Taxus brevifolia var. polychaeta

First prepared 11 Oct 2006, published in JBRIT, 2007; additional photos June 2010;
page format revised  and updated Aug 2014
 ©The World Botanical Associates Web Page


3c. Taxus brevifolia var. polychaeta Spjut—J. Bot. Res. Inst. Texas 1 (1): 217. 2007. Type: U.S.A., Washington: Thurston Co., Mud Bay, near Tacoma, F. G. Meyer 1589 (holotype: K!), photo shown in key.  Note: A combination attributed to Silba, Taxus brevifolia subsp. polychaeta  with reference to "J. Int. Conifer Preserv. Soc. 17(1): 22. 2010" (n.v.) is inconsistent with taxonomic treatment of subtaxa within the genus (Silba 1984; Spjut 2007a, Spjut 2007b).  Subspecies status in Taxus was applied to geographically differentiated populations by Pilger (1903, 1916, 1926).  Thus, Silba's proposed combination is superfluous and possibly invalid.  See also Ellison AM, Davis CC, Calie PJ, Naczi RFC; Pitcher plants (Sarracenia) provide a 21st-century perspective on infraspecific ranks and interspecific hybrids: A modest proposal for appropriate recognition and usage. Syst. Bot. 39: 939–949, 2014.

Diagnostic features: Tree with erect trunk at base, vegetative reproduction unknown; branches developing in all directions around the trunk, wide spreading to ascending or somewhat erect; ovuliferous shoots cylindrical with numerous scales that are all similar in size (belonging to the primary shoot), the entire shoot complex much longer than seed, sometimes shortly branched near base, or branched above base (Sonoma County, CA), usually with more than one ovule, two subterminal ovules on long primary shoots , 5–8 mm long, in the northern range, or as many five ovules on branched shoot complexes <5 mm long in plants from Sonoma County, CA.

Worm-cone yew: California (Mendocino Co.?, Sonoma Co.), Washington (Mud Bay near Tacoma, extirpated?).  Specimens studied: California: Sonoma Co.: 7 km E of Stewarts Point, redwood-grand fir forest, 200 m, Rich Spjut & Rick Spjut 16021 (wba). Bigelow (Marin, Sonoma or Mendocino Co.), yr 1854 (US); Salmon Creek (Sonoma or Mendocino Co.?), McMurphy 315 (US).  Idaho, near Coeur d'Alene, shipped by Marion Blatch as fresh material via overnight mail from Coeur d'Alene Nursery, 26 June 1992, possibly collected in nearby Washington; collector, locality, and habit of plant unknown, supplied by the USDA Forest Service without data.  A report by Spjut (1977) noted a collection from “Coeur d'Alene National Forest: T51N R8W Sec. 25; 3,800 ft, on north slope near stream” (n.v.).  Additional occurrence of the short shoot form found Aug 2011 in the Klamath Mountains of Southern Oregon along Thompson Creek.

     Taxus brevifolia var. polychaeta is distinguished from the other varieties of the species by long cylindrical fertile shoots that resemble annelid or polychaete worms, hence, the epithet, polychaeta (Spjut 2007b).  The ovule bearing shoots (cones) of var. polychaeta have been distinguished by their development of multiple ovules (this web page, June 2010); however, vars. brevifolia and reptaneta have since been found to have multi-ovulate shoot complexes, but they differ in the shape of the shoot, cylindrical in var. polychaeta, funnelform in the other varieties. The ovules may appear terminal on a ovular shoot complex (var. brevifolia, polychaeta, reptaneta), or in var. polychaeta the secondary shoot may further grow from an aborted development of one of the ovules, leaving the lower ovular region to appear lateral with one or more terminal ovules, or as others have suggested, multiple secondary shoots may arise from a primary shoot over a period of several seasons (Anderson & Owens 1999; Anderson 2001).  It has not been determined whether the primary shoot in var. polychaeta falls with the seed or persists.

     The idea of Taxus having just one terminal ovule on a shoot that arises from the axis of a leaf has generally prevailed in the literature on Taxus reproductive morphology, and because there is no clear homologue to the seed-scale-bract complex in conifers (e.g., Florin 1948, 1951; Cope 1998), Taxus is also stated to lack cones; however, the definition of cone given by Clement-Westerhof (1988) includes the “ovuliferous dwarf-shoot” as it relates to extinct conifers, and further defines bract as “a foliar appendage of the cone axis.”   It should be recognized that the ovular shoot complex of Taxus consists of a primary shoot that arises in the axil of leaf and a secondary shoot that arises in the axil of a scale.  The interpretation of whether cones are present or absent in Taxus may be a question of whether it is the subtending scale of the secondary shoot, or the leaf that subtends the primary shoot is the bract homologue (Miller 1988).  Examples of various multi-ovulate shoots of Taxus are presented in an overview of the genus Taxus.

