</ref><ref>{{cite journal | last1 = Anderson | first1 = J.A.R. | last2 = Muller | first2 = J. | year = 1975 | title = Palynological study of a Holocene peat and a Miocene coal deposit from NW Borneo | url = | journal = Review of Palaeobotany and Palynology | volume = 19 | issue = 4| pages = 291–351 | doi=10.1016/0034-6667(75)90049-4}}</ref><ref>{{cite journal | last1 = Kumar | first1 = M | year = 1995 | title = Pollen tetrads from Palaeocene sediments of Meghalaya, India: comments on their morphology, botanical affinity and geological records | url = | journal = Palaeobotanist | volume = 43 | issue = 1| pages = 68–81 }}</ref> ''Archaeamphora'' is also one of the three oldest known [[genus|genera]] of [[flowering plant|angiosperms]] (flowering plants). Li (2005) wrote that "the existence of a so highly derived Angiosperm in the Early Cretaceous suggests that Angiosperms should have originated much earlier, maybe back to 280 mya as the [[molecular clock]] studies suggested".<ref name=Li />
+The species was originally described as a [[pitcher plant]] with close affinities to [[Extant taxon|extant]] members of the [[family (biology)|family]] [[Sarraceniaceae]]. This would make it the earliest known [[carnivorous plant]] and the only known fossil record of Sarraceniaceae, or the New World pitcher plant family.<ref name=Krutzsch1985>{{cite journal | last1 = Krutzsch | first1 = W | year = 1985 | title = Über ''Nepenthes''-Pollen im europäischen Tertiär | journal = Gleditschia | volume = 13 | pages = 89–93 }}</ref><ref name=Krutzsch1989>{{cite journal | last1 = Krutzsch | first1 = W | year = 1989 | title = Paleogeography and historical phytogeography (paleochorology) in the Neophyticum | journal = Plant Systematics and Evolution | volume = 162 | issue = 1–4| pages = 5–61 | doi = 10.1007/BF00936909 | bibcode = 1989PSyEv.162....5K | s2cid = 10454432 }}</ref><ref>{{cite journal | last1 = Anderson | first1 = J.A.R. | last2 = Muller | first2 = J. | year = 1975 | title = Palynological study of a Holocene peat and a Miocene coal deposit from NW Borneo | journal = Review of Palaeobotany and Palynology | volume = 19 | issue = 4| pages = 291–351 | doi=10.1016/0034-6667(75)90049-4| bibcode = 1975RPaPa..19..291A }}</ref><ref>{{cite journal | last1 = Kumar | first1 = M | year = 1995 | title = Pollen tetrads from Palaeocene sediments of Meghalaya, India: comments on their morphology, botanical affinity and geological records | journal = Palaeobotanist | volume = 43 | issue = 1| pages = 68–81 }}</ref> ''Archaeamphora'' is also one of the three oldest known [[genus|genera]] of [[flowering plant|angiosperms]] (flowering plants). Li (2005) wrote that "the existence of a so highly derived Angiosperm in the Early Cretaceous suggests that Angiosperms should have originated much earlier, maybe back to 280 mya as the [[molecular clock]] studies suggested".<ref name=Li />
-Subsequent authors have questioned the identification of ''Archaeamphora'' as a pitcher plant.<ref name=McPherson2011>McPherson, S., A. Wistuba, A. Fleischmann & J. Nerz 2011. ''[[Sarraceniaceae of South America]]''. Redfern Natural History Productions, Poole.</ref><ref name=eggs>{{cite journal | last1 = Heřmanová | first1 = Z. | last2 = Kvaček | first2 = J. | year = 2010 | title = Late Cretaceous ''Palaeoaldrovanda'', not seeds of a carnivorous plant, but eggs of an insect | url = http://www.nm.cz/publikace/publikace-download.php?name=File1&dir=archiv&table=tabPublikaceArchiv&id=2785 | journal = Journal of the National Museum (Prague), Natural History Series | volume = 179 | issue = 9 | pages = 105–118 | access-date = 2012-10-26 | archive-url = https://web.archive.org/web/20151009191247/http://www.nm.cz/publikace/publikace-download.php?name=File1&dir=archiv&table=tabPublikaceArchiv&id=2785 | archive-date = 2015-10-09 | url-status = dead }}</ref><ref>{{cite journal | last1 = Brittnacher | first1 = J | year = 2013 | title = Phylogeny and biogeography of the Sarraceniaceae | url = | journal = [[Carnivorous Plant Newsletter]] | volume = 42 | issue = 3| pages = 99–106 }}</ref>
