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XI 



PHYTOLOGIA 



An international journal to expedite plant systematic, phytogeographical and 

ecological publication 



Vol. 87 March 2005 No. 1 



J. L. PANERO, New combinations and infrafamilial taxa in the Asteraceae 1 -^ 

J. R. SPENCE, New genera and combinations in Bryaceae (Bryales, Musci) for 
North America j5 -^ 

D. B. WARD, Keys to the flora of Florida - 12, Rubus (Rosaceae) 29 ^ 

B. L. TURNER, A new species of Seymeria (Scrophulariaceae) from Oaxaca, 
Mexico 40 ^ 

B. L. TURNER, A new species of Ageratum (Asteraceae; Eupatorieae) from 
Nayarit, Mexico 44 ^'" 

B. L. TURNER, Bidens melchertii (Asteraceae), a new species from Jalisco, ^ 
Mexico 4^ ' 

B. L. TURNER, A new species of Trixis (Asteraceae: Mutisieae) from Oaxaca, 
Mexico 50 '^ 

B. L. TURNER, Elevation of Stevia lemmonii var. hispidula (Asteraceae: 
Eupatorieae) to specific rank 53 -^ 

Botanical Anecdotes: Bajacalia moranii (Asteraceae: Tageteae), comments regarding its 
discovery 5^ 

Botanical Humor 5g 

Index to new names and new combinations in volume 87(1) 59 

Subscription Information & Information for Authors Inside back Jacket 



PHYTOLOGIA 

An international journal to expedite plant systematic, phytogeographical and 

ecological publication 

Vol. 87 March 2005 No. 1 

J. L. PANERO, New combinations and infrafamilial taxa in the Asteraceae 1 "^ 

J. R. SPENCE, New genera and combinations in Bryaceae (Bryales, Musci) for v. 
North America 15 

D. B. WARD, Keys to the flora of Florida -- 1 2, Rubus (Rosaceae) 29 '^ 

B. L. TURNER, A new species of Seymeria (Scrophulariaceae) from Oaxaca, 
Mexico 40 ^ 

B. L. TURNER, A new species of Ageratum (Asteraceae; Eupatorieae) from 
Nayarit, Mexico 44 ' 

B. L. TURNER, Bidens melchertii (Asteraceae), a new species from Jalisco, ^^ 
Mexico 47 

B. L. TURNER, A new species of Trixis (Asteraceae: Mutisieae) from Oaxaca, 
Mexico 50 

B. L. TURNER, Elevation of Stevia lemmonii var. hispidula (Asteraceae: 
Eupatorieae) to specific rank 53 

Botanical Anecdotes: Bajacalia moranii (Asteraceae: Tageteae), comments regarding its 
discovery 56 

Botanical Humor 58 

Index to new names and new combinations in volume 87(1) 59 

Subscription Information & Information for Authors Inside back Jacket 



.<\ M' 




PHYTOLOGIA 

(ISSN 00319430) 

Phytologia, a journal of plant systematics, phytogeography and 
vegetation ecology, is published three times a year by the Warner 
Herbarium, a research unit of the Department of Biological Sciences, 
Sam Houston State University. 



Editor 



Justin K. Williams 
e-mail: bioJkw@shsu.edu 
PHONE: (936) 294-1552 
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Phytologia 

P.O. Box 2012 

Sam Houston State University 

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Copyright 2005 Phytologia. Printed by the Sam Houston Press. 



Phytologia (Mar 2005) 8 7(1) 1 



NEW COMBINATIONS AND INFRAFAMILIAL TAXA IN THE 
ASTERACEAE 

Jose L. Panero 

Section of Integrative Biology, University of Texas, Austin TX 78712, 

USA 

ABSTRACT 

Molecular studies of Asteraceae support the recognition of tribe 
Doroniceae of subfamily Asteroideae, subtribes Anisopappinae, 
Athroisminae, and Centipedinae of tribe Athroismeae, 
Chromolepidinae, Dugesiinae, Enceliinae, and Spilanthinae of tribe 
Heliantheae, and Dyscritothamninae and Jaegeriinae of tribe 
Millerieae. The combinations Euphrosyne acerosa, Euphrosyne 
dealbata, and Euphrosyne nevadensis are also proposed. 

KEY WORDS: Athroismeae, Asteraceae, Asteroideae, Classification, 
Euphrosyne, Heliantheae, Millerieae. 



Comparative studies of DNA sequence data for several coding 
regions of the chloroplast DNA of the Asteraceae have revealed several 
lineages that require names to maintain a classification that reflects 
monophyletic groups. These studies expand on the results of Panero 
and Funk (2002) and are aimed at elucidating the major lineages of the 
Asteraceae recognized at the tribal level. 

Doroniceae Panero, tribus nov. -Type: Doronicum L. 

A Asteroideae ceteris, differt characteribus conjuncte: habitu 
herbaceo; foliis altemiis, petiolatis, laminis integris; capitulis ligulatis; 
phyllariis 2-3 seriatis; receptaculis epaleatis; flosculis radiorum 
pistillatis; flosculis discorum bisexualibus, corollis actinomorphis, lobis 
5; antheris ecaudatis; ramis stylorum truncatis; cypselae teretibus 10- 
angularibus; pappi radiorum presentibus vel absentibus, pappi 



2 Phytologia (Mar 2005) 87(1) 



discorum presentibus, pappi 1-3 seriatis, setis papporum albis vel albo- 
flavis. 

Perennial rhizomatous herbs. Leaves alternate, petiolate, 
sometimes clasping, blades ovate, elliptic or obovate. Capitulescences 
terminal, solitary or open paniculiform cymes. Capitula radiate. 
Involucres hemispherical, phyllaries in 2-3 series, herbaceous to 
somewhat chartaceous. Ray florets fertile, bifid or trifid, corollas 
yellow. Disc florets bisexual, fertile, corollas yellow, essentially 
glabrous or sparsely to moderately pubescent at base. Cypselas 
cylindrical to obovate with 10 ribs, brown to greenish or blackish, 
essentially glabrous to pubescent, pappose, sometimes absent on ray 
florets, pappus of capillary bristles arranged in 1-3 series, white to 
yellowish. Base chromosome number, x = 30. 

Monotypic, Doronicum contains 26 species found mostly in Asia, 
Europe and northern Africa (Alvarez-Fernandez, 2003) 

The genus Doronicum, because of its distinctive chemistry and 
multiseriate involucres, has traditionally been allied to Arnica L. and 
relatives of the Madieae (Nordenstam 1977). Nordenstam (1994) 
believed Doronicum to be one of the most plesiomorphic members of 
the Senecioneae and consider the genus as an outgroup in his cladistic 
studies of the Calenduleae. Comparative DNA studies support the 
distinctive nature of Doronicum as an independent lineage of the 
Asteroideae with affinities to the Calenduleae clade. For this reason, I 
propose the recognition of tribe Doroniceae. 

New subtribes of the Athroismeae 

Athroisminae Panero, subtrib. nov. -Type: Athroisma DC. 

A Athroismeae ceteris, differt characteribus conjuncte: habitu 
herbaceo; foliis altemiis vel fasciculatis, capitulis discoiformis; 
phyllariis absentibus vel 1 -seriatis, receptaculis paleatis; flosculorum 
peripherialum pistillatis, coroUis cylindraceis, flosculi discorum 



Phytologia (Mar 2005) 87(1) 3 



bisexualibus vel fungenter staminatis, lobis (4-) 5; antheris caudatis, 
brevioribus; cypselae compressis, pappi vel squamis. 

Perennial herbs, shrubs or small trees. Leaves alternate or 
fasciculate in brachyblasts, petiolate or sessile, petioles sometimes with 
a basal spine; blades linear, lanceolate to ovate, obovate, venation 
triplinerved or with a single vein. Capitulescence a congested 
glomerule-like solitary cyme or arranged in open to compact, 
paniculiform cymes. Capitula disciform or discoid, homogamous or 
heterogamous, involucre narrowly campanulate, phyllaries 0-2. 
Receptacle paleate. Florets all actinomorphic, female florets present or 
absent, 2 to several, corollas tubular, bisexual florets 2-25, corollas 
campanulate, white, greenish-white, purple or yellow-white, lobes (4-) 
5; anthers (4-) 5, ecalcarate, caudate, thecae white to pale purple, 
appendages sometimes with apical gland; styles divided or filiform and 
undivided, filiform styles associated with functionally staminate florets, 
stigmatic area divided and confluent at style branch apices, sweeping 
hairs present at style apex, or slightly below, rarely present below the 
style branch bifurcation. Cypselae tangentially flattened, ovate to oval, 
obovate to obcordate, elliptic to suborbicular in outline, lateral ridges 
ciliate, black, pappus of fertile floret cypselas either a crown of twin 
trichomes, sometimes with recurved apices or of deeply lacerated and 
fused squamellae, sometimes with two stronger awns at lateral ridge of 
cypselae, pappus of functionally staminate floret ovaries with a 
variously lacerated crown or of a few, free, lacerated scales. Base 
chromosome number, x = 1 

The subtribe contains three genera including Athroisma (12 spp.), 
Blepharispermum Wight ex DC. (15 spp.) and Leucoblepharis Am. (1 
sp.) with most species in eastern Africa, and a few species in western 
Africa, India, Madagascar, southeast Asia, and Indonesia. 

Centipedinae Panero, subtrib nov. -Type: Centipeda Lour. 

A Athroismeae ceteris, differt characteribus conjuncte: habitu 
herbaceo; foliis altemiis, capitulis discoiformis vel radiatis; phyllariis 
1-2 seriatis, receptaculis epaleatis; flosculorum peripherialum 



4 Phytologia (Mar 2005) 87(1) 



pistillatis, corollis zygomorphis, cylindraceis, ligulis brevioribus, lobis 
2-3 vel absentibus; flosculi discorum bisexualibus, corollis 
actinomorphis, lobis 4; antheris caudatis, brevioribus; ramis stylorum 
subtruncatis, brevioribus; cypselae subteretibus, pappi absentibus. 

Annual or perennial herbs. Leaves alternate, sessile, blades 
obovate, variously toothed, rarely entire. Capitulescence sessile to 
shortly pedunculate, axillary, rarely terminal, solitary. Capitula 
disciform or radiate. Involucres campanulate to hemispherical, 
phyllaries in 1-2 series, subequal, herbaceous. Receptacles shallowly 
convex, epaleate. Peripheral/radiate florets in several series, pistillate, 
fertile, sometimes lobes extremely reduced and corolla seemingly 
tubular, corollas creamy white, green, light yellow or purplish. Disc 
florets bisexual, fertile, corollas tetramerous, creamy white, green, light 
yellow or purplish, essentially glabrous with a few glandular trichomes, 
lobes of corollas not vascularized; stamens 4, anthers hyaline, tailed, 
ecalcarate, appendages wanting or if present, minute, oval; styles with 
two vascular strands, style branches with divided stigmatic surfaces, 
concave, urceolate in outline, apices acute, papillose. Cypselas 
subterete, deeply ridged, glabrescent on basal end, cells of distal end 
with raised apical tips, eventually tips elongating and forming biseriate 
trichomes, trichomes increasing in density at apical end forming a 
shallow cup around corolla tube, trichome cells shallowly ridged; 
pappus absent. 

