Monday, April 16, 2018

Resurrect Mahonia sonnei!

Mahonia sonnei was named by Abrams (1934) based on a single gathering from Truckee, California.  The syntype specimens were collected by Charles F. Sonne in August 1884 and again in April 1885 (no lectotype has been as yet designated). 

McMinn pigeon-holed M. sonnei in Berberis (McMinn 1939), as was then and still recently the custom.  We now know that Mahonia is a distinct genus well separated from Berberis, and from the related genera Alloberberis and Moranothamnus in both Asia and the New World (Yu and Chung 2017).

Poor Mahona sonnei: it has died twice.

First it disappeared for 90 years: the species was never collected again until 1973 when James B. Roof and members of the California Native Plant Society organized the founding “rare plant treasure hunt” (Roof 1974).

Mahonia sonnei was listed under the U.S. Endangered Species Act of 1973 in(Federal Register 44, p. 64246, 6 November 6 1979.  In 1993, Michael Williams placed it in synonymy in the first edition of the Jepson Manual (Williams 1993), where it has remained ever since (Williams 2012, Whittemore 1997).  Then it disappeared again - Mahonia repens was delisted – officially removed from the Endangered Species Act in 2003.  Poof!  The delisting was directly a result of a floristic treatment, not a detailed published study.

Abrams (1934) and McMinn (1939) keyed M. sonnei thusly:
Mahonia sonnei: Leaflets bright-green and shining above, teeth 12-16 on each margin, merely bristle-tipped; lower surface not gray
           Mahonia repens: Leaflets dull above; teeth small, bristle-tipped, usually 12 on a side.

In revisiting the Berberidaceae of California, I reviewed Abrams’ original description, and now that we have a wealth of imaged herbarium specimens, sought to do a quick, simple test of the number of  teeth per side of a leaflet.  Marginal teeth were counted on the 3 of the 4 syntype specimens of M. sonnei (NDG19699, US2699, DS95828), and from 70 specimens selected throughout the geographic range of M. repens (Fig. 2)

Figure 1 shows a box plot of the results: M. sonnei averages 15.9 teeth per leaflet side, while M repens averaged 10.2.  The difference is significant at p=<0 .0001="" df="68).<span" style="mso-spacerun: yes;"> 
Imaged specimens of M. repens were selected from throughout its wide range in western North America (Figure 2).

Figure 1.  Box plot of number of teeth on one side of leaflets from 3 of 4 syntypes M. sonnei and from 70 randomly chosen specimens of M. repens.  The difference in means is highly significant (T-test)

Figure 2.  Geographic distribution of M. repens, based on specimens served up on SEINet

This simple comparison suggests that M. sonnei and M. repens exhibit different leaf morphometrics.  Mahonia sonnei has never been DNA sequenced.  My inspection of specimens so far suggests that, overall, M. repens and the syntype specimens M. sonnei occupy differing leaf trait morphometric multivariate space (compare Fig. 3 and Fig. 4).  Additional work on this problem is planned: the expectation is that substantial morphometric differences between M. sonnei and M. repens can be demonstrated.  However, M. sonnei needs to be sequenced.

Figure 3 and 4.  Representative leaflets of M. sonnei (below) and M. repens (above, the leaflet chosen based on the observed mean of n=10 teeth)

Preliminary conclusion: Mahonia sonnei is a distinct taxon.  The baby was thrown out with the bathwater.

Literature Cited

Abrams, Leroy.  1934.  The Mahonias of the Pacific States.  Phytologia 1(2):89-94.
McMinn, Howard E.  1939.  An Illustrated Manual of California Shrubs.  Univ. of California Press, Berkeley, CA
Roof, J.B.  1974.  Found alive: the Truckee barberry.  Four Seasons 4(4):2-18.
Whittemore, A. 1997. Berberis. Pp. 276-286 in Flora of North America Editorial Committee (eds.), Flora of North America North of Mexico, Vol. 3. New York and Oxford.
Williams, M. 1993. Berberis. Pp. 362-364 in Hickman, J.C. (ed.), The Jepson Manual: Higher Plants of California. University of California Press, Berkeley.
Williams, M. 2012. Berberis. Pp. 446-447 in Baldwin et al. (eds.), The Jepson Manual: Higher Plants of California. University of California Press, Berkeley.
Yu, C. C., & Chung, K. F. (2017). Why Mahonia? Molecular recircumscription of Berberis sl, with the description of two new genera, Alloberberis and Moranothamnus. Taxon, 66(6), 1371-1392.

Saturday, June 24, 2017

A new metric of floristic botany

Astronomers are not want to utilize relative measures: astronomical units, a measure of apparent brightness magnitude adjusted for distance, serves as an example. If cosmology can advance with such relative units, why not floristic botany? 

Introducing the Steyermark

Julian Steyermark (1909-1988) collected over 130,000 numbers of vascular plant specimens, as noted in the Guiness Book of World Records His maximal collection number I found is 132,006 (ref 1).  I therefore propose a unit of floristic botany, the Steyermark (StyM), set equal to 132,006 gatherings.

Floristic exploration depends on field work.  Voucher specimens are requisite for field work, hence for floristic documentation.  Gathering, processing, data entry, labeling and mounting of specimens is work, tedious work.  Once prepared, a specimen nowadays needs to be imaged before it is filed.   Any given botanist, however equipped and dedicated, contributes a finite number of specimens.   Their, and thus my magnitude, are finite.