+Subsequent authors have questioned the identification of ''Archaeamphora'' as a pitcher plant and a taxon of [[Flowering plant|angiosperm]] at all.<ref name=McPherson2011>McPherson, S., A. Wistuba, A. Fleischmann & J. Nerz 2011. ''[[Sarraceniaceae of South America]]''. Redfern Natural History Productions, Poole.</ref><ref name=eggs>{{cite journal | last1 = Heřmanová | first1 = Z. | last2 = Kvaček | first2 = J. | year = 2010 | title = Late Cretaceous ''Palaeoaldrovanda'', not seeds of a carnivorous plant, but eggs of an insect | url = http://www.nm.cz/publikace/publikace-download.php?name=File1&dir=archiv&table=tabPublikaceArchiv&id=2785 | journal = Journal of the National Museum (Prague), Natural History Series | volume = 179 | issue = 9 | pages = 105–118 | access-date = 2012-10-26 | archive-url = https://web.archive.org/web/20151009191247/http://www.nm.cz/publikace/publikace-download.php?name=File1&dir=archiv&table=tabPublikaceArchiv&id=2785 | archive-date = 2015-10-09 | url-status = dead }}</ref><ref>{{cite journal | last1 = Brittnacher | first1 = J | year = 2013 | title = Phylogeny and biogeography of the Sarraceniaceae | journal = [[Carnivorous Plant Newsletter]] | volume = 42 | issue = 3| pages = 99–106 | doi = 10.55360/cpn423.jb578 | s2cid = 247114275 | doi-access = free }}</ref> The fossils more probably represent leaves (needles) of the [[Conifer|coniferous]] ''Liaoningocladus boii'' deformed by insect [[Gall|galls]].<ref name="Wong-2015" />
 ==Etymology==
-The [[Name of a biological genus|generic name]] ''Archaeamphora'' is derived from the [[Greek language|Greek]] [[wikt:αρχαίος|αρχαίος]], archaíos ("ancient"; combining form in [[Latin]]: ''archae-''), and  [[wikt:ἀμφορεύς|ἀμφορεύς]], amphoreus ("pitcher"). The [[specific name (botany)|specific epithet]] ''longicervia'' is derived from the [[Latin]] ''longus'' ("long") and ''cervicarius'' ("with a neck"), in reference to the characteristic constriction in the pitcher-like structures of this species.<ref name=Li />
+The [[Name of a biological genus|generic name]] ''Archaeamphora'' is derived from the [[Greek language|Greek]] [[wikt:αρχαίος|αρχαίος]], {{transl|el|archaíos}} ("ancient"; combining form in [[Latin]]: {{lang|la|archae}}-), and  [[wikt:ἀμφορεύς|ἀμφορεύς]], {{transl|el|amphoreus}} ("pitcher"). The [[specific name (botany)|specific epithet]] ''longicervia'' is derived from the [[Latin]] {{lang|la|longus}} ("long") and {{lang|la|cervicarius}} ("with a neck"), in reference to the characteristic constriction in the pitcher-like structures of this species.<ref name=Li />
 ==Fossil material==
-All known fossil material of ''A. longicervia'' originates from the Jianshangou Formation in [[Beipiao]], western [[Liaoning]], [[China]]. These [[Early Cretaceous]] beds constitute the lower part of the [[Yixian Formation]],<ref>Sun, G., S.-L. Zheng, D.L. Dilcher, Y.D. Wang & S.W Mei 2001. ''Early Angiosperms and their Associated Plants from Western Liaoning, China''. Shanghai Scientific and Technological Education Publishing House, 227 pp.</ref><ref>{{cite journal|last1=Sun |first1=G. |last2=Dilcher |first2=D.L. |last3=Zheng |first3=S.-L. |last4=Zhou |first4=Z.-K. |year=1998 |title=In Search of the First Flower: A Jurassic Angiosperm, ''Archaefructus'', from Northeast China |url=http://ephedra.hip.uni-heidelberg.de/lehre/ZV-Literatur.pdf |journal=Science |volume=282 |issue= 5394|pages=1692–1695 |doi=10.1126/science.282.5394.1692 |pmid=9831557 |url-status=dead |archiveurl=https://web.archive.org/web/20110719103627/http://ephedra.hip.uni-heidelberg.de/lehre/ZV-Literatur.pdf |archivedate=2011-07-19 }}</ref> which is dated at 124.6 million years old.<ref>{{cite journal | last1 = Swisher | first1 = III | last2 = Wang | first2 = Y-Q. | last3 = Wang | first3 = X.-L. | last4 = Xu | first4 = X. | last5 = Wang | first5 = Y. | year = 1999 | title = Cretaceous age for the feathered dinosaurs of Liaoning, China | doi = 10.1038/21872| journal = Nature | volume = 400 | issue = 6739| pages = 58–61 }}</ref> Nine specimens of ''A. longicervia'' have been found, including [[holotype]] CBO0220 and [[paratype]] CBO0754.<ref name=Li />