Monotypic, Centipeda contains five species found in Australia, 
southeast Asia, Africa and southern South America. 

The genus Centipeda has been placed in several tribes including 
Anthemideae and Astereae but Bremer (1994), in his excellent account 
of the family, considered the genus difficult to place but maintained it 
in Asteroideae, unassigned to a tribe. Nesom (1994) believed the genus 
to be a member of the Astereae. Comparative studies of the ITS region 
of the nuclear ribosomal DNA by Wagstaff and Breitwieser (2002) 
found strong support for the exclusion of the genus from the Astereae. 
Their studies revealed Centipeda to be an isolated lineage sister to 



Phytologia (Mar 2005) 87(1) 5 



Athroismeae and the Heliantheae alliance. More recent studies (Panero 
et al., in prep.) support the inclusion of Centipeda in the Athroismeae. 

Anisopappinae Panero, subtrib nov. -Type: Anisopappus Hook. 
& Am. 

A Athroismeae ceteris, differt characteribus conjuncte: habitu 
herbaceo; foliis altemiis; capitulis radiates plerumque discoideis; 
phyllariis 1-2 seriatis; receptaculis paleatis plerumque epaleatis; 
flosculi radiorum pistillatis; flosculi discorum bisexualibus, corollas 
actinomorhis plerumque zygomorphis, lobis 5; antheris caudatis, 
brevioribus; cypselae subteretibus, pappi absentibus vel multi squamis. 

Annual or perennial herbs. Leaves alternate, petiolate to 
subsessile, blades linear to ovate sometimes subcordate, simple to 
pinnatifid, triplinerved, sometimes pinnate. Capitulescences terminal, 
solitary, open paniculiform cymes, rarely subumbilliform cymes. 
Capitula radiate, rarely discoid. Involucres campanulate to mostly 
hemispherical. Receptacles convex, rarely shallowly conical, paleate, 
rarely epaleate. Ray florets pistillate, sometimes with staminodes, 
fertile, corollas yellow. Disc florets bisexual, fertile, corollas 5-lobed, 
golden-yellow; stamens 5, anthers yellowish to brown, shallowly 
calcarate or ecalcarate, tailed, appendages ovate; style branches with 
divided stigmatic surfaces, with obtuse papillae. Cypselae subterete, 
edges wanting, epidermis without crystals, pappus of small scales or 
absent. 

Monotypic, Anisopappus contains 40 species found mostly in 
Africa and Madagascar, with one ruderal species extending into 
southeast Asia and China. 

New subtribes of the Heliantheae 

Chromolepidinae Panero, subtribus nov.-Type: Chromolepis 
Benth. 



6 Phytologia (Mar 2005) 87(1) 



A Heliantheae ceteris, differt characteribus conjuncte: habitu 
herbaceo, palustris 

foliis altemiis, petiolatis, laminis integris vel lobatis/runcinatis; 
capitulis ligulatis; phyllariis 2-3 seriatis, dimorphis; receptaculis 
paleatis; flosculis radiorum pistillatis; flosculis discorum bisexualibus, 
coroUis actinomorphis, lobis 5; antheris ecaudatis; ramis stylorum 
sublinearis, deltatis; cypselae radiorum compressis; cypselae discorum 
qudratis; pappi radiorum absentibus, pappi discorum coronniformibus, 
breviorum. 

Rosette-like, perennial, semiaquatic herbs. Leaves alternate, 
petiolate, semi-succulent, ovate to narrowly lanceolate, entire or 
pinnatifid. Capitulescences axillary, solitary. Capitula radiate. 
Involucres campanulate, phyllaries dimorphic in 2-3 series, subequal, 
membranaceous with black markings. Receptacles convex, paleate. 
Ray florets pistillate, fertile, corollas white. Disc florets bisexual, 
fertile, corollas pentamerous, yellow; stamens 5, anthers yellow- 
hyaline sometimes with dark connectives, appendages papillose; styles 
with two vascular strands, style branch with divided stigmatic surfaces, 
apices broadly acute to shallowly deltate. Ray cypselas tangentially 
flattened, triquetrous to weakly quadrate in cross section, glabrous. 
Disc cypselas obpyramidal, quadrate, sparsely pubescent, 
conspicuously smaller than ray cypselas, pappus a crown of minute 
awns and squamellae, absent in ray cypselas. 

Monotypic, Chromolepis contains one species found in vernal 
pools in mid elevation grasslands of central and western Mexico. 

The Chromolepidinae occupy a derived position within the 
Heliantheae, sister to Dugesiinae and collectively sister to Zaluzaniinae 
(Panero et al. 2001). Robinson (1981) placed Chromolepis in the 
Zaluzaniinae because of its fertile ray florets and shallowly quadrartic 
cypselas. The genus is unusual among Mexican sunflowers because of 
its aquatic, rosette-like habit, and conspicuous blackish colorations on 
phyllaries and abaxial side of ray corollas. The heterochromatic 
capitula and dirnorphic involucre of Chromolepis are characteristics not 
seen elsewhere in Zaluzaniinae. 



Phytologia (Mar 2005) 8 7(1) 1 



Dugesiinae Panero, subtribus nov. -Type: Dugesia A. Gray 

A Heliantheae ceteris, differt characteribus conjuncte: habitu 
herbaceo; foliis altemiis, petiolatis, laminis runcinatis, lobatis; capitulis 
ligulatis; phyllariis 2-3 seriatis; receptaculis paleatis; flosculis radiorum 
pistillatis; flosculis discorum fungenter staminatis, corollis 
actinomorphis, lobis 5; antheris ecaudatis; ramis stylorum sublinearis; 
cypselae compressis; pappi radiorum absentibus, plerumque 
coroniformibus breviorum. 

Prostrate, stoloniferous perennial herbs. Leaves alternate, 
petiolate, blades obovate to oval in outline, runcinate to pinnatifid. 
Capitulescences axillary, solitary, or small paniculiform cymes. 
Capitula radiate. Involucres hemispherical, phyllaries in 2-3 series, 
subequal, outermost foliaceous. Receptacles flat to slightly convex, 
paleate. Ray florets pistillate, in two series, fertile, corollas lemon to 
golden-yellow with greenish to black veins on abaxial side, corolla 
apices deeply 2- rarely 3-lobed. Disc florets functionally staminate, 
corollas pentamerous, yellow; stamens 5, anthers black, appendages 
oval to trullate with a visible constriction between appendage and 
thecae, shallowly carinate; style branches of disc florets narrowly 
tapered and papillose, style branches or ray floret spreading with broad, 
divided stigmatic surfaces. Ray cypselas tangentially flattened, 
biconvex, essentially glabrous with a few trichomes on the neck, 
margins with shallowly lacerate wings, pappus absent or of a minute 
crown. 

Monotypic, Dugesia contains one species found in disturbed, 
gravelly or sandy soils, in the dry highlands of eastern-central Mexico. 

The Dugesiinae are sister to Parthenium with which they share 
strongly bifid ray corollas, tangentially flattened ray cypselas, and 
functionally staminate disc florets. More extensive sampling of the 
Heliantheae in the future may reveal subtribe Ambrosiinae to be 
paraphyletic and that Parthenium L. and Parthenice A. Gray should be 
included in the Dugesiinae. 



8 Phytologia (Mar 2005) 87(1) 



Enceliinae Panero, subtribus nov.-Type: Encelia Adans. 

A Heliantheae ceteris, differt characteribus conjuncte: habitu 
herbaceo vel fruticoso, foliis altemiis, petiolatis, laminis integris 
plerumque lobatis/dissectis; capitulis ligulatis plerumque discoideis; 
phyllariis 2-5 seriatis; receptaculis paleatis, paleae deciduis; 
flosculis radiorum plerumque pistillatis; flosculis discorum 
bisexualibus, corollis actinomorphis, lobis 5; antheris ecaudatis; 
ramis stylorum sublinearis; cypselae compressis; pappi absentibus 
vel 1-2 squamis. 

Annual or perennial herbs, shrubs, rarely trees. Leaves alternate, 
rarely opposite, petiolate, rarely sessile, sometimes with resinous 
exudates, blades linear to ovate or trullate, sometimes laciniate, 
pinnately veined or triplinerved, rarely pentanerved. Capitulescences 
terminal, solitary and scapose, or in paniculiform or corymbiform 
cymes. Capitula radiate or discoid. Involucres turbinate, campanulate 
or hemispherical, phyllaries in 2-5 series, subequal, rarely graduated, 
herbaceous, rarely chartaceous sometimes with resinous exudates. 
Receptacles flat to convex, pales deciduous. Ray florets neuter or 
rarely pistillate and sterile, corollas golden-yellow sometimes with 
biseriate trichomes on tube, rarely on limb. Disc florets bisexual, 
fertile, corollas 5-lobed, golden-yellow or purple, mostly without fibers 
embedding the vascular strands, lobes sometimes with thickened cells 
or with glandular or multicelular trichomes on abaxial surfaces of the 
lobes; stamens 5, anthers yellow, brown or black, appendages ovate to 
linear, sometimes with glandular trichomes, endothecium cells 
fusiform, rarely quadrate with 1-3 polar bridges; style branches with 
fused stigmatic surfaces, some species with two stigmatic surfaces that 
fuse slightly above style bifurcation point, densely papillose below, 
apices acute to broadly acute. Cypselae radially flattened, rarely 
thickened or terete, obovate to oblong, densely sericeous, rarely 
glabrous, sometimes with conspicuous wings or corky edges on sides 
and neck, pappus of two slender awns (rarely absent) with or without 
squamellae in between them, rarely of awns fused to a crown 
surrounding the neck. 



Phytologia (Mar 2005) 87(1) 9 



The subtribe contains five genera including Encelia (15 spp.), 
Enceliopsis (A. Gray) A. Nelson (4 spp.), Flourensia A. DC. (33 spp.), 
Geraea Torr. & A. Gray (2 spp.), Helianthella Torr. & A. Gray (8 spp.) 
found mostly in western North and South America. 

The Enceliinae are an isolated lineage of the Heliantheae nestled 
between the Engelmanniinae and the clade containing the Ambrosiinae, 
Chromolepidinae, Dugesiinae, Helianthinae, Spilanthinae, 
Zaluzaniinae, and Zinniinae (Panero et al. 2001). The subtribe shares 
with subtribe Helianthinae sterile ray florets. In addition, the strongly 
flattened and sericeous cypselas, along with the caducous pales of the 
Enceliinae are only seen elsewhere in the Heliantheae in a few 
members of the Helianthineae (e.g., Syncretocarpus S. F. Blake, and 
some species of Viguiera Kunth). 

Spilanthinae Panero, subtribus nov.-Type: Spilanthes Jacq. 

A Heliantheae ceteris, differt characteribus conjuncte: habitu 
herbaceo vel fruticoso; foliis oppositiis, petiolatis plerumque sessilis, 
laminis integris plerumque lobatis; capitulis ligulatis prlerumque 
discoideis; phyllariis 1-5 seriatis; receptaculis paleatis plerumque 
epaleatis, conoideis; flosculis radiorum pistillatis; flosculis 
bisexualibus, corollis actinomorphis, lobis 5 plerumque 4; antheris 
ecaudatis; ramis stylorum sublinearis; cypselae compressis; pappi 
presentibus, 1-3 squamis. 