Two California botanists of note of note are mentioned here: Robert Folger Thorne (1920-2015) collected ca. 63,000 numbers (G. Wallace, pers. comm.).  John Thomas Howell (1903-1994) gathered ca. 55,000 numbers.  

Accordingly:  Thorne =0.477 StyM, while Howell=0.407 StyM.  Many active California botanists have reached the 0.10 to 0.12 StyM range.  This author is 0.16 StyM, but not likely to graduate beyond 0.2!  

We can also set a relative measure of floristic inventory: specimens per unit area (ref 2).  For the western U.S.A., let us here take this to be 90% percentile of collection density as determined on a county basis: 5.55 specimens/square kilometer.  Given this, only 3,797,482 additional herbarium specimens are needed - ~28 StyM.   Presently, the western U.S.A. is at a density equal to much less.

Hay bailing has its value.

1     1.) Ann. Missouri Bot. Garden 76: 652-790. 1989.   Phytoneuron 2014-53: ISSN 2153 733X

Tuesday, June 20, 2017

Ackerson Meadow acquisition adds only 4 vascular plants to the Yosemite National Park Flora?

In 2015, Yosemite National Park moved to acquire 415 acres at Ackerson Meadow, adjacent to the western Park boundary in Tuolumne County (in the vicinity of Mather and the road to Hetch Hetchy). This acquisition was facilitated by a donation of the owners to the Trust for Public Land, and then by conveyance to the National Park Service.

Ackerson Meadow is a rare feature in the Sierra Nevada: a mid elevation (~4000 feet) non-forested setting supporting wet to dry meadows.   Ordinarily, such an addition to the Park ought to add considerably to the documented Yosemite National Park flora, being that the western Park boundary generally excludes much low to mid-elevation settings of the western slope of the Sierra. 

The region of Ackerson Meadow was part of Yosemite National Park in 1890 (act of Congress, October 1st 1890).  It was then removed, and any Public land in the vicinity was conveyed to the Forest Service by Congress acting on June 11, 1906.  By the time that John Muir has successfully argued for an expanded Yosemite in 1890, the land in the vicinity of Ackerson Meadow had been granted to James F. Ackerson in 1882 and 1884.  Portions of the Ackerson Meadow system are now managed by two agencies and under two administrative directives: about 200 acres of the meadow system remains within the Stanislaus National Forest.

Generally, the flora of the Yosemite region is well documented (Taylor 2010, Baldwin et al. 2017).  The four counties flanking Yosemite are rank about the 90th percentile for the western U.S. (Taylor 2014).  In 2014, Yosemite National Park raked in the 97th percentile for specimen density.  My analysis, based on records of herbarium specimens in the Consortium of California Herbaria, the acquisition adds only 4 potential new vascular plant records for the Park.

Species Added to YNP

Eryngium vaseyi var. vallicola (Apiaceae, JEPS101766) – this vernal pool coyote thistle is perhaps a new species: Preston et al (2012) stated :”plants keying here from cSNH (Tuolumne Co.) may be an undescribed taxon.”  Obviously, this possibility alone is worthy of funding for additional study.

Erythranthe utahensis (Phrymaceae) – a single collection is attributed to Ackerson Meadow (R. K. Vickery 191, August 26, 1949. UT).  The record was accepted by Nesom (2012), with the provision “have the taller habit of E. utahensis but with fewer flowers as in E. corallina and an apparently intermediate vesture”.  Obviously, this possibility alone is worthy of funding for additional study: E. utahensis is a CNPS rare plant, List 2B.1

Polygonum polygaloides ssp. confertiflorum (Polygonaceae, JEPS96724) – an annual of open dry sites that are wet to saturated in spring.  This location is the southerly reported station for this taxon.

Hosackia pinnata (Fabaceae, JEPS100871) – This wetland perennial also reaches its southerly geographic distribution in the Yosemite region.  The more common sister species in Yosemite, Hosackia oblongifolia var. oblongifolia, has been collected ~50 times in the Park.
The purpose of this post is to emphasize that a detailed floristic inventory of the acquisition, using rigorous methodology (Groom & Whild 2017), is needed.  The Yosemite Conservancy has undertaken an initial inventory of Ackerson Meadow: extensive, vouchered floristic inventory over a period of several years will be required to ascertain the exact value of this addition to Yosemite NP.  

Groom, Q. J. and S.J. Whild.  2017.  Characterization of false-positive observations in botanical surveys.  PeerJ 5:e3324; DOI 10.7717/peerj.3324

Nesom, G. 2012.  Taxonomy of Erythranthe sect. Simiola (Phrymaceae) in the USA and Mexico. Phytoneuron 40: 1–123.  Published 16 May 2012. ISSN 2153 733X Corrections to Map 9 and Map 15, 21 May 2012

Taylor, D.W. 2014.  Large inequalities in herbarium specimen density in the western United States. Phytoneuron 2014-53: 1–8.  Published 2 June 2014. ISSN 2153 733X

Taylor, D. W. 2010.  Flora of the Yosemite Sierra.  Lulu Press, Raleigh, NC. 382 pp.  ISBN 780557500529

Baldwin, B.G. et al.   2017. Species richness and endemism in the native flora of California.  Amer. J. Bot. 104 (3): 487 – 501, 2017  doi:10.3732/ajb.1600326

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.

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.

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