Taxaceae
{{Short description|Family of conifers}}
{{Automatic taxobox
| fossil_range = {{fossil range|Early Jurassic|Present|earliest=Late Triassic}} (possible Late Triassic records)
| image = Yew 1r.jpg
| image_caption = Foliage and mature arils of a yew plant
| taxon = Taxaceae
| authority = S.F.Gray
| subdivision_ranks = Genera
| subdivision = *Amentotaxus
- Austrotaxus
- Cephalotaxus
- {{extinct}}Diploporus
- Pseudotaxus
- Taxus
- Torreya
- Taxaceoxylon (fossil wood)
| synonyms = * Austrotaxaceae Neger 1907
- Cephalotaxaceae Kudo & Yamamoto 1931
}}
Taxaceae ({{IPAc-en|t|æ|k|ˈ|s|eɪ|s|i|.|iː|,_|-|ˌ|aɪ}}), commonly called the yew family, is a coniferous family which includes six extant and two extinct genera, and about 30 species of plants, or in older interpretations three genera and 7 to 12 species.
Description
They are many-branched, small trees and shrubs. The leaves are evergreen, spirally arranged, often twisted at the base to appear 2-ranked. They are linear to lanceolate, and have pale green or white stomatal bands on the undersides.
The plants are dioecious, or rarely monoecious. The catkin like male cones are {{convert|2|–|5|mm|in}} long, and shed pollen in the early spring. They are sometimes externally only slightly differentiated from the branches. The fertile bracts have 2-8 pollen sacs.{{Cite book |last=Phillips |first=Edwin Percy |title=The genera of South African flowering plants |publisher=Government Printer |year=1951 |publication-place=South Africa}}
The female 'cones' are highly reduced. Only the upper or uppermost bracts are fertile and bear one or rarely two seeds. The ovule usually exceeds the scale, although ovules are sometimes rarely enclosed by it. They may be found on the ends of branches or on the branches. They may grow singly or in tufts or clumps.
As the seed matures, a fleshy aril partly encloses it. The developmental origin of the aril is unclear, but it may represent a fused pair of swollen leaves.{{cite journal | last1=Dörken | first1=Veit Martin | last2=Nimsch | first2=Hubertus | last3=Rudall | first3=Paula J | title=Origin of the Taxaceae aril: evolutionary implications of seed-cone teratologies in Pseudotaxus chienii | journal=Annals of Botany | publisher=Oxford University Press (OUP) | volume=123 | issue=1 | date=2018-08-22 | issn=0305-7364 | doi=10.1093/aob/mcy150 | pmid=30137225 | pmc=6344100 | pages=133–143}} The mature aril is brightly coloured, soft, juicy and sweet, and is eaten by birds which then disperse the hard seed undamaged in their droppings. However, the seeds are highly poisonous to humans, containing the poisons taxine and taxol.[https://www.nlm.nih.gov/medlineplus/ency/article/002877.htm Yew Poisoning: MedLine Plus Medical Encyclopedia]
Distribution
Classification
Taxaceae is now generally included with all other conifers in the order Pinales, as DNA analysis has shown that the yews are phylogenetically nested in the Pinales,{{cite journal |last1=Chase |first1=M. W. |last2=Soltis |first2=D. E. |last3=Olmstead |first3=R. G. |last4=Morgan |first4=D. |last5=Les |first5=D. H. |last6=Mishler |first6=B. D. |last7=Duvall |first7=M. R. |last8=Price |first8=R. A. |last9=Hills |first9=H. G. |last10=Qiu |first10=Y.-L. |last11=Kron |first11=K. A. |last12=Rettig |first12=J. H. |last13=Conti |first13=E. |last14=Palmer |first14=J. D. | last15=Manhart | first15=J. R. | last16=Sytsma | first16=K. J. | last17=Michaels | first17=H. J. | last18=Kress | first18=W. J. | last19=Karol | first19=K. G. | last20=Clark | first20=W. D. | last21=Hedren | first21=M. | last22=Gaut | first22=B. S. | last23=Jansen | first23=R. K. |last24=Kim | first24=K.