Erect or decumbent annual or perennial herbs, sometimes rooting 
at the nodes, rarely scandent shrubs. Leaves opposite, petiolate or 
subsessile, blades linear to ovate, sometimes reniform, entire, 
triplinerved. Capitulescences axillary or terminal, solitary, simple to 
congested cymes, peduncles sometimes fistulose. Capitula discoid or 
radiate. Involucres campanulate to hemispherical, phyllaries in 1-5 
series, subequal, rarely dimorphic, mostly herbaceous, sometimes bases 
indurate and coriaceous. Receptacles convex to conical, especially 
with age, mostly paleate, rarely epaleate, pales chartaceous, rarely 
coriaceous. Ray florets pistillate, fertile, corollas purplish, yellow- 
orange or white. Disc florets, bisexual, fertile, corollas pentamerous. 



1 Phytologia (Mar 2005) 87(1) 



rarely tetramerous, purplish, white, or yellow; stamens 5, rarely 4, 
anthers brown or black, appendages ovate or deltate with or without 
glands; style branches with fused stigmatic surfaces, apices acute and 
papillose, without appendages. Ray cypselae tangentially flattened, 
triquetrous, obovoid, sparsely to densely ciliate, trichomes with 
recurved tips with age. Disc cypselae radially flattened, sometimes 
peripheral ones triquetrous, terete, sometimes quadrate and narrowly 
rhombic, rarely square in outline, mostly shallowly winged or 
sometimes conspicuously winged, corky, ciliate, glabrous to sparsely 
pubescent, pappus of a minute crown, a single awn fused to a broad 
ring around the neck of the cypsela, or more commonly of 2-3 slender 
awns as a continuation of the wings, sometimes with squamellae in 
between. 

The subtribe contains five genera including Acmella Rich, ex Pers. 
(30 spp.), Oxycarpha S. F. Blake (1 sp.), Salmea DC. (10 spp.), 
Spilanthes (6 spp.), and Tetranthus Sw. (2-4 spp.). Most species of the 
subtribe are found in the New World tropics, but a few species of 
Acmella and Spilanthes are endemic to the tropical and subtropical 
regions of the Old World. 

The Spilanthinae are sister to the Zinniinae (Panero et al 2001) and 
share with them opposite leaves, the tendency for solitary, long- 
pedunculate capitulescences, and conical receptacles. The Spilanthinae 
differ from the Zinniinae in lacking marcescent ray corollas. 

New subtribes of tribe Millerieae 

Dyscritothamninae Panero, subtribus nov. - 

Type: Dyscritothamnus B. L. Rob. 

A Millerieae ceteris, differt characteribus conjuncte: habitu 
herbaceo vel fruticoso foliis oppositiis plerumque altemiis, petiolatis 
vel sessilis, laminis integris plerumque lobatis/dissectis; capitulis 
ligulatis plerumque discoideis; phyllariis 1-4-seriatis; receptaculis 
paleatis plerumque epaleatis; flosculis radiorum pistillatis; flosculis 
discorum bisexualibus, coroUis actinomorphis, lobis 5; antheris 



Phytologia (Mar 2005) 87(1) 



ecaudatis; ramis stylorum sublinearis; cypselae subteretibus; pappi 
presentibus, setis papporum plumosis. 

Annual or perennial herbs or shrubs. Leaves mostly opposite, 
sometimes alternate, petiolate or sessile, sometimes clasping or 
perfoliate, blades linear to broadly ovate, sometimes trilobed to deeply 
lobed, rarely deltate or acicular, triplinerved, rarely uninerved. 
Capitulescences terminal, solitary and scapose or open paniculiform 
cymes, rarely congested cymes. Capitula radiate, rarely discoid. 
Involucres cylindrical, campanulate or hemispherical, phyllaries in 1-4 
series, subequal, sometimes graduated and dimorphic with inner series 
chartaceous and much longer than the outer herbaceous series, 
sometimes reduced to only 4 fleshy, herbaceous phyllaries enclosing 
florets in a pyramid-like, valvate bud. Receptacles convex to conical, 
paleate, rarely epaleate, pales sometimes chartaceous, hyaline. Ray 
florets fertile, corollas golden-yellow, white, pink or magenta. Disc 
florets bisexual, fertile, corollas 5-lobed, golden-yellow, green, white, 
pink or purplish; stamens 5, anthers yellow or brown, appendages 
ovate; style branches with divided stigmatic surfaces. Cypselae mostly 
terete to shallowly radially flattened or quadrate, glabrous to densely 
sericeous, pappus of multiple bristles or scales, rarely a very reduced 
crown. 

The subtribe contains five genera including Bebbia Greene (2 
spp.), Cymophora B. L. Rob. (5 spp.), Dyscritothamnus (2 spp.), 
Tetragonotheca L. (4 spp.), and Tridax L. (30 spp.). Most species are 
distributed in montane, tropical North America with a few species in 
the north and central Andes of South America. 

The Dyscritothamninae are sister to the Melampodiinae (Panero et 
al. 2001). The monophyly of the Dyscritothamninae and their subtribal 
relationships are well supported. 

Jaegeriinae Panero, subtribus nov.-Type: Jaegeria Kunth 

A Millerieae ceteris, differt characteribus conjuncte: habitu 
herbaceo, terrestris, palustris vel aquaticus; foliis oppositiis, petiolatis 



12 Phytologia (Mar 2005) 87(1) 



vel sessilis, laminis integris; capitulis ligulatis; phyllariis 1- seriatis; 
receptaculis paleatis; flosculis radiorum pistillatis; flosculis discorum 
bisexualibus, coroUis actinomorphis, lobis 5; antheris ecaudatis; ramis 
stylorum sublinearis; cypselae teretibus; pappi absentibus plerumque 
coroniformibus breviorum. 

Annual or perennial herbs, rooting at the nodes. Leaves opposite, 
sessile to petiolate, blades linear to ovate. Capitulescences axillary or 
terminal, solitary or of open paniculiform cymes. Capitula radiate. 
Involucres campanulate to hemispherical, phyllaries in 1 series, 
subequal, herbaceous with hyaline, ciliate wings wrapping around the 
ray cypselae, caducous and shed along with cypsela as a unit 
(perigynia). Receptacles strongly convex to conical, paleate. Ray 
florets pistillate, fertile, corollas white, white-pink, white-purple, 
yellow, marcescent, tube short or absent. Disc florets bisexual, fertile, 
corollas yellow or yellow-green, sometimes whitish gray, lobes 5 rarely 
4, tube abruptly narrowed into the throat; stamens 5, anthers yellow, 
appendages ovate; style branches recurved, apices round to obtuse. 
Cypselae terete to obscurely angled, clavate, bases tapered, glabrous, 
shiny black, ray cypselas slightly larger than disc cypselas otherwise 
equivalent; pappus absent or a minute crown. Base chromosome 
number, x = 9. 

Monotypic, Jaegeria contains nine species found in the New 
World tropics with most species concentrated in the highlands of 
Mexico. 

The Jaegeriinae are sister to the clade containing Milleriinae, 
Desmanthodiinae, Espeletiinae, and Galinsoginae (Panero et al. 2001). 

New Combinations in subtribe Ambrosiinae. 

Studies of Heliantheae subtribe Ambrosiinae by Miao et al. (1995) 
support the expansion of the genus Euphrosyne DC. to include five 
species. This new concept of Euphrosyne requires the creation of the 
following combinations: 



Phytologia (Mar 2005) 87(1) 1 3 



Euphrosyne acerosa (Nutt.) Panero, comb. nov. Oxytenia acerosa 
Nutt., J. Acad. Phil. N. S. 1:172. 1847. 

Euphrosyne dealbata (A. Gray) Panero, comb. nov. Iva dealbata A. 
Gray, PI. Wright. 1:104. 1852. 

Euphrosyne nevadensis (M. E. Jones) Panero, comb. nov. Iva 

nevadensis M. E. Jones, Amer. Natur. 17: 973. 1883. 

ACKNOWLEDGMENTS 

I wish to thank Billie L. Turner and Blanca Leon for helpful 
comments and improving the manuscript. 

LITERATURE CITED 

Alvarez Fernandez, I. 2003. Systematics of Eurasian and North 
African Doronicum (Asteraceae: Senecioneae). Ann. Missouri 
Bot.Gard. 90:319-389. 

Bremer, K. 1994. Asteraceae: cladistics and classification. Timber 
Press, Portland, pp. 1-752. 

Miao, B., B. L. Turner, and T. J. Mabry. 1995. Systematic 
implications of chloroplast DNA variation in the subtribe 
Ambrosiinae (Asteraceae: Heliantheae). Amer. J. Bot. 82: 924- 
932. 

Nesom, G. L. 1994. Subtribal classification of the Astereae 
(Asteraceae). Phytologia 76:193-274. 

Nordenstam, B. 1977. Senecioneae and Liabeae-systematic review. 
In: Heywood, V. H., J. B. Harbome, B. L. Turner (eds), The 
Biology and Chemistry of the Compositae 2. Academic Press. 
London, pp. 799-830. 



14 Phytologia (Mar 2005) 87(1) 



Nordenstam, B. 1994. Tribe Calenduleae. In: Bremer K. (ed.) 
Asteraceae cladistics and classification, Timber Press. Portland, 
Oregon, pp. 365-376. 

Panero, J. L., B. G. Baldwin, E. E. Schilling, and J. A. Clevinger. 

2001. Molecular phylogenetic studies of members of tribes 
Helenieae, Heliantheae, and Eupatorieae (Asteraceae) 3. General 
systematics and proposed taxonomic changes in current 
classification. Abstracts of the AIBS 2001 meeting, Albuquerque, 
New Mexico, p. 132. 

Panero, J. L. and V. A. Funk. 2002. Toward a phylogenetic 
subfamilial classification for the Compositae (Asteraceae). Proc. 
Biol. Soc. Washington 115: 909-922. 

Robinson, H. 1981. A revision of the tribal and subtribal limits of the 
Heliantheae (Asteraceae). Smithsonian Contr. Bot. 51: 1-102. 

Wagstaff, S. J. and I. Breitwieser. 2002. Phylogenetic relationships 
of New Zealand Asteraceae inferred from ITS sequences. PI. Syst. 
Evol. 231: 203-224. 



Phytologia (Mar 2005) 8 7(1) 1 5 



NEW GENERA AND COMBINATIONS IN BRYACEAE 
(BRYALES, MUSCI) FOR NORTH AMERICA 

John R. Spence 

National Park Service, Glen Canyon National Recreation Area, P.O. 