+All known fossil material of ''A. longicervia'' originates from the Jianshangou Formation in [[Beipiao]], western [[Liaoning]], [[China]]. These [[Early Cretaceous]] beds constitute the lower part of the [[Yixian Formation]],<ref>Sun, G., S.-L. Zheng, D.L. Dilcher, Y.D. Wang & S.W Mei 2001. ''Early Angiosperms and their Associated Plants from Western Liaoning, China''. Shanghai Scientific and Technological Education Publishing House, 227 pp.</ref><ref>{{cite journal|last1=Sun |first1=G. |last2=Dilcher |first2=D.L. |last3=Zheng |first3=S.-L. |last4=Zhou |first4=Z.-K. |year=1998 |title=In Search of the First Flower: A Jurassic Angiosperm, ''Archaefructus'', from Northeast China |url=http://ephedra.hip.uni-heidelberg.de/lehre/ZV-Literatur.pdf |journal=Science |volume=282 |issue= 5394|pages=1692–1695 |doi=10.1126/science.282.5394.1692 |pmid=9831557 |url-status=dead |archive-url=https://web.archive.org/web/20110719103627/http://ephedra.hip.uni-heidelberg.de/lehre/ZV-Literatur.pdf |archive-date=2011-07-19 }}</ref> which is dated at 124.6 million years old.<ref>{{cite journal | last1 = Swisher | first1 = III | last2 = Wang | first2 = Y-Q. | last3 = Wang | first3 = X.-L. | last4 = Xu | first4 = X. | last5 = Wang | first5 = Y. | year = 1999 | title = Cretaceous age for the feathered dinosaurs of Liaoning, China | doi = 10.1038/21872| journal = Nature | volume = 400 | issue = 6739| pages = 58–61 | bibcode = 1999Natur.400...58S | s2cid = 4424629 | url = http://doc.rero.ch/record/14906/files/PAL_E2048.pdf }}</ref> Nine specimens of ''A. longicervia'' have been found, including [[holotype]] CBO0220 and [[paratype]] CBO0754.<ref name=Li />
 ==Description==
-''Archaeamphora longicervia'' was a [[herbaceous]] plant growing to around {{convert|50|mm|in|abbr=on}} in height. The stem, at least {{convert|21|mm|abbr=on}} long by {{convert|1.2|mm|in|abbr=on}} wide, bore distinctive vertical ridges and grooves. The pitcher-like structures were [[wikt:ascidiate|ascidiate]] in form and {{convert|30|to|40|mm|in|abbr=on}} long. Mature pitchers and underdeveloped pitchers or [[Sarracenia#Growth cycle|phyllodia]]-like leaves were arranged spirally around the stem. Pitchers consisted of a tubular base, expanded middle section, constriction around the mouth, and a vertical, spoon-shaped lid. A single wing ran down the [[wikt:adaxial|adaxial]] side of each pitcher. Three to five parallel major veins were present on the pitchers, along with a few intercostal veins and numerous small veinlets.<ref name=Li />
+''Archaeamphora longicervia'' was supposed to be a [[herbaceous]] plant growing to around {{convert|50|mm|in|abbr=on}} in height. The stem, at least {{convert|21|mm|abbr=on}} long by {{convert|1.2|mm|in|abbr=on}} wide, bore distinctive vertical ridges and grooves. The pitcher-like structures were [[wikt:ascidiate|ascidiate]] in form and {{convert|30|to|40|mm|in|abbr=on}} long. Mature pitchers and underdeveloped pitchers or [[Sarracenia#Growth cycle|phyllodia]]-like leaves were arranged spirally around the stem. Pitchers consisted of a tubular base, expanded middle section, constriction around the mouth, and a vertical, spoon-shaped lid. A single wing ran down the [[wikt:adaxial|adaxial]] side of each pitcher. Three to five parallel major veins were present on the pitchers, along with a few intercostal veins and numerous small veinlets.<ref name=Li />
