Howellanthus dalesianus , Recognition of a New Genus and Species in Tribe Phacelieae (Boraginaceae)

2010·5.1 MB·English

by  Genevieve K Walden

Madrono, Vol. 57, No. 4, pp. 268-273, 2010
HOWELLANTHUS DALESIANUS, RECOGNITION OF A NEW GENUS AND
SPECIES IN TRIBE PHACELIEAE (BORAGINACEAE)
Genevieve K. Walden' and Robert Patterson
Department of Biology, San Francisco State University,
1600 Holloway Avenue, San Francisco, California 94132
[email protected]
Abstract
Howellanthus (Constance) Walden and R. Patt. is recognized as a new genus, containing a single
species H. dalesianus (J. T. Howell) Walden and R. Patt., based on morphological evidence.
Howellanthusdalesianus(Boraginaceae) is a small perennial herb, endemicto northern Californiaand
restricted to ultramafic soils. The species was first collected in 1936 by Ella Dales Cantelow and
Herbert Clair Cantelow, described by John Thomas Howell as Phacelia dalesicma J. T. Howell, and
later placed in the monotypic Phacelia subg. Howellanthus by Lincoln Constance.
Key Words: Boraginaceae, Howellanthus, Hydrophyllaceae, Hydrophylloideae, Phacelia, Phacelieae,
serpentine.
Phacelia dalesiana J. T. Howell was first loideae; and the ecologic specialization on
described in 1937, and named in honor of Ella ultramafic substrate, although it lacks the high
Dales Miles Cantelow, a long-time friend and ploidy level characterizing other paleoendemics
correspondent of John Thomas Howell (Camp- (Constance 1953; Favarger and Contandriopou-
bell n.d.; Howell 1954^1955, 1955, 1955-1957). los 1961; Stebbins and Major 1965). It occurs in
Ella Dales Cantelow, with her husband, Herbert the Siskiyou-Trinity mountains, an area of high
Clair Cantelow, collected specimens on a trip to concentration of Arcto-Tertiary relictual species
the Scott Mountains (Siskiyou Co., California) in (Stebbins and Major 1965).
1936. A duplicate was sent to Howell at the The strongest argument to retain the taxon
California Academy of Sciences for identifica- within Phacelia has been common, shared char-
tion, and the "remarkable new phacelia" piqued acters between species across several infrageneric
his interest sufficiently to request an expedition to divisions in the genus: the scorpioid cyme,
collect mature fruiting material (Fig. 1) (Cante- although few flowered, lax and axillary from a
low 1937-1940; Howell 1937). The following year basal rosette in P. dcdesiana (Figs. 2 and 3);
Howell and Alice Eastwood traveled to Scott presence of interstaminal corolla scales (Fig. 4),
Mountain, guided by the Cantelows, to collect modified in subg. Cosmanthus or absent in sect.
the type specimen. Howell returned again to Whitlavia and sect. Gyinnohythus; deeply parted
Scott Mountain in July to collect additional style, which can be lobed in some species of the
material with mature fruit for the description, genus;entireleavesandperennialhabit; and/7 = 8,
and published the new taxon (Howell 1937). shared with P. stebbinsiiConstance and Heckard,
HowCeolnlsatnatnJicues((/1?9=538))etroeccotnetdaithnethmeodniosttiynpcitcspseucbige.s NP.uttn.i,aracnesdceP.nscjEuaicsktiwi.J.eTx.JH.owFe.llM.acHborw.e,veP.r,gltahbersae
(see Constance 1953, p. 201 for scientific illustra- characters are also shared to varying degree
tion), based upon chromosome number differenc- with Draperia Torr., Hesperochiron S. Watson,
es and taxonomic revisions of infrageneric divi- Romanzoffia Cham., and Tricardia Torr. ex S.
sionsin Pluicelia(CaveandConstance 1942, 1944, Watson (Table 1), which left P. dcdesiana in an
1947, 1950; Constance 1949, 1950, 1951). The uncertain and isolated relationship with these
subgenus has since remained monotypic in treat- genera and Phacelia (Walden 2010).
