Tuesday, January 17, 2017

Minority opinion on classification of Ponderosa Pine

The recent study of Willyard et al (2017) on lineages of Ponderosa Pine (Pinus ponderosa) is important and at the same time highly problematic.  The study strongly demonstrates that Pinus ponderosa is a complex lineage containing several distinct taxa which merit taxonomic treatment. 
Willyard et al (2017) suggest treatment of the taxa in the lineage as full species:  Pinus ponderosa, P. benthamiana, P. scophulorum and P. brachyptera.   

This will not find wide acceptance in the non-botanical world of commerce.  A long history of practical use, common understanding, and governmental inertia will result in default back to a broadly circumscribed, single species model.  Unfortunately, the single taxon model fails to account for geographic and evolutionary distinct units within this clade, and is not a scientifically supported, proper classification.  Put crudely, the guy at the lumber yard is not going to care...!

The principal purpose of any classification is utility.  Treatment of the taxa of the Pinus ponderosa lineage as subspecies would have a much higher probability of gaining wide application.   Without reference to geography, the identification of individual trees is not easy, even with mature cones and bud-color features at hand, because the overall morphogical similarity between the individual taxa is substantial (Callaham 2013).  Users of a classification which employs infrataxa can refer only to Pinus ponderosa generally in instances where their determination of subspecies is unimportant (i.e. this timber is ponderosa pine) or in geographic areas where the taxa overlap (i.e. in the KR-CaRH region of California, where both P. ponderosa and P. benthamiana haplotypes are present). 

The five validly published taxa treated as subspecies are:
Pinus ponderosa P. Lawson & C. Lawson ssp. ponderosa, Agric. Man. 354 (-355). 1836
P. ponderosa ssp. ssp. benthamiana (Hartw.) Silba -- J. Int. Conifer Preserv. Soc. 16(1): 30. 2009
P. ponderosa ssp. brachyptera (Engelm.) Silba -- J. Int. Conifer Preserv. Soc. 18(1): 16. 2011
P. ponderosa ssp. scopulorum (Engelm.) A. E. Murray -- Kalmia 12: 23. 1982
P. ponderosa ssp. washoensis (H. Mason & Stockw.) A. E. Murray, Kalmia 12: 23. 1982.

The treatment of the taxa of Pinus ponderosa as subspecies is consistent with broad practice for many higher plants.  Treatment of the clade as varieties (where not all of the required names are valid) is not consistent with treatment models which have survived the test of time (Simpson 1945, Clausen 1951 Dobzhansky 1951, Stebbins 1966, Mayr 1982) and is inconsistent with common practice in the larger portions as applied to higher plants (Briggs & Walters 1971) worldwide.

In plant taxonomy, my opinion is that we devalue the worth of our effort when we do not defer to the practical needs of society at large whom consume our names.  

The five ponderosa pines are subspecies.

References:
Briggs, D., & S. M. Walters.  1971    Plant Variation and Evolution.   White & Gilbert, U.K.
Callaham , R. Z. 2013 . Pinus ponderosa : Geographic races and subspecies based on morphological variation. USDA Forest Service, Pacific Southwest Research Station PSW-RP-265. Washington, D.C.
Clausen J.  1961.  Stages in the evolution of plant species.  Cornell Univ. Press, Ithaca, N.Y.
Dobzhansky, T.  1951.  Genetics and the origin of species.  Columbia Univ. Press, New York, N.Y.
Mayr, E.  1982.  Growth of Biological Thought: Diversity, Evolution, and Inheritance. Harvard Univ. Press Cambridge, M.A.
Simpson, G. G. 1945. The Principles of Classification and a Classification of Mammals. Bull. American Museum Nat. Hist. 85: 1-23.
Stebbins, G. L. 1966.  Processes of organic evolution.  Prentice Hall, Engelwood Cliffs, N.J.
Willyard, A. W. et al. 2017.  Pinus ponderosa: A checkered past obscured four species.  Amer. J. Bot. 104(1):1-21.

Wednesday, December 16, 2015

Horkelia marinensis in Santa Cruz County - self-compatible?

Horkelia marinensis is a California endemic, endangered plant known from limited geographic regions along the California coast.  At the southerly geographic extent, in Santa Cruz County, there are 5 extant occurrences known (and three other old records that have not been georeferenced).