-J. |last25=Wimpee | first25=C. F. |last26=Smith |first26=J. F. | last27=Furnier | first27=G. R. | last28=Strauss | first28=S. H. | last29=Xiang | first29=Q.-Y. | last30=Plunkett | first30=G. M. |last31=Soltis | first31=P. S. | last32=Swensen | first32=S. M. | last33=Williams | first33=S. E. | last34=Gadek | first34=P. A. | last35=Quinn | first35=C. J. | last36=Eguiarte | first36=L. E. | last37=Golenberg | first37=E. |last38=Learn | first38=G. H. | last39=Graham | first39=S. W. | last40=Barrett | first40=S. C. H. | last41=Dayanandan | first41=S. | last42=Albert | first42=V. A. |display-authors=2| title=Phylogenetics of Seed Plants: An Analysis of Nucleotide Sequences from the Plastid Gene rbcL | journal=Annals of the Missouri Botanical Garden | publisher=JSTOR | volume=80 | issue=3 | year=1993 | issn=0026-6493 | doi=10.2307/2399846 | page=528|jstor=2399846 |url=https://spectrum.library.concordia.ca/6741/1/Dayanandan_AnnalsMissouriBotanicalGardens_1993.pdf | hdl=1969.1/179875 | hdl-access=free }} a conclusion supported by micromorphology studies.{{cite journal|last1=Anderson|first1=E.|last2=Owens|first2=J.N.|year=2003|title=Analyzing the Reproductive Biology of Taxus: Should It be Included in Coniferales?|url=https://www.actahort.org/books/615/615_22.htm|journal=Acta Horticulturae|publisher=International Society for Horticultural Science (ISHS)|issue=615|pages=233–234|doi=10.17660/actahortic.2003.615.22|issn=0567-7572|access-date=2021-06-22|archive-date=2022-11-26|archive-url=https://web.archive.org/web/20221126082249/https://www.actahort.org/books/615/615_22.htm|url-status=dead|url-access=subscription}} Formerly they were often treated as distinct from other conifers by placing them in a separate order Taxales. Ernest Henry Wilson referred to Taxaceae as "taxads" in his 1916 book.{{cite book | last=Wilson | first=Ernest Henry | title=The conifers and taxads of Japan. Issued December 30, 1916. | url=https://archive.org/details/coniferstaxadsof00wilsuoft | publisher=University Press | location=Cambridge | year=1916 | doi=10.5962/bhl.title.17457 }} Taxaceae is thought to be the sister group to Cupressaceae, from which it diverged during the early-mid Triassic. The clade comprising both is sister to Sciadopityaceae, which diverged from them during the early-mid Permian.{{Cite journal|last1=Stull|first1=Gregory W.|last2=Qu|first2=Xiao-Jian|last3=Parins-Fukuchi|first3=Caroline|last4=Yang|first4=Ying-Ying|last5=Yang|first5=Jun-Bo|last6=Yang|first6=Zhi-Yun|last7=Hu|first7=Yi|last8=Ma|first8=Hong|last9=Soltis|first9=Pamela S.|last10=Soltis|first10=Douglas E.|last11=Li|first11=De-Zhu|date=July 19, 2021|title=Gene duplications and phylogenomic conflict underlie major pulses of phenotypic evolution in gymnosperms|url=https://www.nature.com/articles/s41477-021-00964-4|journal=Nature Plants|language=en|volume=7|issue=8|pages=1015–1025|doi=10.1038/s41477-021-00964-4|pmid=34282286 |bibcode=2021NatPl...7.1015S |s2cid=236141481 |issn=2055-0278}} The oldest confirmed member of Taxaceae is Palaeotaxus rediviva from the earliest Jurassic (Hettangian) of Sweden. Fossils belonging to the living genus Amentotaxus from the Middle Jurassic of China indicate that Taxaceae had already substantially diversified during the Jurassic.{{Cite journal |last1=Dong |first1=Chong |last2=Shi |first2=Gongle |last3=Herrera |first3=Fabiany |last4=Wang |first4=Yongdong |last5=Herendeen |first5=Patrick S |last6=Crane |first6=Peter R |date=2020-06-18 |title=Middle–Late Jurassic fossils from northeastern China reveal morphological stasis in the catkin-yew |journal=National Science Review |language=en |volume=7 |issue=11 |pages=1765–1767 |doi=10.1093/nsr/nwaa138 |issn=2095-5138 |pmc=8288717 |pmid=34691509 |doi-access=free}}