     Taxus brevifolia var. polychaeta apparently occurs with the typical variety (var. brevifolia), judging from other specimens—collected by Fred Meyer reportedly at the type locality under the same number (e.g., specimen at US)—that have typical cones, and also by Peter Baye in Sonoma County.  While Spjut (2007b) had  earlier questioned whether var. polychaeta  has biological 'species' significance (Spjut 2007b), he also noted that unique genotypes in T. brevifolia have been recognized in relationship to other geographically differentiated genotypic populations (El- Kassaby & Yanchuk 1994), but without consideration to morphological and reproductive differences (Spjut 2007b).  Two varieties of T. mairei, var. mairei and var. speciosa, which are sympatric throughout most of their range in southern China, are distinguished by their branching patterns and by their seed shape and color (Spjut 2007b), and seem to correspond to two haplotypes that occur together throughout their range (Gao et al. 2007). Taxus brevifolia var. brevifolia also occurs with var. T. brevifolia var. klamathensis (Spjut ineditus, abstract submitted, var. nov.).  It may be noted that Möller et al (2013) do not accept taxonomic differences in Taxus unless they are geographically distinct, possibly because disjunct populations of a particular species or variety of Taxus may be associated with different species of Taxus at the different locations, in which there are likely to be genetic differences resulting from historical hybridization and introgression.

     The occurrence of var. polychaeta along the California coastal region is rare (Baye 2004, ltr to Calif. Dept. Forestry; Spjut 2007).  One record cited for Sonoma County by Spjut (2007) was from Annapolis (Milo Baker s.n.).  In a brief survey of this area by Rich Spjut and Rick Spjut in August 2006,  we found only one tree (Rich Spjut & Rick Spjut 16021; Spjut 2007).

     However, additional plants have been found by Peter Baye at widely scattered spots along Fuller Creek and its north-facing drainages as noted by him in a report (letter) to the California Dept. of Forestry (; no longer available), and subsequently has found it at other locations (Peter Baye, pers. comm., Mar-June 2010), some of which have been studied by Spjut (Spjut & Baye proposed publ. on Taxus in Sonoma County).  Generally, plants of var. polychaeta occur as isolated individuals at intervals of 300–500 m, or may be found in small numbers of up to three or more plants within 50 m of each other.   Nonetheless, the occurrences are still considered  rare, especially in view of the narrow ecological zone in which they seem to occupy (Spjut & Baye proposed publ.).  Peter Baye has also found plants of var. brevifolia substantiated by a photo of arillocarpia sent to Spjut in June 2010 for confirmation.

     Spjut has since found var. polychaeta along Thompson Creek in Jackson Co.—in the Klamath Mountains Region of southern Oregon—in early August 2011 while conducting a survey for Veratrum californicum. Additionally one image placed on CalPhotos in 2011, reportedly taken from Opal Creek  Wilderness in the Cascade Range—showing a single arillocarpium—is similar to var. polychaeta in the relatively visible primary shoot, but the shoot scales overlap only slightly, appearing fewer in number and in series,  3 or 4 seriate.

     Other coastal occurrences of Taxus brevifolia were reported from a ravine on Sea Ranch in southern Mendocino County, an area reportedly logged in early 1900’s and again in the early 1990’s (Web report previously cited but no longer available online, or has changed), and also one specimen provided by the participants in the California Consortium of Herbaria collected from near the town of  Mendocino;; Fri Sep 3 14:49:50 2010; Wheeler & Gladys L. Smith 2681 (collected 29 Jul 1981), HSC83650.     

     It was reported that the specimen with a male cone (strobilus) collected by John Milton Bigelow from California corresponded to a male plant of var. polychaeta (Spjut 2007b, also shown below).  However, male plants in the general vicinity where female plants have been found of var. polychaeta appear similar to male cones of var. brevifolia, except for possibly being smaller in size.   Bigelow is known to have collected in Marin, Sonoma and Mendocino Counties (CNPS, Marin Chapter, website).  More field studies are needed during the spring, along with comparative DNA and morphological analyses.  This may clarify whether more than one variety or species of Taxus occurs in Sonoma County. 

Taxus brevifolia var. polychaetaCalifornia.  Sonoma Co.: near Mendocino Co. line, 7 km E of Stewarts Point; Rich Spjut & Rick Spjut 16021, Aug. 2006.  Redwood forest with grand fir, Douglas fir, and Torreya.  Here both Taxus and Torreya occur next to each other, their branches tangled (second photo from left).   Taxus rare, only one tree seen.  Seeds appearing immature, without aril (except for one shown in lower right photo), the ovules probably not pollinated due to lack of male plant in the vicinity.  Photo on top row far left shows single trunk of Taxus and also that of Torreya.  Lower row of photos by Rick Spjut (PhD student—Mathematics, Univ. Santa Barbara) shows close-up of bark, leaves with broad leaf margins, stipitate-like seed and enlarged scales around one developing ovule.