This note reports that Horkelia marinensis is probably self-compatible.   In 2005, I salvaged several plants that were being taken by instillation of a electrical transmission line pole.  The site (JEPS109793) is at 2500 feet on Ben Lomond Mountain.  The Horkelia occurrence here contained about 1000 plants, most of them growing in the road under the powerline for perhaps half a mile, the adjacent brush being very dense (see power line ROW) 



All but one of the plants dug died, most of the plants I dug had large taproots and hence did not make it.  The survivor, now 10 years old, is about one-half a meter in diameter and flowers nicely.   Several years ago, seedlings from this plant began to grow in pavement and walkway cracks.  This photo shows one on the side of the driveway.  That plant is perhaps 5 years old now. 



The apparent fecundity of this single individual of Horkelia marinensis suggests that it is self-compatible.

Also:  
Two Santa Cruz county occurrences do not yet appear in NDDB
No. 1 UCSC8932 on a ridge draining to San Vicente Creek
No. 2 Moore Creek Greenbelt site, Santa Cruz

Wednesday, October 7, 2015

David Douglas in California, and the type of the Santa Cruz Tarplant (Holocarpha macradenia)

Type stations are vague: many type specimens of California endemic plants collected long ago cannot be attributed to a specific date or specific place.

The type specimen of the Santa Cruz tarplant was collected by David Douglas.  In, 1836 DeCandolle described it as Hemizonia macradenia in Prodromus (“Systematis Naturalis Regni Vegetabilis”), a 17-volume treatise on botany last updated by his son A. de Candolle in October 1873. 

The protologue statements are: “in Nova California legit cl. Douglas” and “v. s. comm. ab hon. Soc. hort. Lond.).  [recall here that typesetting was then manual and abbreviations were used prolifically, but not exactly diagnosed: none of these abbreviations derive specific problems here]. 

There are three specimens of the original Douglas collection, two at Key and one at the de Candolle herbarium in Geneva: barcodes K001079845, K001079846 and G00453655.  All three sheets are attributed to Douglas and to 1833.  Based on the narrative detailing Douglas travels in California (in “California Frontier Naturalists” Biedelman 2006 pages 116-125, the best approximation is that Douglas collected the fall of 1831 and somewhere within a day’s ride of Monterey.  The most probable station therefore being in far northern Monterey County.  One proviso is that H. macradenia is often in peak bloom in September and early October, and the two Kew specimens are clearly in nice flower.  If Douglas had ventured north to Santa Cruz in the fall of 1831, then no mention is given in Biedelman, hence the type station is not very probably directly attributed to Santa Cruz county.  Accordingly, this raises the possibility that H. macradenia might have been VERY extensive in northern Monterey County, and since has contracted significantly. Presently, there is but a singe Monterey County occurrence.  Threrefore, Douglas may have obtained the type somewhere in the vicinity of present day occurrences at Elkhorn Slough and vicinity. 

Based on the narrative in Biedelman (2006) Douglas specifically visited Santa Cruz in February, 1831, but H. macradenia would not have been in flower at that time.

Regardless, the type was not likely to have been collected in 1833, because on November 4th, 1833 Douglas only arrived at San Francisco via Ft. Vancouver, having been away from California.  Douglas had been at destinations other than those in the known geographic range of H. macradenia between March 1832 and August 1832, thence he departed San Francisco for the Sandwich Islands (Hawai’i) in November, then returned again, departing the final time at the end of November 1833.  Holocarpha macradenia has never been documented on the San Francisco peninsula. 


Accordingly, I attribute the holotype to: Monterey County, vicinity “Bolsa Neuva Moho Cojo” [36.80023/- 121.71204], David Douglas s.n. , ±September 1831.

Tuesday, October 6, 2015

Plants that merit California Rare Plant Rank Status

Presently, 2345 plants enjoy California Rare Plant Rank (CRPR) status.  Status review for new additions proceeds at a pace of 30-40 taxa yearly.  Consequently, there is presently quite a backlog of newly described endemic plants California region plants that may qualify for CRPR status but which have not been reviewed.  Below I list these, and offer a preliminary recommendation for potential CRPR status.