Box 1507, Page, AZ 86040-1507, U.S.A. Email: 

John_Spence@nps.gov 

ABSTRACT 

The genus Ptychostomum Homsch. is resurrected for species of 
Bryum in sections Amblyophyllum and Caespitibryum, with 32 well 
established species found in North America transferred. The section 
Leptostomopsis of Brachymenium is raised to generic rank, with the 
one species found in North American transferred. The genus 
Plagiobryoides is newly described for the neotropical Bryum 
incrassatolimbatum. Two species of Mielichhoferia in North America, 
M. macrocarpa and M. tehamensis, are transferred to the neotropical 
genus Haplodontium. 

Key words: mosses. North America, Bryaceae, Haplodontium, 
Plagiobryoides, Ptychostomum 

Introduction 

A considerable amount of new research has been published 
recently on the Bryaceae, and in particular the genus Bryum (Cox & 
Hedderson 1998, 2003; Cox et al 2000; Pedersen 2002; Pedersen & 
Hedenas 2002; Pedersen et al 2003; Spence 1987, 1996; Spence & 
Ramsay 1996, 1999, 2002). Bryum is an extremely large and diverse 
genus, united primarily by plesiomorphic sporophytic characters found 
throughout the Bryalean altemate-diplolepidous taxa (cf Pedersen et 
al 2003). Although there is considerable morphological variation 
within the genus, no attempts have been made to divide Bryum into 
smaller and presumably more natural groups. To date, the various 
phylogenetic studies using chloroplast DNA sequences and 



16 Phytologia (Mar 2005) 87(1) 



morphology have not converged on a particular taxonomic solution, 
other than indicating that Bryum and the related genus Brachymenium 
are polyphyletic. 

Although trying to understand the relationships among various 
proposed genera, species and sections of Bryum s.l. seems at times to 
be hopeless, I believe that it is possible to define natural groups within 
the genus based on a combination of characters from both generations. 
Dr. Helen Ramsay and I, while monographing Bryum for the Flora of 
Australia (Spence & Ramsay 2005), have identified groups among the 
gametophytic diversity in the genus. We have focused primarily on 
characters of the gametophyte generation, as all work to date suggests 
that the capsule orientation and peristome features, traditionally used to 
delimit genera in the family, are not indicative of evolutionary 
relationships above the species or perhaps sectional levels. We feel that 
restricting analysis to primarily gametophyte characters provides a 
valuable and relatively stable basis for the delimitation of natural 
groups (genera) in the Bryaceae (c/ Zander 1993 for an analogous 
situation in the Pottiaceae). There are several implications of our work 
and other studies based on DNA characters, which are discussed below. 

Our work clearly supports the presence of two major lineages 
within Bryum, in addition to several smaller groups. Interestingly, 
these two major groups receive at least partial support from the other 
DNA and morphological studies cited above {cf. Pedersen et al. 2003). 
One lineage includes the many traditional Bryum species first grouped 
as the informal "Cemuibryum" by one of us (Spence 1987), along with 
Acidodontium, Brachymenium section Brachymenium, Rhodobryum 
and Rosulabryum. The second lineage includes the numerous small 
gemmiferous species of Bryum in sections Bryum, Apalodictyon, 
Alpiniformia, and Doliolidium, along with Anomobryum and 
Brachymenium sect. Dicranobryum. We have pointed out that true 
Bryum {e.g., Bryum argenteum) is closely related morphologically to 
Anomobryum, and could easily be accomodated in that genus (Spence 
& Ramsay 2002). However, E.G. Britton lectotypified the genus 
Bryum with B. argenteum, which we regard as an unfortunate choice 
(Spence & Ramsay 1999). Anomobryum, and the type of Bryum are 



Phytologia (Mar 2005) 87(1) 1 7 



sufficiently distinct from other members of the genus to warrant 
generic status. There are two potential taxonomic implications of this 
lectotypification if revisions are attempted. The first is that B. 
argenteum and its allies could be treated as a genus closely related to 
Anomobryum, with all other Bryum species given a new name. The 
second is that the two taxa might be combined, in which case a new 
name would still have to be found for all other Bryum species. For 
Australia, we chose the latter of the two (Spence & Ramsay 2002). We 
also argued for the conservation of the name Bryum with a new type, 
for which we selected the first species in Hedwig, B. caespiticium 
(Spence & Ramsay 1999). However, the Bryophyte Committee on 
Nomenclature rejected our proposal (Zijistra 2002), based in part on a 
concern for the use of Bryum caespiticium as the conserved type, and 
also because further studies on Bryum were pending. Some of these 
presumed studies have now been either published {e.g., Pedersen et al. 
2003) or are cited in Pedersen (2002), and as noted above they do not 
agree with each. This leaves two choices, continue to treat Bryum in a 
very broad sense, knowing full well that it is polyphyletic, as Ochi 
(1992) chose to do, or come up with a new classification of the genus 
based on the evidence to date from all studies, both genetic and 
morphological. The first approach is unacceptable, so this paper as 
well as the treatment in Spence & Ramsay (2005) represents the first 
attempt since Fleischer and Brotherus (1925) to generically re-classify 
Bryum 

Perhaps not surprisingly, the 1 9^^ century also witnessed confusion 
with the exact delimitation and nature of Bryum. Hedwig (1801) 
included a variety of species in his concept of the genus, many of 
which now reside elsewhere. Fairly soon thereafter both Homschuch 
(1822) and Bridel (1826) described new genera long since 
synonymized under Bryum. In Homschuch 's case, it was 
Ptychostomum, described for several species now residing in Bryum, 
including B. algovicum and B. uliginosum. Bridel published 
Cladodium for the species B. marratii and B. inclinatum (^B. 
amblyodon). Most of the other species they included in these new 
genera have since been transferred to Pohlia, Mnium and other genera. 



1 8 Phytologia (Mar 2005) 87(1) 



Validly Published Name 


Year 


Current Name 


C. calophyllum (R. Br.) Brid. 


1826 


Bryum calophyllum 


C. demissum (Hook.) Nees 


1836 


Plagiobryum demissum 


*C. inclinatum (Sw. ex Brid.) Brid. 


1826 


Bryum amblyodon 


C. rhamphostegium Hampe 


1865 


Acidodontium rhamphostegium 


C. socorrense Hampe 


1869 


Bryum limbatum 


C. uliginosum Brid. 


1827 


Bryum uliginosum 



Table 1 . Summary of validly published species of Cladodium Bridel, 
year of publication, and currently accepted name. The lectotype species 
is indicated by an asterisk. 

For some reason, these generic names never caught on with later 
bryologists, and have languished under Bryum ever since, probably 
because the first species of Bryum in Hedwig (1801), 5. caespiticium, 
is morphologically similar to both B. algovicum and B. amblyodon. All 
those species still remaining in Bryum that were placed by these two in 
Cladodium and Ptychostomum are in the informal group 
"Cemuibryum" while B. argenteum is in the group "Bryum" of Spence 
(1987). 

Because of the above considerations, I propose here to re-instate 
the genus Ptychostomum Homschuch for those species of Bryum in the 
informal group "Cemuibryum" of Spence (1987). Since all three 
species first cited by both Bridel and Homschuch are related, either 
genus could be used, but Ptychostomum has priority over Cladodium 
by four years. Interestingly, my work also suggests two closely related 
groups within Ptychostomum, one corresponding to the species B. 
algovicum, and the second to those species related to B. uliginosum. 
For now I prefer to keep these two groups together, although future 
work may indicate that they represent distinct but closely related 
genera. The studies of Cox and Hedderson (2001) and Pedersen et al. 
(2003) both show a well supported clade that consists of species in 
Ptychostomum, although including the species Bryum donianum, which 
based on its gamteophyte may be a Rosulabryum. 



Phytologia (Mar 2005) 87(1) 19 



Validly Published Name 


Year 


Current Name 


P. caespiticium Brid. 


1827 


Bryum algovicum 


♦P. cernuum (Hedw.) Homsch. 


1822 


Bryum uliginosum 


P. compactum Homsch. 


1822 


Bryum algovicum 


P. pendulum Homsch. 


1822 


Bryum algovicum 


P. puchellum R. Br. 


1823 


Bryum algovicum 


P. radiculosum Brid. 


1826 


Bryum uliginosum 



Table 2. Summary of validly published species of Ptychostomum 
Homsch., year of publication, and currently accepted name. The 
lectotype species is indicated by an asterisk. 

Tables 1 and 2 list all validly published species in Cladodium and 
Ptychostomum. Pfeiffer (1873) lectotypified Cladodium with the 
species C. inclinatum, but Ptychostomum has not yet been 
lectotypified. Homschuch's understanding of the genus was based on 
certain characters of the sporophyte, primarily the peristome. His 
essential characters included "Peristomium duplex: exterius dentibus 
sedecim erectis, interius membrana hyalia plicata, peristomio externo 
adhaerens et denta ejusdem inter se conjungens ". The adherence of 
the endostome to the exostome is a critical character in the 
circumscription of the genus. The two species Homschuch included in 
Ptychostomum are currently known as Bryum algovicum and B. 
uliginosum. Both these species are characterized by reduced peristomes 
with the endostome adherent to the exostome. I have selected P. 
cernuum {=B. uliginosum) as the lectotype because P. pendulum (^B. 
algovicum) is closely related to the type of Cladodium, C inclinatum. 
This selection was made in the interests of preserving both generic 
names, in case Cladodium is also recognized at the generic level at 
some point in the future. Ptychostomum cernuum is in a different 
section than C. inclinatum and is not closely related to it. 

Ptychostomum Homsch., Flora 5, 2: syll. 62, 1822. Lectotype 
species: Ptychostomum cernuum Homsch., Flora 5, 2: syll. 64, 1822. 
In addition, Homschuch included the species currently known as 
Bryum algovicum (Brid.) B.S.G. in the genus as Ptychostomum 
pendulum Homsch., Flora 5, 2: syll 62, 1822. 



20 Phytologia (Mar 2005) 87(1) 



The following 27 species found in North America are also 
transferred to the genus. 

Ptychostomum archangelicum (B.S.G.) J.R. Spence, comb. nov. 
Basionym: Bryum archangelicum B.S.G., Bryol. Eur. 4: 153, 
1846. 

Ptychostomum arcticum (R. Br.) J.R. Spence, comb. nov. 

Basionym: Pohlia arctica R. Br., Chlor. Melvill. 38. 1823. 

Ptychostomum badium (Brid.) J.R. Spence, comb. nov. 

Basionym: Bryum caespiticium var. badium Brid., Bryol. Univ. 1 

850. 1827. 

Ptychostomum bimum (Schreb.) J.R. Spence, comb. nov. 

Basionym: Mnium bimum Schreb., Bot. Zeit. (Regensburg) 1: 79. 
1802. 

Ptychostomum calophyllum (R. Br.) J.R. Spence, comb. nov. 

Basionym: Bryum calophyllum R. Br., Chlor. Melvill. 38. 1823. 
Synonym: Cladodium calophyllum (R. Br.) Brid., Bryol. Univ. 1: 
620. 1826. 

Ptychostomum cryophilum (Mart.) J.R. Spence, comb. nov. 

Basionym: Bryum cryophilum Mart., K.V.A. Afh. Natursk. 15: 
183, 1956. Nom. nov. B. obtusifolium Lindb., Ofvers. Forh. 
Konsch. Svenka Vetensk.-Akad. 23: 544. 1866; not 5. 
obtusifolium Brid., Muscol. Recent 2(3): 52. 1803. 