 Two unusual bag-like structures were present on each pitcher, one on either side of the central wing. Similar but semi-circular structures were found on the margin of the lid. These structures exhibited strong yellow-green intrinsic [[fluorescence]] when exposed to [[visible light]] with a [[wavelength]] of 500 [[nanometre|nm]] ([[Visible spectrum|blue-green]]).<ref name=Li />
@@ Line 39: / Line 36: @@
 ==Taxonomy==
-The fossil material of ''A. longicervia'' was subjected to chemical analysis for [[oleanane]], considered a key marker differentiating [[angiosperm]]s from [[gymnosperm]]s.<ref>{{cite journal | last1 = Moldowan | first1 = J.M. | last2 = Dahl | first2 = J. | last3 = Huizinga | first3 = B.J. | last4 = Fago | first4 = F.J. | last5 = Hickey | first5 = L.J. | last6 = Peakman | first6 = T.M. | last7 = Taylor | first7 = O.W. | year = 1994 | title = The molecular fossil record of oleanane and its relationship to Angiosperms | url = http://www.sciencemag.org/cgi/content/abstract/265/5173/768 | journal = Science | volume = 265 | issue = 5173| pages = 768–771 | doi=10.1126/science.265.5173.768| pmid = 17736275 }}</ref> Oleanane was detected in these specimens, suggesting that they belong to the angiosperms.<ref name=Li />
+The fossil material of ''A. longicervia'' was subjected to chemical analysis for [[oleanane]], considered a key marker differentiating [[angiosperm]]s from [[gymnosperm]]s.<ref>{{cite journal | last1 = Moldowan | first1 = J.M. | last2 = Dahl | first2 = J. | last3 = Huizinga | first3 = B.J. | last4 = Fago | first4 = F.J. | last5 = Hickey | first5 = L.J. | last6 = Peakman | first6 = T.M. | last7 = Taylor | first7 = O.W. | year = 1994 | title = The molecular fossil record of oleanane and its relationship to Angiosperms | url = http://www.sciencemag.org/cgi/content/abstract/265/5173/768 | journal = Science | volume = 265 | issue = 5173| pages = 768–771 | doi=10.1126/science.265.5173.768| pmid = 17736275 | s2cid = 9389874 }}</ref> Oleanane was detected in these specimens, suggesting that they belong to the angiosperms.<ref name=Li />
 ===Pitcher plant interpretation===
+[[File:Archaeamphora longicervia.png|thumb|Artist's reconstruction of ''Archaeamphora longicervia'' as pitcher plant, after Li (2005)<ref name="Li" />]]
 According to Li (2005), several morphological features of ''A. longicervia'' indicate a close relationship to [[Sarraceniaceae]]: both taxa exhibit one or two pitcher wings, a smooth [[peristome]], and pitchers that extend vertically from the top of a short [[petiole (botany)|petiole]].<ref name=Li />
 [[File:Sarracenia purpurea0.jpg|thumb|Pitchers of ''[[Sarracenia purpurea]]'']]
-Li (2005) suggests that ''A. longicervia'' is morphologically similar to modern ''[[Sarracenia purpurea]]''. It shares with this species the spiral arrangement of its pitchers and [[Sarracenia#Growth cycle|phyllodia]]-like tubular leaves with parallel major veins. ''Archaeamphora longicervia'' also shows a resemblance to species of the genus ''[[Heliamphora]]'' in having pitchers with a long neck and upright lid. Of particular note is the similarity between the thick semi-circular structures on the lid of ''A. longicervia'' and the large [[nectar]]-secreting "bubble" present on the upper posterior portion of ''[[Heliamphora exappendiculata]]''{{refn|group=nb|name=exa|This taxon was recognised as a [[variety (botany)|variety]] of ''[[Heliamphora heterodoxa]]'' when ''Archaeamphora'' was described in 2005.<ref name=Li /> It was raised to species rank the following year.<ref>{{cite journal | last1 = Nerz | first1 = J. | last2 = Wistuba | first2 = A. | year = 2006 | title = ''Heliamphora exappendiculata'', a clearly distinct species with unique characteristics | url = http://www.carnivorousplants.org/cpn/Species/v35n2p43_48.html | journal = [[Carnivorous Plant Newsletter]] | volume = 35 | issue = 2| pages = 43–51 }}</ref>}} pitchers.<ref name=Li />