Merit List 1B (14 plants)
Apiaceae                Spermolepis infernensis G. L. Nesom         List 1B.1
Phrymaceae           Erythranthe percaulis G. L. Nesom             List 1B.1
Rhamnaceae          Ceanothus decornutus V. T. Parker            List 1B.1
Brassicaceae          Streptanthus glandulosus Hook. var. josephinensis M. S. Mayer      List 1B.2
Brassicaceae          Streptanthus tortuosus Kell. var. truei Al-Shebaz   List 1B.2
Fabaceae               Trifolium piorkowskii Rand. Morgan & A. L. Barber              List 1B.2
Brassicaceae          Streptanthus glandulosus Hook. var. arkii M. S. Mayer        List 1B.3
Brassicaceae          Streptanthus glandulosus Hook. var. rachiei M. S. Mayer   List 1B.3
Caryophyllaceae     Silene krantzii T. R. Stoughton      List 1B.3
Cyperaceae            Carex xerophila Janeway & Zika   List 1B.3
Lamiaceae             Monardella mohavensis  Elvin & A. C. Sanders       List 1B.3
Polemoniaceae       Gilia ochroleuca M. E. Jones ssp. lanosa Hrusa      List 1B.3
Polygonaceae         Chorizanthe minutiflora Rand. Morgan & Reveal   List 1B.3

Merit List 4 (14 plants)
Ericaceae              Vacciniumn shastense J. K. Nelson & L. Lindstrand ssp. shastense   List 4.1
Polemoniaceae      Navarretia paradoxiclara L. A. Johnson & D. Gowen            List 4.1
Polemoniaceae      Navarretia paradoxinota L. A. Johnson & D. Gowen            List 4.1
Rosaceae              Holodiscus dumosus var. cedrorum Raiche & Reveal           List 4.1?
Asteraceae           Cirsium scariosum Nutt. var. robustum D. J. Keil    List 4.2
Convolvulaceae     Calystegia collina ssp. apricum Brummitt & Namoff       List 4.2
Polygonaceae        riogonum umbellatum Torr. var. nelsoniorum      List 4.2
Pteridaceae          Adiantum shastense Huiet & A. R. Sm.       List 4.2
Themidaceae        Brodiaea rosea Baker ssp. vallicola Preston            List 4.2
Brassicaceae        Streptanthus purpureus Sanchez-Mata et al.          List 4.3
Ericaceae             Vacciniumn shastense J. K. Nelson & L. Lindstrand
                             ssp. nevadense  J. K. Nelson & L. Lindstrand       List 4.3
Hydrophyllaceae   Nemophila hoplandensis M. Baar               List 4.3
Phrymaceae         Diplacus compactus (D.M. Thompson) G. L. Nesom             List 4.3

Uncertain rank status (4 plants)
Lamiaceae            Monardella australis Abrams ssp. gabrielensis Elvin & A. C. Sanders
Lamiaceae            Monardella australis Abrams ssp. occidentalis Elvin & A. C. Sanders
Polemoniaceae    Linanthus dichotomus Benth. var. pattersonii J. M. Porter
Polemoniaceae    Linanthus maculatus (Parish) Milken
                             ssp. emaculatus J. M. Porter, D. S. Bell & R. Patt.

Reject List (recommended CBR rank)
Campanulaceae
Nemacladus tenuis (McVaugh) Morin var. aliformis Morin
Cyperaceae
Calliscirpus brachythrix C. N. Gilmour, J. R.Starr & Naczi
Cyperaceae
Carex orestera Zika
Ericaceae
Arctostaphylos glandulosa Eastw. ssp. leucophylla J. E. Keeley, M. C. Vasey & V. T. Parker
Lamiaceae
Monardella linoides A. Gray ssp. sierrae  Elvin & A. C. Sanders
Orchidaceae
Spiranthes stellata P. M.Br., Dueck & K. M.Cameron
Papaveraceae
Eschscholzia papastilii Still
Pinaceae
Abies magnifica A. Murray bis var. critchfieldii Lanner
Poaceae
Festuca roemeri (Pavlick) E. B. Alexeev var. klamathensis B. L. Wilson
Pteridaceae
Pentagramma triangularis (Kaulf.) Yatsk. et al. 
var. rebmanii Winner & M. G. Simpson
Rosaceae
Drymocallis pseudorupestris (Rydb.) Rydb. var. crumiana Ertter
Rosaceae
Rosa pisocarpa A. Gray ssp. ahartii Ertter & W. H. Lewis
Cactaceae
Cylindropuntia chuckwallensis M. A. Baker & M. Cloud-Hughs
Onagraceae
Clarkia tembloriensis Vasek ssp. longistyla Vasek
Orchidaceae
Spiranthes stellata P. M.Br., Dueck & K. M.Cameron ssp. perexilis Sheviak



Friday, July 3, 2015

California Botany Batting Average

The Consortium of California Herbaria database continues to grow.  The rate plot shown below suggests that for the past year the rate of additions has become constant.