Ptychostomum curvatum (Kaur. & Am.) J.R. Spence, comb. nov. 
Basionym: Bryum curvatum Kaur. & Am., Bot. Not. 67. 1897. 

Ptychostomum cyclophyllum (Schwaegr.) J.R. Spence, comb. nov. 
Basionym: Mnium cyclophyllum Schwaegr. Sp. Muse. Suppl. 2, 
2(2): 160, pi. 194. 1827. 



{ 



Phytologia (Mar 2005) 87(1) 2 1 



Ptychostomum inclinatum (C. Muell.) J.R. Spence, comb. nov. 
Basionym: Cladodium inclinatum (Sw. ex Brid.) Brid., Bryol. 
Univ. 1:620, 1826. 

Ptychostomum intermedium (Brid.) J.R. Spence, comb. nov. 

Basionym: Pohlia intermedia Brid., Muscol. Recent 2(3): 144, pi. 
2,f. 12. 1803. 

Ptychostomum knowltonii (Barnes) J.R. Spence, comb. nov. 

Basionym: Bryum knowltonii Barnes, Bot. Gaz. 14: 44, 1889. 

Ptychostomum lonchocaulon (C. Muell.) J.R. Spence, comb. nov. 
Basionym: Bryum lonchocaulon C. Muell., Flora 2(6): 90, 1819. 

Ptychostomum longisetum (Bland, ex Schwaegr.) J.R. Spence, comb, 
nov. Basionym: Bryum longisetum Bland, ex Schwaegr., Spec. 
Frond. Muse. Suppl. 1, 2: 105, pi. 74, 1816. 

Ptychostomum marratii (Hook. & Wils.) J.R. Spence, comb. nov. 
Basionym: Bryum marratii Hook. & Wils., Bryol. Brit., p. xi 
(add.), pi. XXXIIb, 1855. 

Ptychostomum meesioides (Kindb.) J.R. Spence, comb. nov. 

Basionym: Bryum meesioides Kindb., Bull. Torrey Bot. Club 16: 
95, 1889. 

Ptychostomum neodamense (Itzigs.) J.R. Spence, comb. nov. 

Basionym: Bryum neodamense Itzigs. in C. Muell., Syn. Muse. 
Frond. 1:258. 1848. 

Ptychostomum pallens (Sw.) J.R. Spence, comb. nov. 

Basionym: Bryum pallens Sw., Monthl. Rev. Lond. 34: 538. 1801. 

Ptychostomum pallescens (Schleich. ex Schwaegr.) J.R. Spence, comb, 
nov. Basionym: Bryum pallescens Schleich. ex Schwaegr., Sp. 
Muse. Suppl. 1, 2: 107, pi. 75. 1816. 



22 Phytologia (Mar 2005) 87(1) 



\ 



Ptychostomum purpurascens (R. Br.) J.R. Spence, comb. nov. 

Basionym: Pohlia purpurascens R. Br., Clor. Melvill. 39. 1823. 

Ptychostomum reedii (Robins.) J.R. Spence, comb. nov. f 

Basionym: Bryum reedii Robins., Bryol. 69: 107, 1966. 

Ptychostomum rutilans (Brid.) J.R. Spence, comb. nov. 

Basionym: Bryum rutilans Brid., Bryol. Univ. 1: 684. 1826. 

Ptychostomum salinum (Hag. ex Limpr.) J.R. Spence, comb. nov. 
Basionym: Bryum salinum Hag. ex Limpr. Laubm. Deutschl. 2: 
334. 1892. 

Ptychostomum schleicheri (Schwaegr.) J.R. Spence, comb. nov. 

Basionym: Bryum schleicheri Schwaegr., Sp. Muse. Frond. Suppl. 
1,2: 113,pl. 73p.p. 1816. 

Ptychostomum turbinatum (Hedw.) J.R. Spence, comb. nov. 

Basionym: Mnium turbinatum Hedw., Sp. Muse. Frond. 191. 1801. 

Ptychostomum warneum (Rohl.) J.R. Spence, comb. nov. 

Basionym: Mnium caespiticium var. warneum Rohl., Deutsch. Fl. 
(ed. 2), Kryptog. Gew. 3: 95. 1813. 

Ptychostomum weigelii (Spreng.) J.R. Spence, comb. nov. 

Basionym: Bryum weigelii Spreng., Mant. Prim. Fl. Hals. 55. 
1807. 

Ptychostomum wrightii (SuU. & Lesq.) J.R. Spence, comb. nov. 

Basionym: Bryum wrightii Sull. & Lesq., Proc. Amer. Acad. Arts 
Sci.4:278. 1860. 

Several additional species of Bryum that may belong in 
Ptychostomum have been reported from North America. Most are 
poorly known Arctic species, and more work is needed to determine 
their status. These are: B. acutiforme Limpr., B. axel-blyttii Philib., B. 



Phytologia (Mar 2005) 87(1) 23 



bryoides (R. Br.) Angstr., B. brachyneuron Kindb., B. hagenii Limpr., 
B. nitidulum Lindb., B. subneodamense Kindb., and B. teres Lindb. 

The following three species of Bryum also occur in Australia, and 
are transferred for the Flora of Australia (Spence & Ramsay 2005). 

Ptychostomum angustifolium (Brid.) J.R. Spence & H.P. Ramsay, 
comb. nov. Basionym: Bryum angustifolium Brid., Muse. Recent. 
Suppl. 3: 31, 1817. Synonym: Bryum caespiticium Hedw., Sp. 
Muse. Frond., 1801, not Ptychostomum caespiticium Brid., Bryol. 
Univ. 1:837, 1827. 

Since the combination Ptychostomum caespiticium has priority as 
a synonym of Ptychostomum pendulum a new name is needed for the 
species known as B. caespiticium Hedw. The first name that still 
remains valid within that species is B. angustifolium Bridel. 

Ptychostomum creberrimum (Taylor) J.R. Spence & H.P. Ramsay, 
comb. nov. Basionym: Bryum creberrimum Taylor, Lond. J. Bot. 
5:54. 1846. 

Ptychostomum pseudotriquetrum (Hedw.) J.R. Spence & H.P. Ramsay, 
comb. nov. Basionym: Mnium pseudotriquetrum (Hedw.) 
Schwaegr., Spec. Muse. Suppl. 1, 2: 110. 1816. 

Leptostomopsis (C. Muell.) J.R. Spence & H.P. Ramsay, stat. nov. 
Lectotype: L. systylium (C. Muell.) J.R. Spence & H.P. Ramsay, 
comb. nov. Basionym: Bryum systylium C. Muell., Syn. Muse. 
Frond. 1:320. 1848. 

The species in section Leptostomopsis of Brachymenium comprise 
a distinctive group that superficially resembles Leptostomum. They are 
characterized by densely compact cushions on bark or rock, dense 
lamina areolation, a long hyaline spinose hairpoint, and erect capsules 
with reduced peristomes. In an earlier paper (Spence 1996) I first 
suggested that the section may represent a distinct genus. The work of 
Cox & Hedderson (2003) and Pedersen et al. (2003) support this. 



24 Phytologia (Mar 2005) 87(1) 



suggesting that the group is basal to the remainder of the Bryaceae, and 
only distantly related to the rest of Brachymenium as well as Bryum. 
This section is sufficiently distinct from other members of the Bryaceae 
in both genetic and morphological characters to warrant generic rank. 
There are about 8-10 species, pantropical and southern hemispheric in 
distribution in drier tropical and subtropical regions. One species, the 
type, is known from southern areas of the U.S.A. 

Plagiobryoides J.R. Spence, gen. nov. 

Caules uniformiter foliati. Folia ovata, in statu sicco contorta, 
areolatione laminali subheterogenea, cellulis distalibus brevibus 
irregulariter subquadratis vel brevi-rhombicis, in parte mediana atque 
proximali cellulis paulo longi-rhombicis, marginibus distincte limbatis 
valde bi-vel multistratosis, costa in sectione transversali cellulis ducum 
in strato unico adaxialiter supra stratum stereidarum bene effectum sitis 
praedita. Gemmae asexuales nullae. Plantae dioicae. Capsulae 
subzygomorphae, ore obliquo, inclinatae vel subrectae; peristomium 
diplolepidum, segmentis endostomialibus longitudine acquis 
exostomio, ciliis imperfectis vel nuUis. Sporae parvae, 10-15 um. 

Stems elongate, to 30 mm, simple, rarely branched, in transverse 
section with weakly developed central strand, with large thin-walled 
cells exterior, and somewhat smaller and thicker-walled peripheral 
cells; evenly foliate; rhizoids abundant, red-brown, papillose. Leaves 
pink, red to brown, ovate, keeled, contorted and twisted when dry, 
spreading when wet, concave, 1.0-2.5 mm long, apex acute to obtuse, 
moderately to strongly and longly decurrent; costa broad, strong, not 
reaching apex to rarely percurrent, in cross-section with enlarged 
guide-cell like layer, one to two-layered, ventral to a well developed 
stereid band, sometimes guide-like cells lacking; lamina areolation 
somewhat heterogenous, thin-walled, with short irregularly 
subquadrate to short-rhomboidal cells distally and somewhat longer 
irregularly rhomboidal cells in median and proximal part, typically, 60- 
100 fxm X 12-30 |im wide, margins with strongly differentiated border, 
bi- to multistratose, of thick-walled cells. Specialized asexual 
gemmae lacking. Sexual condition dioicous. Seta slender, flexuose to 



Phytologia (Mar 2005) 87(1) 25 



bent near base. Capsules suberect, somewhat zygomorphic and 
asymmetric with oblique mouth, apophysis slender, well differentiated, 
operculum conic, without apiculus, peristome double, exostome teeth 
lanceolate, somewhat united at base, yellowish, basal membrane 
hyaline, high, segments slightly keeled, with narrow perforations or 
slits, the same length as exostome, cilia rudimentary or lacking. Spores 
small, 11-14 um (description of sporophyte from Ochi and Salazar- 
Allen 1990). 

Type species: Plagiobryoides incrassatolimbata (Card.) J.R. 
Spence, comb. nov. Basionym: Bryum incrassatolimbatum Card., 
Rev. Bryol. 36: 114, 1909. 

Plagiobryoides incrassatolimbata is a distinctive species that 
cannot be confused with any other member of the Bryaceae. The 
extremely lax lamina areolation and often multistratose limbidium are 
diagnostic. Remarkable for the family, lamina cells near the leaf tip are 
often nearly isodiametric to subquadrate. The capsule is similar to that 
found in Plagiobryum and its allies in the broad sense, including Bryum 
cellulare and related species. This distinctive plant is found on wet 
calcareous rock and is a Neotropical species, distributed from the 
southwestern U.S.A. through Mexico and Central America. 