(m/ezn=ts11o)fatnhdesguebngu.s,Cosseupiaarnattleidusf(rnom=s9u)b(g.CoPnhsatcaenlciea asTahmeesmpebceiresoifsesausbiflyamrielcyogHnyidzreodpmhoyrlplhooildoegaiec,albluyt
1963; Constance and Chuang 1982; Halse et al. is distinct from other members oftribe Phaceheae
1993; Ferguson 1998; Garrison 2007). andwarrants recognition asa separategenusfrom
Phacelia dalesiana has been considered a Phacelia. The taxon can be distinguished from
paleoendemic due to several factors: the system- Phacelia by the combination of perennial, acau-
atic isolation of the taxon within Pluicelia; a lescent habit, axillary few-flowered scorpioid
hypothesized relictual lineage within Hydrophyl- inflorescence, presence ofmulticellular glandular
trichomes with unicellular ellipsoidal heads,
Uni'vCeurrsrietnytoafddCraelsisf:orDneiap,arBtemrkeenlteyo,fCInAteg9r4a7t2i0v.e Biology, dseemeipolrybicpualratredcorsotlyllae,scpalaeisr,end =pe8n,daanndttroivcuollepso,-
2010] WALDEN AND PATTERSON: A NEW GENUS IN THE BORAGINACEAE 269
Fig. 1. Herbert Clair Cantelow, John Thomas Howell and Ella Dales Miles Cantelow on porch at Scott Ranch
(1936 photo by grandson Herbert Park Cantelow, courtesy ofthe Cantelow family and ofElla Dales and Herbert
Cantelow papers, California Academy ofSciences Archives).
Fig. 2. Habit ofHoweUcmthus dalesiamis. Bar = 1 cm.
MADRONO
270 [Vol. 57
Fig. 3. Close up of corolla, showing semiorbicular FiG. 4. Sideviewofopencorollaandunexpandedbud
scales at base offilaments. Bar = 1 cm. with presence ofabundant glandular trichomes. Bar =
cm.
1
rate pollen with exine striato-rugulate (Constance
andChuang 1982; Di FulvioandDottori 1995; Di umque 2, raro 4, alveolata; chromosomatum
Fulvio et al. 1999; Walden 2010). numerus /? = 8; species endemica terrae oriundae
ex rupibus ultramaficis.
Taxonomic Treatment Perennial herb, 5-15 cm, densely hairy due to
presence of two types of trichomes, unicellular
Howellanthus (Constance) Walden and R. Patt., eglandular verrucose-walled conical trichomes,
gen. et stat. nov. Phacelia subg. Howellanthus and multicellular glandular trichomes with uni-
Constance, Madrono 11:198-203. 1953.—Type cellular ellipsoidal heads. Basal leaves rosulate
species Howellanthus dalesianus (J. T. Howell) from caudex, cauline leaves alternate, sometimes
Walden and R. Patt., comb, nov., hoc loco appearing opposite on stems, reduced upward.
designatus. Phacelia dalesiana J. T. Howell, Leaf blades oblong to elliptic, simple, margins
mm
Leaflets of Western Botany, 2:51. 1937.— entire, with lateral incised venation, 10-50
Type: USA, California, Trinity Co., Summit long, blades more or less equal to petiole.
ofScott Mountain, 25 June 1937, A. Eastwood Inflorescence a lax scorpioid cyme from axils,
mm
andJ.T.Howell5014 (holotype: CAS!; isotypes one- to few-flowered, pedicel 10-20 in fruit.
CAS!, UC!, POM!). Sepals fused at base, lobes 5, unequal, oblanceo-
mm mm
Genus habitu Hesperochiron, differt cymis laxis late, accrescent, 3-5 long in flower, 4-7
mm
paucifloris, vix scorpioideis; trichomata glanduli- long in fruit. Corolla 5-10 in diameter,
fera, stipula multicellularia, capitataellipsoideaet campanulate, fused at base, deciduous, white,
unicellularia; corolla alba maculis purpureis in sometimes fading lavender in age, throat purple-
fauce; squamae interstaminales semiorbiculares; marked. Corolla scales semiorbicular, fused to
mm
filamenta glabra basibus non dilatatis; stylus base ofcorolla throat but not to filaments, 2
partitus ad basem; capsula suborbicularis, ovar- long. Filaments adnate to corolla at base,
ium quasi divisum placentis parietalibus spisses- glabrous, slender, included to slightly exserted,
mm
cens, ovulis 2^, geminatis, pendulis; pollinis 6-8 long, anthers purple, pollen tricolporate,
grana 3-colporata, striata-rugulata; semina pler- colpi nearly smooth, exine striato-rugulate. Style