The broadly defined Taxaceae (including Cephalotaxus) comprises six extant genera and about 30 species overall. Cephalotaxus is now included in Taxaceae, rather than being recognized as the core of its own family, Cephalotaxaceae. Phylogenetic evidence strongly supports a very close relationship between Cephalotaxus and other members of Taxaceae,{{cite journal | last1=Quinn | first1=C. J. | last2=Price | first2=R. A. | last3=Gadek | first3=P. A. | title=Familial Concepts and Relationships in the Conifer Based on rbcL and matK Sequence Comparisons | journal=Kew Bulletin | publisher=JSTOR | volume=57 | issue=3 | year=2002 | issn=0075-5974 | doi=10.2307/4110984 | page=513| jstor=4110984 | bibcode=2002KewBu..57..513Q | s2cid=83816639 }}{{cite journal | last1=Rai | first1=Hardeep S. | last2=Reeves | first2=Patrick A. | last3=Peakall | first3=Rod | last4=Olmstead | first4=Richard G. | last5=Graham | first5=Sean W. | title=Inference of higher-order conifer relationships from a multi-locus plastid data set. | journal=Botany | publisher=Canadian Science Publishing | volume=86 | issue=7 | year=2008 | issn=1916-2790 | doi=10.1139/b08-062 | pages=658–669| s2cid=14007221 }}{{cite journal | title=One thousand plant transcriptomes and the phylogenomics of green plants | journal=Nature | publisher=Springer Science and Business Media LLC | volume=574 | issue=7780 | year=2019 | issn=0028-0836 | doi=10.1038/s41586-019-1693-2 | pmid=31645766 | pmc=6872490 | pages=679–685| author1=One Thousand Plant Transcriptomes Initiative }} and morphological differences between them are not substantial. Previous recognition of two distinct families, Taxaceae and Cephalotaxaceae (e.g.,{{Cite journal|last=Hart|first=Jeffrey A.|date=1987|title=A cladistic analysis of conifers: preliminary results|journal=Journal of the Arnold Arboretum|volume=68|issue=3|pages=269–307|doi=10.5962/p.185944 |jstor=43782212|s2cid=88860959 |issn=0004-2625|doi-access=free}}), was based on relatively minor morphological details: Taxaceae (excluding Cephalotaxus) has smaller mature seeds growing to {{convert|5|-|8|mm|in}} in 6–8 months, that are not fully enclosed by the aril; in contrast, Cephalotaxus seeds have a longer maturation period (from 18–20 months), and larger mature seeds ({{convert|12|-|40|mm|in}}) fully enclosed by the aril. However, there are also very clear morphological connections between Cephalotaxus and other members of Taxaceae,{{cite journal | last=Doyle | first=James A. | journal=Annual Review of Ecology and Systematics | publisher=Annual Reviews | volume=29 | issue=1 | year=1998 | issn=0066-4162 | doi=10.1146/annurev.ecolsys.29.1.567 | pages=567–599| title=Phylogeny of Vascular Plants | s2cid=85631751 }}{{cite journal|last1=Stützel|first1=Thomas|last2=Röwekamp|first2=Iris|year=1999|title=Female reproductive structures in Taxales|url=https://linkinghub.elsevier.com/retrieve/pii/S0367253017308939|journal=Flora|publisher=Elsevier BV|volume=194|issue=2|pages=145–157|doi=10.1016/s0367-2530(17)30893-9|bibcode=1999FMDFE.194..145S |issn=0367-2530|url-access=subscription}} and considered in tandem with the phylogenetic evidence, there is no compelling need to recognize Cephalotaxus (or other genera in Taxaceae) as a distinct family.