Haplodontium Hampe 

Haplodontium consists of species with lateral sporophytes and an 
extremely reduced peristome, typically of one layer. Shaw (1985) 
transferred all species in the genus to Mielichhoferia Homsch. 
However, the recent work using DNA analysis (Cox and Hedderson 
2003; Cox et_al. 2000; Pedersen 2002; Pedersen and Hedenas 2002; 
Pedersen et al . 2003) indicates that the type of Mielichhoferia, M. 
mielichhoferiana, is more closely related to Pohlia, while many other 
species originally in Haplodontium are deeply nested with the 
Bryaceae. These include the two North American species M 
macrocarpa and M. tehamensis, which are gametophytically very close 
to other species oi Haplodontium and Plagiobryum, and very similar to 



26 Phytologia (Mar 2005) 87(1) 



the type of the genus, H. megalocarpum Amott. These two species are 
thus transferred to a resurrected Haplodontium as: 

Haplodontium macrocarpum (Hook, ex Drumm.) J.R. Spence, comb, 
nov. Basionym: Weissia macrocarpa Hook, ex Drumm. Musci 
Amer., Brit. N. Amer. 74. 1828. 

Haplodontium tehamense (Showers) J.R. Spence, comb. nov. 

Basionym: Mielichhoferia tehamensis Showers, Bryol. 83: 365, 
1980. 

Acknowledgments 

This work was supported in part by a grant from the Flora of North 
America Association. The Missouri Botanical Gardens provided office 
space. Thanks are due to Richard Zander, Marshall Crosby and Bruce 
Allen for their support during my stay. I am grateful to Patricia Eckel 
for providing the Latin diagnoses. Marshall Crosby and Richard 
Zander kindly reviewed an early draft of this work. 

Literature Cited 

Bridel, S.E. 1 826. Bryologia Universa. Vol. 1 . Leipzig. 

Brotherus, V.F. 1925. Musci (Laubmoose). 2. Halfte. In: Engler, A & 
K. Prantl (eds), Die Natiirlichen Pflanzenfamilien. Leipzig. 

Cox, C.J. and T.A.J. Hedderson. 1998. Phylogenetic relationships 
among the ciliate arthrodontous mosses: evidence from chloroplast 
and nuclear DNA sequences. Plant Syst. Evol. 215: 1 19-139. 

. 2003. Phylogenetic relationships within the moss family 

Bryaceae based on chloroplast DNA evidence. J. Bryol. 25: 31-40. 



Phytologia (Mar 2005) 87(1) 27 



Cox, C.J., B. Goffinet, A.E. Newton, A.J. Shaw and T.A.J. 
Hedderson. 2000. Phylogenetic relationships among the 
diplolepidous-ahemate mosses (Bryidae) inferred from nuclear and 
chloroplast DNA sequences. Bryol. 103: 224-241. 

Hedwig, J. 1801. Species muscorum frondosorum descriptae. Leipzig. 

Hornschuch, F. 1826. Flora 5, 2: Syll. 62. 1822. 

Ochi, H. 1992. A revised infrageneric classification of the genus 
Bryum and related genera (Bryaceae, Musci). Bryobrothera 1: 231- 
244. 

Ochi, H. and N. Salazar- Allen. 1990. Bryum incrassatolimbatum new 
to Panama, with first description of its sporophyte. Bryol. 93: 286- 
287. 

Pedersen, N. 2000. A cladistic overview of the Bryaceae (Musci) 
based on morphological and anatomical data and with emphasis on 
the genus Bryum. J. Bryol. 22: 193-206. 

Pedersen, N. 2002. Phylogeny and taxonomy of the acrocarpous moss 
family Bryaceae with emphasis on the genus Bryum Hedwig. 
Botaniska Institutionen, Stockholms Universitet, Stockholm, 
Sweden. 

Pedersen, N. and L. Hedenas. 2002. Phylogenetic relationships 
between Bryum and supposedly related genera. J. Bryol. 24: 277- 
289. 

Pfeiffer, L. 1873. Nomenclatur Botanicus. 1(1). Cassellis, sumptibus 
Theodori Fischeri, vi + 808 pp. 

Pedersen, N., C.J. Cox, and L. Hedenas. 2003. Phylogeny of the moss 
family Bryaceae inferred from chloroplast DNA sequences and 
morphology. Syst. Bot. 28: 471-482. 



28 Phytologia (Mar 2005) 87(1) 



Shaw, A.J. 1985. Nomenclatural changes in the Bryaceae subfamily 
Mielichhoferioideae. Bryol. 88: 28-30. 

Spence, J.R. 1987. A proposed reclassification of Bryum, 
Anomobryum, and Brachymenium (Musci, Bryaceae). J. Bryol. 14: 
659-676. 

Spence, J.R. 1996. Rosulabryum genus novum (Bryaceae). Bryol. 99: 

221-225. 

Spence, J.R. and H.P. Ramsay. 1996. New and interesting species of 
Bryum from Australia. J. Adelaide Bot. Gard. 17: 107-1 18. 

Spence, J.R. and H.P. Ramsay. 1999. Proposal to conserve 
the name Bryum (Musci, Bryaceae) with a conserved 
type. Taxon48: 827-828. 

Spence, J.R. and H.P. Ramsay. 2002. The genus Anomobryum 
Schimp. (Bryopsida, Bryaceae) in Australia. Telopea 9: 777-792. 

Spence, J.R. and H.P. Ramsay. 2005. Bryaceae. Flora of Australia, 
Vol. 5 1 . Government Printing Service, Canberra (in press). 

Zander, R.H. 1993. Genera of the Pottiaceae: Mosses of harsh 
environments. Bull. Buffalo Soc. Nat. Sci. 32: 1-378. 

Zijistra, G. (sec). 2002. Report of the committee for bryophytes: 7. 
Taxon 51: 793-794. 



Phytologia (Mar 2005) 87(1) 29 



KEYS TO THE FLORA OF FLORIDA - 12, RUBUS 
(ROSACEAE) 

Daniel B. Ward 

Department of Botany, University of Florida 
Gainesville, Florida 3261 1, U.S.A. 

ABSTRACT 

The genus Rubus (Rosaceae) is represented in Florida by 4 species: 
R. flagellaris Willd. is found in the state only in the Florida panhandle; 
R. argutus Link, R. cuneifolius Pursh, and R. trivialis Michx., are 
widely distributed, although rare or absent in South Florida. Specific 
names based upon 1 5 Florida Rubus types are assigned to these taxa. 
An extended commentary is provided in support of the thesis that the 
great majority of specific names in Rubus have been given to 
agamospermic segregates that, although morphologically 
distinguishable by the specialist, are too subtly characterized to be 
usefiilly employed by the working taxonomist, and that a less precise 
classificatory structure centered upon the sections into which the genus 
is divisible, is to be preferred. An amplified key is given to the Florida 
species. 

KEY WORDS: Rubus, Rosaceae, Florida flora. 



"What is the blackberry situation at this hour? It is 
indeed an unhappy heritage. Where angels had feared to tread 
the ground has been traversed, and so unforbearingly, 
notwithstanding the briers, that not any semblance of a 
pathway has been suffered to exist." E. P. Bicknell (Bull. 
Torrey Bot. Club 37:393. 1910) 



Alongside Crataegus, Rubus (Rosaceae) shares the unenviable 
reputation of being among the most intractable of North American 



30 Phytologia (Mar 2005) 87(1) 



genera in terms of the ease by which a specimen or even an entire plant 
may be assigned to its correct species. The muhiplicity of specific 
epithets found in floristic works, together with the subtle 
morphological differences by which the various entities are separated, 
has presented the user with an excessive number of alternatives and 
little prospect that his choice will be correct. Reproducibility ~ the 
bedrock of scientific enquiry ~ in this case the probability that 
independent observers of competent background will assign the same 
name to a given Rubus clone, has been replaced by a quicksand of 
individual interpretation. 

This imprecision of identification has produced in the minds of 
many workers distaste for the genus Rubus and a disinclination to 
spend time and attention in the resolution of its problems. The hiatus 
has been filled by a few dedicated and exceedingly patient students 
whose very expertise has resulted in the recognition of ever more taxa 
and a further widening of the barrier to conventional enquiry. 

A casual observer whose interests lie primarily in the identification 
of plants of a limited area may not fully appreciate the explosive 
proliferation in the names assigned to Rubus in eastern North America. 
Until the last years of the 1 9th century less than a dozen names were of 
common usage, and included the wholly distinct raspberries, flowering 
raspberries, and cloud-berries. The blackberries, where most of the 
taxonomic difficulties lie, received no serious examination until W. H. 
Blanchard studied them in the field from Newfoundland to Florida and 
west to the geographic limit of the group. By 1911, although 
Blanchard had concluded that "eight species include the great bulk of 
our blackberries, perhaps ninety percent of them," he felt obligated to 
recognize and name 36 additional specific entities (Bailey, Gentes 
Herbarum 1:142. 1923). 

Other authors then saw need to name still other species. M. L. 
Femald, although he accepted many of Blanchard's names, gave 24 
additional specific epithets to North American blackberries (Gray 
Herbarium index). P. A. Rydberg described a further 24, H. A. and T. 
Davis described 9 more, while still others were described and named 



Phytologia (Mar 2005) 87(1) 3 1 



by T. C. Porter, N. L. Britton, W. W. Ashe, J. K. Small, and L. H. 
Shinners. 

But the author of greatest importance, whose intensive field studies 
and voluminous publications on Rubus span more than half a century, 
was L. H. Bailey. His interest in Rubus grew slowly, with a single 
specific epithet published in 1898 and a second in 1902, By 1934 
Bailey had published a total of 52 specific names. His perceptions by 
then were acutely sensitized, and a profusion of new names appeared: 
42 in 1941, 79 in 1943, 40 in 1944, 74 in 1945, and 81 in 1947. With a 
few additional names in following years, Bailey was responsible for the 
description, the naming, and in nearly all cases the illustration of a total 
of 383 species of Rubus. 

Yet Bailey did not feel that the reservoir of undescribed species 
was near depletion. His compendious Species Batorum (Gentes 
Herbamm 5:1-932. 1941-45) provided coverage of the 390 species of 
Rubus then recognized by him in North America. In the next two years 
he accumulated a further 76 species, as described and illustrated in his 
supplementary Studies in Rubus (Gentes Herbarum 7:193-349. 1947). 
But the 466 species he thus recognized were no more than, in Bailey's 
opinion, "nearly or quite one-half the number of species native on the 
continent." 

Although North America is, in Bailey's words, "probably... the 
most fertile area on the globe for Rubus,'' Europe has fared no less well 
in terms of the number of specific epithets applied to the genus. 
Perhaps reflecting the greater density of botanists more than the 
European subcontinent's relatively impoverished vascular flora, the 
basic texts of Europe treat large numbers of minutely distinguished 
Rubus species. The volume of recognized names has compelled a 
stratagem for their handling, by the segregation of species into 
categories, or levels of importance, with only the principal species or 
"circle-species" given full treatment. In central Europe, H. Huber (in 
Hegi, Flora von Mitteleuropa IV/2A:274-41 1. 1964-66) recognized 33 
basic species and 252 subordinate species; these latter were described, 
but in different type size and without inclusion in the main key to 



32 Phytologia (Mar 2005) 87(1) 



species. In the most comprehensive modem European floristic 
treatment, Y. Heslop-Harrison (Flora Europaea 2:7-25. 1968) 
acknowledged 75 species of Rubus, of which 66 were "circle-species" 
representative of groups which contained an additional 374 
undescribed and unkeyed related specific names. Since there is no 
provision in the International Code, nor accepted systematic dogma, for 
the existence of species of more than one level of biological 
importance, the European stylistic practice, though maintaining the 
semblance of a workable structure, is as yet unexplored in its more 
fundamental implications. 