1
2010] WALDEN AND PATTERSON: A NEW GENUS IN THE BORAGINACEAE 271
parted nearly to base, included to slightly
mm
exserted, 6-7 long, glabrous or sometimes
pubescent-hairy at base. Ovaries one-celled,
13 ovules paired (2-4 per ovary), pendant. Capsule
mm
4 long, subspheric. Seeds 2^, ovoid, brown,
''oa 2.5-4 mm long, surface alveolate, pubescent, n =
8. Blooming May-August.
i2 ;3
^3
Distribution
6 Hcnvellanthus dalesicmus occurs in the Scott
Mountains and Trinity Alps, near thejunction of
(aU
'a3 Shasta, Trinity, and Siskiyou counties (Fig. 5).
The type location is Scott Mountain Summit in
"o
"5- '1 Trinity County, intersected by State Highway 3
o (cOU Ccc« X) ^ ^tn 8 taynpdetlhoecalPiatcyifiisctChreesmtosTtraciolll(eFcetreldatstieng1l9e78p)o.puTlhae-
tion represented in herbaria, due to the accessi-
bility provided by the maintained highway and
forest trails. However, while the type locality may
appear accessible today, we speculate that the
ici« i3S -c3 species was not collected prior to 1936 due to
morphological similarity to Hesperochiron cali-
^ 2 fornicus (Benth.) S. Watson, which also occurs on
Scott Mountain, and the limited presence of
botanical collectors in the Scott Mountains prior
to 1930.
Forthree decades Scott Mountain was the only
13
knownlocality forthe species, but currently seven
B^ -^5 'oo populations have been vouchered in herbaria,
although three are from single collections. The
<U CD X 5t3 —T II S-H"^ USDA National Forest Service Shasta Trinity
C« C/2 X) and Klamath have surveyed and continue to
manage fiftypopulations forthe species on public
lands, with additional unvouchered reports of
^ S «^ ^ X -^5 ooO 73 CpaolpiufloartniioansDepoanrtpmreinvatteoflaFnisdh haonlddiGngasmeinNatth-e
O ural Diversity Database (Adamson and Kier-
stead-Nelson 1991). The USDA Forest Service
'o lists Phacelia dalesiana as a sensitive species, and
'oa S o t: the California Native Plant Society lists PJuicelia
II dalesiana as 4.3 (limited distribution, not very
threatened in California), California Endemic,
S3.3 (vulnerable) and G3 (vulnerable, no current
'o
'a threats known) (CNPS 2010; Showalter 1991).
o 'o The elevation ofthe species ranges from 1600
2200 m, on generally flat topography in openings
or dry meadows in upper montane coniferous
3 "o C3 forest (Adamson and Kierstead-Nelson 1991). All
> o
00 populations are on soils derived from ultrabasic
^
co a"^ n II 8 .Si r(oKcrkucaksesboceiragted19w8i4t;h tAhdeaTmrisnoitny UalntdramKaifeircstsehaede-t
Nelson 1991). Hcnvelkinthus dalesiauus has been
documented on disturbed areas, with variation in
litter cover within mature populations, although
the species requires bare soil for germination and
establishment (Adamson and Kierstead-Nelson
(U O 11999931);. TKhieersspteecaide-sNebllsooonmsafnrdomEnMgasytrtoomAug1u9s9t1,-
O a at: with flowering closely tied to snowmelt for a
U c/l ^ U Oh U short season, and fruits until late September.
MADRONO
272 [Vol. 57
Fig. 5. DistributionofvoucheredcollectionsofHoyveJlanthusdalesianusin Siskiyou,Trinity, andShastacounties,
California. Starred locality indicates type locality on map.
Additional Specimens Examined China Mountain quadrangle 1 July 1978, Clifton
and Ground 1662 (UC); Summit ofrd over Scott
USA. CALIFORNIA. Shasta Co.: Trinity Mountain, sandy flat of rocky meadow, 1 June
Mountains, saddle between Rattlesnake Hill 1946, L.Constance and R. H. Shan 3070 (CAS,
and Chicken Hawk Hill, at summit on Forest UC); Trail from Deer Flat to Shiny Lake ca.
RW.d 3Ta8yNl2o1r,(aJtEPSSa)r.diSnieskSipyroiungC,o.1:54JumneN1E993S,coDt.t D1/i4Tommaasboov6e00De(eUrC)C;reAelkonFglatU,S8FSJultyrail19786W,1/3.