Added are an average of 341 specimen records per day

The ability of individual botanists and governmental agencies to keep up with the constant, essentially relentless, addition of data suggests to me a PROBLEM.  We should not to stop adding records, but we do need a mechanism that new records get moved between platforms (vis. Calflora, CNDDB) instantly.

Conservation of the endemic rich California flora is dependent on good data.  Funding to support programming/application development that would result in real-time data ought to be a distinct objective that merits the attention of botanists.


Monday, March 23, 2015

Sagebrushes that Almost make California

California is far from deprived of sagebrush vegetation, but is deprived of some sagebrushes (Artemisia Subg. Tridentatae).  Several species approach the eastern California region, and should be sought out, either in Modoc County or in the mountains and eastern Mohave desert.  (Schultz 2012) displays a small (thus low resolution) ecological array depicting sagebrush habitat selection pattern (which I reproduce here).  California has species that fill her entire array (green stars); these ‘missing’ species are the red stars (this also discounts a report of A. tripartita from CA)

Artemisia filifolia
“Sand sage” is distributed over much of Arizona, New Mexico, nw Texas, north in the western Great Plains to South Dakota.  Artemisia filifolia is absent from far southern Nevada (Schultz 2009).  It has been collected often within Washington County in far sw Utah, and in adjacent parts of Arizona.  One record from Mohave County, Hualpai Mountains, : R. Darrow s.n., June 1 1942 (ARIZ52484) if correct would be the most proximal to CA, within ca. 50 miles.    Artemisia filifolia was not reported for the Hualpai Mountain flora (Butterwick et al. 1991), nor was the specimen cited by or mapped by (Schultz 2009).  Absent from the Whipple Mountains (DeGrout 2007)

Artemisia papposa
Centered in the Snake River plains of Idaho, occurrences are known from ncNV in Elko County.  There are very few records generally, and there are no PNW Consortium collections from far se Oregon, where Packard 76-165 (UTC154100) was gathered.  Described in the Pocket Sagebrush Guide (Schultz 2012) as occurring in ‘pockets’ of soil in barren (their term, equal to recent?) lava fields, at 1400-2100 meters elevation (the elevation given both in Intermountain Flora and in FNA Vol. 19).  BONAP records an Elko County, Nevada record.  The center of distribution is in the Owyhee Desert bioregion of Intermountain Flora.  The geographic range west of the mapped occurrences in SEINet are poorly collected.   Mansfield (2000) did not report it in the vicinity of the Steens Mountains, Harney County, Oregon.  If A. papposa were disjunct to the Modoc Plateau (ca. 200 km westward) it would be expected only in far northeast Modoc County.  Artemisia papposa is a low subshrub rarely >2 dm tall, and was not treated in Artemisia Subg. Tridentatae by (Schultz 2009); in FNA Vol. 19 she placed it in Sect. Artemisia and remarked it show alliance to Sphaeromeria.

Artemisia pygmaea
Schulz (2009) maps a record in the vicinity of Clark County, Nevada but does not cite the specimen.  No record is reported from the Spring Mountains (Niles & Leary 2007).  In far northern Arizona, where it is a rare plant, there are records on the Grand Canyon north rim, and a redacted record for Mohave County.  The elevation range given is 1500-2000 meters (Schultz 2009) or 1500-1800 meters in FNA Vol. 19.  Reported from ‘barren outcrops’, ‘fine textured clay soils’ or ‘cemented ash’ (Schulz 2012) or gypsum and shale (Schultz 2009). This compact, low, aromatic sage, would not be unexpected, but would be rare if it were to occur in Inyo or northeastern San Bernardino County, California.

Artemisia rigida
In Steens Mountains ‘rare on rocky slopes below 1600 meters Mansfield (2000).  One occurrence in Harney County T. M. Barkley et al. s.n. in 1956 Squaw Butte Experiment Station, on Juniper Ridge north of the Headquarters” is ca. 115 km from Modoc County.  BONAP does not record A. rigida from Klamath County, but does have a record for Douglas County in a cismontane setting (which is suspect...).  Capable of sprouting after fire (Schultz 2009).  Occurs on loams or montmorillonic clays (which characteristically swell in volume when wet), and at 1500-1800 meters (Schultz 2009). 