+Li (2005) suggests that ''A. longicervia'' is morphologically similar to modern ''[[Sarracenia purpurea]]''. It shares with this species the spiral arrangement of its pitchers and [[Sarracenia#Growth cycle|phyllodia]]-like tubular leaves with parallel major veins. ''Archaeamphora longicervia'' also shows a resemblance to species of the genus ''[[Heliamphora]]'' in having pitchers with a long neck and upright lid. Of particular note is the similarity between the thick semi-circular structures on the lid of ''A. longicervia'' and the large [[nectar]]-secreting "bubble" present on the upper posterior portion of ''[[Heliamphora exappendiculata]]''{{refn|group=nb|name=exa|This taxon was recognised as a [[variety (botany)|variety]] of ''[[Heliamphora heterodoxa]]'' when ''Archaeamphora'' was described in 2005.<ref name=Li /> It was raised to species rank the following year.<ref>{{cite journal | last1 = Nerz | first1 = J. | last2 = Wistuba | first2 = A. | year = 2006 | title = ''Heliamphora exappendiculata'', a clearly distinct species with unique characteristics | url = http://www.carnivorousplants.org/cpn/Species/v35n2p43_48.html | journal = [[Carnivorous Plant Newsletter]] | volume = 35 | issue = 2| pages = 43–51 | doi = 10.55360/cpn352.jn909 | s2cid = 247149195 | doi-access = free }}</ref>}} pitchers.<ref name=Li />
 Li (2005) mentions the discovery of another type of "pitcher plant" from the same formation. This variety differs from the type material of ''A. longicervia'' in having pitchers that lack any constriction before the mouth, instead gradually expanding from the [[petiole (botany)|petiole]] into a hollow trumpet-like shape. He suggests that it "should be a different species" from ''A. longicervia''. An intermediate form with a wider neck is also reported, suggesting that these plants were already a diversified group in the Early Cretaceous.<ref name=Li />
@@ Line 57: / Line 55: @@
 <blockquote>Serious doubt is emerging that reduces the likelihood that ''Archaeamphora longicervia'' belongs in the Sarraceniaceae lineage, or was even a pitcher plant at all. [...] Although ''Archaeamphora'' might well be a representative of the earliest flowering plants on Earth [...] it is very unlikely that it represents an ancestor of Sarraceniaceae since it is much too old to be part of the advanced "[[crown group]]" of [[Ericales]] to which Sarraceniaceae belong. [...] Another contradiction is that except for ''Archaeamphora'', there is no other evidence to suggest that Sarraceniaceae evolved outside the [[New World]], to which all extant members of the family are endemic.</blockquote>
+Wong et al. (2015) put forward a new perspective as follows:<ref name="Wong-2015">{{cite journal | last1 = Wong | first1 = WO | last2 = Dilcher | first2 = DL | last3 = Labandeira | first3 = CC | last4 = Sun | first4 = G | last5 = Fleischmann | first5 = A | year = 2015 | title = Early Cretaceous ''Archaeamphora'' is not a carnivorous angiosperm | doi = 10.3389/fpls.2015.00326 | pmid = 25999978 | pmc = 4423337 | journal = Frontiers in Plant Science | volume = 6 | page = 326 | doi-access = free }}</ref>
-Wong et al. (2015) put forward a new perspective [16] as follows:
 <blockquote>''Archaeamphora longicervia'' H. Q. Li was described as an herbaceous, Sarraceniaceae-like pitcher plant from the mid Early Cretaceous Yixian Formation of Liaoning Province, northeastern China. Here, a re-investigation of ''A. longicervia'' specimens from the Yixian Formation provides new insights into its identity and the morphology of pitcher plants claimed by Li. We demonstrate that putative pitchers of ''Archaeamphora'' are insect-induced leaf galls that consist of three components: (1) an innermost larval chamber; (2) an intermediate zone of nutritive tissue; and (3) an outermost wall of sclerenchyma. ''Archaeamphora'' is not a carnivorous, Sarraceniaceae-like angiosperm, but represents insect-galled leaves of the previously reported gymnosperm ''Liaoningocladus boii'' G. Sun et al. from the Yixian Formation.</blockquote>
 ==Habitat==