The redundancy of specific epithets in Rubus, both in Europe and 
America, has not failed to bring forth skeptics who questioned the 
biological significance of the named entities, even their very existence 
and the motivation of their authors. The American E. P. Bicknell, 
himself the author of 70 obscurely defined species in Sisyrinchium, 
expressed his unease at Blanchard's many Rubus names with an article 
plaintively entitled "Have we enough New England blackberries?" 
(Bull. Torrey Bot. Club 37:393-403. 1910). Defense of the reality of 
the new entities has rested largely with L. H. Bailey who provided a 
rationale with each of his major treatments of the genus. At no time 
was Bailey unaware of the unconventional image that his many species 
produced in the minds of classical systematists. He remarked in 1923 
(Gentes Herbarum 1 :143), "In Rubus. .At is not possible always to apply 
the formal species concept of ante-evolutionary days with either 
precision or satisfaction." In 1941 (G. H. 5:18) he said, "The reader 
may suppose that I have split the species finely. The opposite is the 
truth. I could have described any number more if I had cared to pursue 
a separatist course." In 1943 (G. H. 5:233) he commented, "With so 
many kinds now separated, the reader may wonder whether every 
colony is not a distinct species. This is a natural reaction..." He 
cautioned in 1944 (G. H. 5:508), "...persons not critical in Rubus will 
have little success with either pictures or specimens. The mind must 
first be free of notions and then the eye must be able to discriminate." 
And he rested his arguments in 1947 (G. H. 7:194) with the defense, "I 
have never made a species; I have only recognized, named and 
described them." 



Phytologia (Mar 2005) 87(1) 33 



Bailey gave repeated attention to the supposition that the many 
closely defined species recognized by him were the product of prolific 
hybridization among a limited number of true species. Bicknell had 
been insistent that the entities named by Blanchard could be accounted 
for as hybrids between no more than 1 1 species. Unwillingness to 
accept such an explanation was a theme that persisted through all of 
Bailey's works. His hostility to what he considered a glib and 
superficial interpretation was scarcely concealed (G. H. 5:6. 1941): 
"Early in the present century began the singular hybridity postulate in 
Rubus work, whereby hybrids were freely assumed from herbarium 
specimens so fragmentary that not even the species themselves can be 
determined; thereby was the fear of making new species escaped and 
the difficulty of understanding the plants was assuaged." He did not 
deny the possibility of natural hybridity in the genus, but challenged 
that its existence had yet to be proved and insisted that the entities 
described by him were readily recognized by a person of observant eye 
and sufficient field experience. 

Yet even as students of Rubus determinedly pursued the self- 
immolating course of recognizing and naming a seemingly endless 
series of dubious new species, and as Bailey shrugged off the shallow 
proposals that nothing more was involved than a massive hybrid 
swarm, contemporaries in the fields of embryology and cytogenetics 
were disclosing a framework of understanding on which a workable 
taxonomic structure could be hung. This was the discovery that 
reproduction by seed was a less than universally sexual process. In 
Europe, influenced particularly by the work and writings of A. 
Gustaffson, the concept came to be held that many genera of vascular 
plants display the phenomenon of agamospermy, or reproduction by 
seeds but without fertilization, as a special case within the general 
process of asexual reproduction or apomixis. Rubus was among the 
numerous genera in which agamospermic reproduction was suspected 
or identified. 

In North America, and particularly among classical systematists, 
such a concept was not quickly applied or perhaps understood. Bailey 



34 Phytologia (Mar 2005) 8 7(1) 



seems never to have given written expression of awareness that the 
individuals constituting many of his species might be of asexual origin, 
although his activities were indeed largely prior to widespread 
acceptance of the agamospermous pathway. He referred to the process 
only once, by indirect inclusion (G. H. 7:197. 1947): "I am asked for 
proof or at least for opinion that my novelties are not hybrids or 
apomicts or other irresponsibilities... of course no man has such 
proofs...." 

But Bailey was nonetheless in search of such proofs, whichever 
way they might point. In 1944 he had begun the support and 
encouragement of John Einset in New York State, in his study of the 
cytology and embryology of various Rubus clones. By 1951 (Amer. J. 
Botany. 38:768-772) Einset was able to publish the first satisfactory 
evidence that in North America Rubus there operated certain apomictic 
phenomena that had previously been demonstrated only in Old World 
species. 

Einset worked with 24 wild selections of Rubus which he brought 
into cultivation and which Bailey identified. He found the 
chromosome numbers to form a regular series of multiples of the basic 
7, ranging from 14 through 63 in the species studied, with only a single 
clone having an aneuploid number outside this series. Triploids, with 
the somatic number of 21, formed the most common grouping, 
representing a third of the clones examined. As had European workers 
previously, he found that pollination was necessary for seed 
production. By crossing clones with different chromosome numbers 
and by counting the chromosomes of the resultant seedlings, if the 
seedling count matched that of the seed parent and differed from that of 
the pollen parent he could assume that agamospermy was present. 

Einset's work strongly supports the assumption of apomixis in the 
American blackberries. When the chromosome numbers of the seed 
parent and the pollen parent were different, a high proportion of the 
seedlings (80 per cent in the case of tetraploid seed parents, 96 per cent 
with triploid seed parents) gave the same chromosome count as that of 
the seed parent. Had there been reduction of chromosome number with 



Phytologia (Mar 2005) 87(1) 35 



a chromosomal contribution from each parent, as in sexual 
reproduction, intermediate counts would have been obtained with much 
greater frequency. Einset could only conclude that a high percentage 
of his Rubus progeny resulted from the parthenogenetic development of 
unreduced eggs. 

The mechanics of reproduction in Rubus thus became clearer. 
Blackberries spread by vegetative means, with runners and rooting 
shoot-tips increasing the extent of the colony. Seeds are also formed, 
and serve as a means of dissemination across natural barriers and over 
distance. At times these seeds are produced by familiar sexual 
processes, and generate individuals that show the minute differences 
characteristic of genetic recombination. Perhaps also disparate 
individuals combine at times, to yield hybrids which differ from the 
offspring of conventional sexual reproduction only in the magnitude of 
their variability. 

But in Rubus a less familiar generative mode is also present, and 
may well form the dominant reproductive pattern within the genus. 
This is the agamospermous pathway, in particular the pollination- 
requiring variant known as pseudogamy. The progeny thus produced, 
while simulating the offspring of sexual reproduction, are identical in 
genetic composition and essentially identical in morphology to their 
maternal parent. By the agamospermous replication of these 
individuals in their turn, large numbers of clones may be generated. 
With seed-eating birds as vectors, their distribution will be limited only 
by factors of habitat availability and physiologic adaptability. 

A student inexperienced in Rubus who detects one of these 
agamospermic multi-clonal series is compelled to view it as he would 
an undescribed species. He finds the plants to have a coherent 
distribution, occur in predictable habitats, require pollination and 
reproduce by seed, and vary morphologically within a narrow range. 
His novelty is recognizable to him, and in almost every regard 
possesses the criteria that he associates with conventional species. It is 
perhaps inevitable that he should wish to bring legitimacy to his 



36 Phytologia (Mar 2005) 87(1) 



discovery by publishing its description and by coining a new name to 
serve as its label. 

And thus conflict arises in the study of Rubus. On the one hand is 
the insistence of close students of the genus, most of them of 
unquestioned taxonomic competence and extensive field experience, 
that they can recognize entities that are uniform, at times in many 
colonies distributed over hundreds of miles. On the other hand is the 
practical inability of less practiced workers to distinguish reliably 
among entities so very numerous that only the finite energies and 
lifetimes of their human identifiers appear to restrict their numbers. 

This conflict has no elegant solution. The organisms themselves 
create the hierarchy to be described, and it is neither linear nor 
consistent in its structure. The series of equivalent units of a sexual 
system does not have a parallel in an agamospermic complex. No 
usefiil purpose is achieved by insisting that apomictic microspecies can 
be recognized with sufficient study, for they can neither be keyed by 
the specialist nor identified by the workaday taxonomist. A coarser, 
less precise classificatory structure seems to be the only feasible 
approach. 

Thus one is led to a system that recognizes as species only major 
groupings of the genus Rubus. In the blackberries only one species 
appears usefully recognized in each section. In North America, Bailey 
(Gentes Herbarum 5:45-46. 1941) acknowledged ten sections in the 
true blackberries; the present scenario, if rigorously followed, would 
reduce the North American blackberries to this number of species. 
Among the raspberries, flowering raspberries, and cloud-berries the 
same degree of reduction is perhaps unwarranted, for it is not clear that 
the agamospermic process is as dominant there. But the blackberries, 
at least, appear more easily handled as a few aggregations of related 
forms than as constellations of numerous related microspecies. 

Such an approach implies the assumption that each aggregation 
consists of a, single basic species and its derived apomicts. Extensive 
work on Rubus in Europe, much of it by Gustafsson (see V. Grant, 



Phytologia (Mar 2005) 87(1) 37 



Plant Speciation 325-331. 1971), has shown that diploid sexual species 
form the phylogenetic foundation of the European blackberry flora. A 
similar evolutionary origin may reasonably be attributed to North 
American species. The ancestral diploids, as deduced from the 
European example, may indeed no longer be extant, being represented 
in modem times only by their polyploid pseudogamous offspring. 
Further, certain aggregations of apomicts appear derived, not from a 
single diploid, but from hybrids formed by crossings in distant times 
between two of the diploid species. One must therefore not expect that 
the natural groupings of microspecies will necessarily be demarked by 
the presence of a sexual diploid, nor that all clones will fall within the 
larger aggregations, however they may be arranged. 

Certain blackberry clones have been selected from the wild, and 
additional forms will undoubtedly be selected in the future, that possess 
characteristics of fruit, of flowers, or of vigor superior to those of the 
general population from which they come. These selections may have 
been recognized and named by previous students of the genus, or they 
may as yet be unnamed. The horticultural and other commercial 
importance of the selection may be such that a formal name is desired. 
In such situations, rather than a formal botanical binomial, the use of 
the flexible cultivar nomenclature would seem preferable. As 
examples, Rubus trivialis Michx. 'Marvel' (or Rubus trivialis cv. 
Marvel) and Rubus trivialis 'Okeechobee' are fully adequate 
replacements for Rubus minis Bailey and Rubus okeechobeus Bailey, 
respectively. Rubus flagellaris Willd. 'Almus' and R. flagellaris 'Foster 
Thomless' are among other listed cultivar names. 