Taxus brevifolia var. polychaetaCalifornia.  Northwestern Sonoma Co., Rich Spjut & Peter Baye 16710, 22 Jun 2010.  Redwood forest  ecotone to grand fir-mixed evergreen forest with Umbellularia, Lithocarpus, Calycanthus, Cornus, Corylus, Euonymus, Vaccinium, Salix and other shrubs on north-facing rocky benches along stream.  Note branching of primary-secondary shoot complex, all containing more than one ovule—as many as five in top and bottom right images; thus, there is no clear terminal ovule.  This plant was found by Amelia Ryan in June 2010.  Bottom row image on left by Peter Baye, cneter by Richard Spjut, shoots with two arillocarpia by Peter Baye (Sep 2010)..

Taxus brevifolia var. polychaetaCalifornia.  Northwestern Sonoma Co., Rich Spjut & Peter Baye 16708, 22 Jun 2010.  See 16710 for more details on habitat.  Male plant. Lower right (bottom) photo by Peter Baye, 12-13 June 2010. Collected along river banks where scattered female plants of T. brevifolia had ovuliferous shoots described for var. polychaeta  Male strobili unlike the Bigelow specimen shown below, more like male strobili of var. brevifolia but not studied in detail.



Taxus brevifolia var. polychaetaCalifornia.  Northwestern Sonoma Co., Rich Spjut & Peter Baye 16712, 22 Jun 2010.

Taxus brevifolia var. polychaeta.  

Washington, Mud Bay. Possibly Mud Bay Road near Olympia. F. G. Meyer 1589 (Holotype K)A search was conducted in the vicinity of Mud Bay near Olympia in July 2010. No yew plants could be found.

Taxus brevifolia var. polychaeta

California. Salmon Creek, McMurphy 315

Idaho. Shipped overnight mail fresh from Coeur d'Alene by Marion Blatch, shipping clerk at Coeur d'Alene Nursery, to Richard Spjut at the USDA Agricultural Research Service in Beltsville, MD, without information on the collector and collector's number, locality of collection, and habit of the plant.  Sample collected by the USDA Forest Service as part of a number of samples collected in the Pacific Northwest during May-June 1992 in regard to an investigation of leaf anatomical characters being used for a taxonomic study of tree vs. shrub yews.  Note two terminal ovules on primary-secondary shoot complex. This specimen had been filed separately in the WBA herbarium because it lacked data, and it was not until reviewing the proofs for Spjut (2007) that this specimen was again seen—after nearly 15 years had passed.  Phone calls were made to various personnel at Coeur d'Alene Nursery in May or June 2007.  No one had any information as to who collected it. One person suggested that it could have come from along the Spokane River, because the environment there was thought to be unique.  Perhaps one reason why data were not sent with this specimen is because the collector may have felt that cones of Taxus should also be considered in the 1992 study.  Additionally, Taxus specimens at the Stillinger Herbarium were reviewed in July 2010; none could be identified as belonging to var. polychaeta.  No yew plants were observed in a search for yew near Coeur d'Alene, July 2010.

Taxus brevifolia var. polychaetaCalifornia.  Bigelow without number, year 1854.


3a. Taxus brevifolia var. brevifolia

3d. Taxus brevifolia var. klamathensis

3c. Taxus brevifolia var. reptaneta

References (Not included in the Introduction, or provided as full citations as mentiond above)


Anderson, E. D. and J. D. Owens. 1999. Megagametophyte development, fertilization and cytoplasmic inheritance in Taxus brevifolia. Int. J. Plant Sci. 160: 459–469.

Baye, P. 2004. Comments. THP-1-04-059-SON Martin (Sleepy Hollow) Timber Conversion Permit/Timber/Harvest Plan (Brushy Ridge, Annapolis, Sonoma County, California), to the California Dept. Forestry, 15pp.. (no longer available).

Clement-Westerhof, J. A. 1988. Morphology and phylogeny of Paleozoic conifers.  Pp. 298–337 in C. S. Beck (ed.), Origina and evolution of gymnosperms. Columbia Univ. Press, New York.

Foster, A. S. & E. M. Gifford, Jr. 1974. Comparative morphology of vascular plants.  2nd ed.  W. H. Freeman and Company, San Francisco.

Harris, T. M. 1976. The Mesozoic gymnosperms. Rev. Paleobot. and Palyn. 21: 119–134.

Judd, W.S., Campbell, S., Kellogg, E.A., Stevens, P. F. & M. J. Donoghue.  2002.  Plant Systematics.  A phylogenetic approach.  Sinauer Associates, Inc., Sunderland, MA.

Robertson, A. 1907. Taxoideae:  A phylogenetic study.  The New Phytologist VI: 92–102.

Spjut, R. W. 2007a. A phytogeographical analysis of Taxus (Taxaceae) based on leaf anatomical characters.  J. Bot. Res. Inst. Texas 1(1): 291–332

Spjut, R. W. 2007b. Taxonomy and nomenclature of Taxus.  J. Bot. Res. Inst. Texas 1(1): 203–289.