Literature Cited
BONAP. 2015.  Biota of North America (www.bonap.org)
Butterwick, M, BD Parfitt, D. Hillyard. 1991.  J. Arizona-Nevada Acad. Sci. 24:31-49.
DeGrout, SJ.  2007.  Aliso 24:63-96.
Mansfield, DH 2000.  Flora of Steens Mountain.
Niles, WE and PJ Leary. 2007. Mentzelia Vol. 8
Schulz, LM.  2009. Systematic Botany Monographs Vol. 89
Schulz, LM. 2006.  Artemisia, in Vol. 19, Flora North America.

Schulz, LM.  2012. Pocket Guide to Sagebrush (www.sagestep.org)

Wednesday, March 4, 2015

Endemic vascular plant genera of the California region

There are on the order of 67 genera endemic to California and the California Floristic Province (‘California region’).  The list below gives my most recent compilation. About half of these are monotypic.  The total number of treated taxa is 155, or about 6% of the endemic flora of the region.

Agavaceae
           Chlorogalum Kunth (8)
Apiaceae
           Oreonana Jeps. (3)
Asteraceae
Achyrachaena Schauer    (1)
Adenothamnus D. D. Keck (1)
Baeriopsis J.T. Howell (1)
Benitoa D. D. Keck (1)
Blepharizonia (A. Gray) Greene (2)
Calycadenia DC. (10)
Constancea B. G. Baldwin (1)
Corethrogyne DC (1)
Eastwoodia Brandegee (1)
Holocarpha Greene (4)
Holozoinia Greene (1)
Jensia B. G. Baldwin (2)
Kyhosia B. G. Baldwin (1)
Hesperevax (A. Gray) A. Gray (3)
Monolopia DC. (5)
Munzothamnus P. H. Raven (1)
Orochaenactis Coville (1)
            Pentachaeta Nutt. (8)
            Phalacroseris A. Gray (2)
            Pseudobahia (A. Gray) Rydb. (3)
            Tracyina S. F. Blake (1)
            Venegasia DC. (1)
Brassicaceae
             Euclisia (Nutt. ex Torr. & A. Gray) Greene (9)*
             Heterodraba Greene (1)
 ‘Kruckeberia’ (10)
             Sibaropsis S. Boyd & T.S. Ross (1)
             Twisselmannia Al-Shebaz (1)        
Cactaceae
               Bergerocactus Britton & Rose (1)
Chenopodiaceae
               Extriplex E. H. Zacharias (2)
Crassulaceae
               Sedella Britton & Rose (30
Cyperaceae
               Calliscripus C.N.Gilmour , J.R.Starr & Naczi (2)
Ericaceae
               Ornithostaphylos Small (2)
Fabaceae
               Pickeringia Nutt. (2)
Hydrophyllaceae
               Draperia Torr. (1)
  Lemmonia A. Gray (1)
Hydrangeaceae
               Carpenteria Torr. (1)
Lamiaceae
               Acanthomintha (A. Gray) A. Gray (5)
  Pogogyne Benth. (8)
Lauraceae
               Umbellularia (Nees) Nutt. (2)
Monotropaceae
               Sacrcodes Torr. (1)
Oleaceae
               Hesperelaea A. Gray (1)
Onagraceae
               Heterogaura Rothr. (1)
Papaveraceae
            Dendromecon Kell. (2)
Ehrendorferia Fukuhara & Lidén (2)
Hesperomecon Greene (1)
Romneya Harv. (2)
Stylomecon G. Taylor (1)
Poaceae
               Neostapfia Burtt-Davy (1)
Polygonaceae
           Acanthoscyphus Small (4)
           Aristocapsa Reveal & Hardham (1)
           Dedeckara Reveal & J. T. Howell (1)
           Gilmania Cov. (1)
Goodmania Reveal & Ertter (1)
           Hollisteria S. Watson (1)
Mucronea Benth. (2)
Sidotheca Reveal. (3)
Systenotheca Reveal & Hardham (1)
Rosaceae
               Chamaebatia Benth. (2)
               Lyanothamnus A. Gray (2)
Saxifragaceae
               Bensoniella C. V. Morton (1)
               Jepsonia Small (3)
Taxodiaceae
               Sequoia Endl. (1)
               Sequoiadendron J. Buchholz (1)
Tecophilaeaceae
               Odontostomum Torr. (1)
Themidaceae
               Bloomeria Kellogg (5)


*genera as indicated in Cacho et al. 2014 (Mol. Phylo. And Evolution 72:71-81), but not yet described