=Phylogeny=
Phylogeny of Taxaceae.{{cite journal |last1=Leslie |first1=Andrew B. |last2=Beaulieu |first2=Jeremy |last3=Holman |first3=Garth |last4=Campbell |first4=Christopher S. |last5=Mei |first5=Wenbin |last6=Raubeson |first6=Linda R. |last7=Mathews |first7=Sarah |display-authors=et al. |year=2018 |title=An overview of extant conifer evolution from the perspective of the fossil record |journal=American Journal of Botany |volume=105 |issue=9 |pages=1531–1544 | doi=10.1002/ajb2.1143 |pmid= 30157290|pmc= |bibcode= |doi-access=free}}{{cite journal |last1=Leslie |first1=Andrew B. |display-authors=et al. |year=2018 |title=ajb21143-sup-0004-AppendixS4 |journal=American Journal of Botany |volume=105 |issue=9 |pages=1531–1544 |doi=10.1002/ajb2.1143 |pmid=30157290 |s2cid=52120430 |url=https://bsapubs.onlinelibrary.wiley.com/action/downloadSupplement?doi=10.1002%2Fajb2.1143&file=ajb21143-sup-0004-AppendixS4.pdf |doi-access=}}
{{clade|style=font-size:100%;line-height:100%
|1={{Clade
|label1=Cephalotaxeae
|1=Cephalotaxus
|label2=Taxoideae
|2={{Clade
|1={{Clade
|1=Amentotaxus
|2=Torreya
}}
|2={{Clade
|1=Austrotaxus
|2={{Clade
|1=Pseudotaxus
|2=Taxus
}}
}}
}}
}}
}}
File:Cephalotaxus sinensis 1.jpg]]
File:Taxus brevifolia Blue Mts WA.jpg]]
{{div col}}
Amentotaxus {{small|Pilg.}} – Catkin-yew
- Amentotaxus argotaenia - Catkin yew
- Amentotaxus assamica - Assam catkin yew
- Amentotaxus formosana - Taiwan catkin yew
- Amentotaxus poilanei - Poilane's catkin yew
- Amentotaxus yunnanensis - Yunnan catkin yew
Austrotaxus {{small|Compton}} – New Caledonia yew
- Austrotaxus spicata - New Caledonia yew or southern yew
Cephalotaxus {{small|Siebold & Zucc. ex Endl.}} – Plum yew
- Cephalotaxus fortunei - Chinese plum-yew
- Cephalotaxus griffithii - Griffith's plum yew
- Cephalotaxus hainanensis - Hainan plum-yew
- Cephalotaxus harringtonii - Korean plum yew, Japanese plum-yew
- Cephalotaxus koreana - Korean plum yew
- Cephalotaxus lanceolata - Gongshan plum yew
- Cephalotaxus latifolia - Broad-leaved plum yew
- Cephalotaxus mannii - Mann's yew plum
- Cephalotaxus oliveri - Oliver's plum yew
- Cephalotaxus sinensis - Chinese plum yew
- Cephalotaxus wilsoniana - Taiwan plum yew, Taiwan cow's-tail pine, or Wilson plum yew
Pseudotaxus {{small|W.C.Cheng}} – White-berry yew
- Pseudotaxus chienii - the whiteberry yew
Taxus {{small|L.}} – Common yew
- Taxus baccata European yew
- Taxus biternata Delicate branch yew
- Taxus brevifolia Pacific yew, western yew
- Taxus caespitosa Caespitosa yew
- Taxus calcicola Asian limestone yew
- Taxus canadensis Canada yew
- Taxus celebica Celebes yew
- Taxus chinensis China yew
- Taxus contorta West Himalayan yew
- Taxus cuspidata Rigid branch yew, Japanese yew
- Taxus fastigiata Irish yew
- Taxus floridana Florida yew
- Taxus florinii Florin yew
- Taxus globosa Mesoamerican yew
- Taxus kingstonii Kingston yew
- Taxus mairei Maire yew
- Taxus obscura Obscure yew
- Taxus ocreata Scaly yew
- Taxus phytonii Phyton yew
- Taxus recurvata English yew
- Taxus rehderiana Rehder yew
- Taxus scutata Scutaceous yew
- Taxus suffnessii Suffness yew
- Taxus sumatrana Sumatera yew
- Taxus umbraculifera Umbrelliform yew
- Taxus wallichiana Wallich yew, East Himalayan yew
Torreya {{small|Arn.}} – Nutmeg yew
- Torreya californica - California torreya
- Torreya fargesii - Farges nutmeg tree
- Torreya grandis - Chinese nutmeg yew
- Torreya jackii - Jack's nutmeg tree, longleaf torreya etc
- Torreya nucifera - kaya, Japanese torreya, or Japanese nutmeg-yew
- Torreya taxifolia - Gopher wood
- {{extinct}}Torreya clarnensis{{Cite journal|last1=Manchester|first1=S.R.|year=1994|title=Fruits and Seeds of the Middle Eocene Nut Beds Flora, Clarno Formation, Oregon|url=https://www.biodiversitylibrary.org/part/253565|journal=Palaeontographica Americana|volume=58|pages=30–31}}
{{div col end}}
Footnotes
{{Commons category|Taxaceae}}
{{Wikispecies|Taxaceae}}
{{Reflist}}
{{Pinophyta}}
{{Taxonbar|from=Q223751}}
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