-The area inhabited by ''A. longicervia'' is thought to have experienced significant climatic fluctuations during the Early Cretaceous, ranging from arid or semi-arid to more humid conditions.<ref>{{cite journal | last1 = Barrett | first1 = P.M. | last2 = Hilton | first2 = J. M. | year = 2006 | title = The Jehol Biota (Lower Cretaceous, China): new discoveries and future prospects | journal = Integrative Zoology | volume = 1 | issue = 1| pages = 15–17 | doi=10.1111/j.1749-4877.2006.00006.x| pmid = 21395985 }}</ref> The [[Substrate (biology)|substrate]] in the region was mostly composed of [[Lake|lacustrine]] sediments and [[volcanic rock]]s.<ref>{{cite journal | last1 = Tan | first1 = J.-J. | last2 = Ren | first2 = D. | year = 2006 | title = ''Ovatucupes'': A New Cupedid Genus (Coleoptera: Archostemata: Cupedidae) From The Jehol Biota (Late Jurassic) Of Western Liaoning, China | url = http://fossilinsects.net/pdfs/Tan_Ren_2006.pdf | journal = Entomological News | volume = 117 | issue = 2| pages = 223–232 | doi=10.3157/0013-872x(2006)117[223:oancgc]2.0.co;2}}</ref>
+The area inhabited by ''A. longicervia'' is thought to have experienced significant climatic fluctuations during the Early Cretaceous, ranging from arid or semi-arid to more humid conditions.<ref>{{cite journal | last1 = Barrett | first1 = P.M. | last2 = Hilton | first2 = J. M. | year = 2006 | title = The Jehol Biota (Lower Cretaceous, China): new discoveries and future prospects | journal = Integrative Zoology | volume = 1 | issue = 1| pages = 15–17 | doi=10.1111/j.1749-4877.2006.00006.x| pmid = 21395985 | doi-access = free }}</ref> The [[Substrate (biology)|substrate]] in the region was mostly composed of [[Lake|lacustrine]] sediments and [[volcanic rock]]s.<ref>{{cite journal | last1 = Tan | first1 = J.-J. | last2 = Ren | first2 = D. | year = 2006 | title = ''Ovatucupes'': A New Cupedid Genus (Coleoptera: Archostemata: Cupedidae) From The Jehol Biota (Late Jurassic) Of Western Liaoning, China | url = http://fossilinsects.net/pdfs/Tan_Ren_2006.pdf | journal = Entomological News | volume = 117 | issue = 2| pages = 223–232 | doi=10.3157/0013-872x(2006)117[223:oancgc]2.0.co;2| s2cid = 84331871 }}</ref>
 ==See also==
@@ Line 68: / Line 66: @@
 ==Notes==
-{{reflist|group=nb}}
+{{Reflist|group=nb}}
 ==References==
-{{reflist|40em}}
+{{Reflist|40em}}
-. {{cite journal | last1 = Wong | first1 = WO | last2 = Dilcher | first2 = DL | last3 = Labandeira | first3 = CC | last4 = Sun | first4 = G | last5 = Fleischmann | first5 = A | year = 2015 | title = Early Cretaceous ''Archaeamphora'' is not a carnivorous angiosperm | doi = 10.3389/fpls.2015.00326 | pmid = 25999978 | pmc = 4423337 | journal = Frontiers in Plant Science | volume = 6 | issue = | page = 326 }}
-{{CarnivorousPlants}}
+{{Carnivorous plants}}
-{{Taxonbar|from=Q144107}}
+{{Taxonbar|from=Q144107|from2=Q27664062}}
 [[Category:Sarraceniaceae]]
-[[Category:Prehistoric angiosperm genera]]
+[[Category:Prehistoric gymnosperm genera]]
 [[Category:Early Cretaceous plants]]
 [[Category:Extinct carnivorous plants]]
 [[Category:Cretaceous angiosperms]]
 [[Category:Fossil taxa described in 2005]]
+[[Category:Pinales]]
+[[Category:Controversial plant taxa]]

v t e Carnivorous and protocarnivorous plants
Carnivorous genera († extinct)	Aldrovanda †Archaeamphora? Brocchinia Byblis Catopsis Cephalotus Darlingtonia Dionaea Drosera †Droserapites †Droserapollis †Droseridites Drosophyllum †Fischeripollis Genlisea Heliamphora Nepenthes †Palaeoaldrovanda? Philcoxia Pinguicula Roridula Sarracenia †Saxonipollis Triantha Triphyophyllum Utricularia
Protocarnivorous genera	Aracamunia Capsella Colura Dipsacus Drymocallis Geranium Ibicella Lathraea Paepalanthus Passiflora Plumbago Proboscidea Stylidium
See also	International Carnivorous Plant Society Insectivorous Plant Society North American Sarracenia Conservancy List of carnivorous plants List of carnivorous plant periodicals Pitcher plant