Mountain, trail from Robbers Meadows to from Deer Flat to Shimmy Lake, 1/4 to 1/2 m E
Kangaroo Lake, 1 June 1977, S. Homer 110 of Deer Flat Camp, 7 July 1976, W. J. Ferlatte
(JEPS); Mt. Shasta Ranger District, Mumbo and J. DiTomaso 1776 (CAS, JEPS); Summit of
Basin at 40N26 and 38N24, 20 June 1991, /. Scott Mountain, N ofCarrville, 24 August 1936,
Kierstead 91-68 (ST); Scott Mountain divide, 17 /. T. Howell 12736 (CAS); Summit of Scott
May 1947, H. L. Mason 14773 (UC); Kangaroo Mountain, N of Carrville, 30 July 1937, /. T
Lake Campground, along streams, 25 July 1969, Howell 13691 (CAS); Scott Mountain summit on
F. W. Oettinger s.n. (CAS, RSA, UC); Kangaroo rd from Carrville to Callahan, 20 May 1980, /. T.
Lake, 25 July 1967, D. Parker and W. Roderick Howell, T. C. Fuller and G. D. Barbe 53545
s.n. (CAS); Scott Mountain Summit, 21 June (CAS); Summit of Scott Mountain, rocky ser-
2005, R. W. Patterson & S. Santos 1982 (SFSU); pentine soil, 9 June 1958, D. Parker 649 (CAS,
Scott Mountain rd summit, upper end of N RSA); Scorpion Lake, Trinity Mountains, west-
meadow, 14 June 1963, W. Roderick s.n. (JEPS); erly upper flank Bonanza King, 13 June 2002, D.
Scott Mountain campground, 18 June 2010, G. W. Taylor 18156 (JEPS). Shasta Co.: Trinity
K. Walden 324 (SFSU); Scott Mountain camp- Mountains, saddle between Rattlesnake Hill and
ground, 7 July 2010, G. K. Walden 332 (SFSU). Chicken Hawk Hill, at summit on Forest Rd
Trinity Co.: Summit of Scott Mountain, 21 May 38N21, at Sardine Spring, 15 June 1993, D. W.
1936, E. D. Cantelow 1276 (CAS); Summit of Taylor 13656 (JEPS).
Scott Mountain, 9 June 1939, E. D. Cantelow
Acknowledgments
2891 (CAS); Summit ofScott Mountain, 23 June
1948, E. D. Cantelow s.n. (CAS); Near Scott This represents a portion of work resulting from a
Mountain Summit, close to the town ofCallahan, master's thesis by GKW, and was previouslypresented.
2010] WALDEN AND PATTERSON: A NEW GENUS IN THE BORAGINACEAE 273
in part, by the authors at the Cahfornia Botanical Di FULVio,T. E. andN. Dottorl 1995. Contribucion
Society Graduate Student Meeting2010, San Jose State al conocimiento de tricomas y emergencias en
University, California, as Pluicclia dalesicma J.T.How- Hydrophyllaceae. Clasificacion y consideraceones
ell, a remarkable paleoendemic phacelia." taxonomicas. Kurtziana 24:19-24.
GKThWisfprroomjeNctSFwaGsRsFuPp,porNtSedF,TiRnEpEa,rt,anbdyNfSunFdiGnKg1t2o Mor,foMl.ogiTa.yCvoassAcu,larainzadcioNn.floDroalttdoerDlrape1r9i9a9,.
fellowships and research grants by SFSU Department Eminouinthe, Hesperochiron. Rouuinzojfia, y Tri-
ofBiology and CNPS chapters. We are grateful to the cardia (Phacelieae, Hydrophyllaceae). Kurtziana
following for field, herbarium, archive, and laboratory 27:187-209.
assistance: Deb Trock, Andrew Doran, Ellen Dean, Favarger, C. and J. Contandriopoulos. 1961.
Sula Vanderplank, Danielle Castronovo, the Cantelow Essai sur fendemisme. Bulletin of the Society
family, Frank Cipriano, Dennis Desjardin, Julie Kier- Botany Suisse 71:383^08.
sotfeGadK-WNe)l.sonW,eJitmhanLiknntbheerge,ditaonrdaTnrdigtgewro(asnerovnicyemoduosg Fergruelsaotnio,nshDi.psMi.n H1y99d8r.ophPyhyllloagceenaeetibcasaendalyosnisndalnidF
reviewers who substantially improved this paper. sequence data. Systematic Botany 23:253-268.
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