Florida does not have a complicated blackberry flora. Only four of 
the ten sections recognized by Bailey are represented within the state: 
Arguti Rydb. (=Frondosi Bailey, Floridi Bailey), Cuneifolii Bailey, 
Flagellares Bailey (=Procumbentes Rydb., Tholiformes Fern.), and 
Verotriviales Bailey (=Persistentes Fern., Triviales Rydb.). Within 
these sections fall all of the 15 Rubus specific names based upon 
Florida types (1 by Rydberg, 14 by Bailey), as well as the uncounted 
but numerous names typified elsewhere but applied to Florida 
blackberries. Among these other names are four that typify the above 



38 Phytologia (Mar 2005) 87(1) 



four sections and should be used to denote the major aggregations of 
Rubus as found in Florida: Rubus argutus Link, Rubus cuneifolius 
Pursh, Rubus flagellaris Willd., and Rubus trivialis Michx. 



Rubus L. Blackberries 



1. Stems trailing or supported by low vegetation, elongate (to 2 m.), 
densely set with both stout prickles and numerous stiff bristles, 
usually mahogany red; leaves persistent throughout winter, 
subcoriaceous, leaflets glabrous, dark glossy; flowers usually 
solitary, on erect pedicels; petals white. Prickly trailing shrub. 
Thickets, open woodlands, brushy fields. Throughout; common. 
Spring. [R. agilis Bailey; R. continentalis Bailey; R. lucidus 
Rydb.; R. mirus Bailey; R. okeechobeus Bailey] 

SOUTHERN DEWBERRY. Rubus trivialis Michx. 

1 . Stems erect, arching at tips, or declining and rooting, with prickles 
but with few or no slender bristles; leaves deciduous, thin, leaflets 
glabrous or densely pubescent below; flowers mostly in 3 -several- 
flowered panicles. 

2. Stems erect when young but soon arching and declining, often 
rooting at tips; prickles weak, sparse, often almost lacking; 
petals white. Prickly arching shrub, to 0.8 m. Mesic 
woodlands, clearings, old fields. Panhandle (east to Madison 
County); infrequent. Spring. [Rubus enslenii Tratt] 
NORTHERN DEWBERRY. Rubus flagellaris Willd. 

2. Stems remaining erect, although often arching toward tip, not 
rooting; prickles stout, numerous, recurved. 

^The "amplified key" format employed here is designed to present in compact form the 
basic morphological framework of a conventional dichotomous key, as well as data on 
habitat, range, and frequency. This paper is a continuation of a series begun in the 1970s 
(vide Phytologia 35:404-413. 1977). I wish to thank David W. Hall and Kent D. Perkins 
for constructively reviewing the manuscript. 



Phytologia (Mar 2005) 87(1) 39 



3. Leaflets ovate to oblong, glabrous or nearly so; stems 
commonly to 2 m. (to 8 m., when supported by 
surrounding vegetation); panicle often several-flowered; 
petals white; fruits sweet. Prickly arching shrub. Moist 
to dry thickets, pond margins, swamps. Throughout 
panhandle and north Florida, south to mid-peninsula 
(Highlands, Okeechobee counties); common. Spring. [R. 
betulifolius Small; R. floridus Tratt; R. harperi Bailey; R. 
penetrans Bailey; R. rhodophyllus Rydb. in Small;' i?. 
tallahasseanus Bailey; R. ucetanus Bailey; R zoae 
Bailey] 

HIGHBUSH BLACKBERRY. Rubus argutus Link 

3. Leaflets obovate, densely gray-pubescent beneath; stems 
usually less than 1 m.; panicle usually 1-3-flowered; 
petals white; fruits bland. Prickly arching shrub. Dry 
sands, old fields, disturbed areas. Nearly throughout 
(excl. south peninsula); common. Spring. [R. audax 
Bailey; R. chapmanii Bailey; R. floridensis Bailey; R. 
humei Bailey; R. inferior Bailey] 

SAND BLACKBERRY. Rubus cuneifolius Pursh 



40 Phytologia (Mar 2005) 87(1) 

A NEW SPECIES OF SEYMERIA (SCROPHULARIACEAE) 
FROM OAXACA, MEXICO 

B. L. Turner 

Plant Resources Center, University of Texas, Austin 78712-0471, USA 

ABSTRACT 

Seymeria mazatecana B. L. Turner, sp. nov., is described from 
northeastern Oaxaca, Mexico. It is closely related to S. decurva but is 
readily distinguished from that taxon by vestiture and shape of its 
mature capsules. A photograph of the holotype is provided. 

KEY WORDS: Scrophulariaceae, Seymeria, Mexico, Oaxaca. 



Routine identification of Mexican plants has revealed the 
following novelty: 

Seymeria mazatecana B. L. Turner, sp. nov. Fig. 1 

Similis Seymeriae decurvae Benth. ex DC, sed plantae herbae 
compactae, caules dense villosi, et capsulae late ovoideae (vice 
capsularum ellipsoidearum) sunt. 

Compact perennial herbs 15-30 cm high, drying black. Stems 
densely villous with multicellular spreading or reflexed hairs 0.5-1.0 
mm long. Leaves pinnately lobed, those at mid-stem mostly 6- 1 2 mm 
long, 5-8 mm wide, pubescent with both short hispidulous hairs and 
capitate-glandular hairs, interspersed among these a smattering of 
longer crinkly hairs such as occur on the stems. Inflorescence a 
terminal leafy raceme up to 12 cm long bearing 10-20 flowers. Pedicels 
8-11 mm long. Calices 6-7 mm long having 5-8 irregular lobes 2-4 
mm long. Corollas yellow, arcuate-urceolate, the tube glabrous 
without, pubescent within, ca. 4 mm long, 3 mm wide, the throats ca. 8 
mm long, 6 mm wide, capitate-glandular, their lobes broadly cordate, 



Phytologia (Mar 2005) 87(1) 4 1 



w 





Fig.l. Seymeria mazatecana, holotype. 



42 Phytologia (Mar 2005) 87(1) 



ca. 4 mm long, 4-5 mm wide, having ciliate margins. Stamens 4; 
anthers 4-5 mm long; filaments 2.0-2.5 mm long, densely pilose to the 
apex or nearly so. Capsules broadly ovoid, ca. 8 mm high, 6 mm 
across, capitate-glandular, the style persistent, 6-8 mm long. Seeds 
(somewhat immature) ca. 15 per locule, ca. 2 mm long, ca. 1.5 mm 
wide, their margins with narrow undulate wings. 

TYPE: MEXICO. OAXACA: Mpio. San Jeronimo Tecoatl, 
"Vereda (antiguo camino real) del Plan de Guadalupe al Cerro Pelon" 
(18 08 10 N X 96 58 20 W) ca. 2640 m, 6 Jul 2001, Xochitl Munn- 
Estrada et al. 1351 (HOLOTYPE: MEXU; ISOTYPE: TEX). 

This species clearly belongs to the Virgatae group of Seymeria as 
treated by Perinell (1925) where it will key to the widespread S. 
decurva. It will also key to the latter in my two treatments of Seymeria 
for Mexico (Turner, 1982; 1995). The present novelty differs from S. 
decurva in being a smaller, more compact plant having a villous 
vestiture on its stems and broadly ovoid capsules (as opposed to ovate- 
ellipsoid capsules as consistently found in S. decurva). 

The only specimen of S. decurva previously collected in the state 
of Oaxaca has been that of Purpus 3264 (vicinity of "Cerro Verde," 
this housed at UC). While I examined the latter in my initial study of 
the group (Turner 1982), I can not recall if it was in fruit or not; at least 
I did not record the capsule as broadly ovoid, nor the stems as being 
villous, characters emphasized here. Nevertheless, it is possible that the 
specimen concerned belongs to S. mazatecana, memories being 
fallible. 

The species is named for the sierra where it was first collected. I 
intended to name the taxon for its principal collector, but when told of 
my intentions she humbly replied," Please no, I deplore such honors, 
besides it would be more meaningfully named for the sierra in which it 
was collected," or words to that effect. But left to myself I would have 
preferred this attractive herb bear the name of its collector, Xochitl 
Munn-Estrada. She is currently enrolled in a graduate program in 



Phytologia (Mar 2005) 87(1) 43 



botany at The University of Texas, Austin working on a floristic study 
of the mountains concerned. 

ACKNOWLEDGMENTS 

I am grateful to my wife Gayle Turner for the Latin diagnosis and 
to her and Ms Munn-Estrada for reviewing the paper. 

LITERATURE CITED 



Pennell, F.W. 1925. The genus Afzelia: a taxonomic study in 
evolution. Proc. Acad. Nat. Sci. Phila. 77: 335-373. 

Turner, B.L. 1982. Revisional treatment of the Mexican species of 
Seymeria (Scrophulariaceae). Phytologia 51: 403-419. 



1995. A new species oi Seymeria (Scrophulariaceae) from 

Mexico. Phytologia 78: 28-35. 



44 Phytologia (Mar 2005) 87(1) 

A NEW SPECIES 0¥ AGERATUM {XST^KACEX^\ 
EUPATORIEAE) FROM NAYARIT, MEXICO 

B. L. Turner 

Plant Resources Center, University of Texas, Austin 78712-0471, USA 

ABSTRACT 

A new species, Ageratum solisii sp. nov., is described from Mpio. 
de Huacori, Nayarit, Mexico. It resembles the widespread, highly 
variable, A. corymbosum but is readily distinguished by a combination 
of characters including deltoid, markedly reticulate leaves, and more 
numerous much smaller heads. A photograph of the holotype is 
provided, 

KEY WORDS: Ageratum, Asteraceae, Eupatorieae, Nayarit, 
Mexico. 



In The Comps of Mexico (Turner 1997) Ageratum is recognized as 
having 11 species; the present novelty brings this total to 12. In my 
treatment, the new species described below {A. solisii) will key to or 
near A. tomentosum (Benth.) Hemsl. because of its suffruticose habit 
and thick deltoid leaves. The latter taxon is confined to the tri-comer 
region of the states of Veracruz, Pueblo, and Oaxaca and can be 
immediately distinguished from A. solisii by its markedly bicolored 
leaves and few-headed capitulescence. In reality, A. solisii is probably 
more closely related to the widespread A. corymbosum Zucc. ex Pers., 
but is readily distinguished from this by a combination of characters, as 
noted in the above abstract . 

Ageratum solisii B. L. Turner, sp. nov. Fig 1. 

Similis Agerato corymboso Zucc. ex Pers. sed laminis foliorum 
deltatis, valde nervatis reticulate infra, et capitulis parvioribus 
pluribusque per capitulescentiam. 



Phytologia (Mar 2005) 87(1) 45 



I 









-<"/-" 





The U~Ptv*r5.?v e^ TesfAa ttsfbiarn 



Fig 1. Holotype of Ageratum solisii. 



46 Phytologia (Mar 2005) 87(1) 



Suffruticose herbs or shrublets up to 1 m high. Stems densely 
hirsute, the vestiture ca. 0.5 mm high. Leaves opposite, pubescent like 
the stems, 2.0-4.5 cm long, 2-3 cm wide; petioles 0.5-1.0 cm long; 
blades deltoid, rugose above, markedly reticulate-veined beneath, 
subcordate at the base; margins irregularly crenate. Heads numerous, 
arranged in terminal corymbose, somewhat flat-topped capitulescences 
12-15 cm across, and about as high, the ultimate peduncles mostly 1-5 
mm long. Heads relatively small, campanulate, the involucres ca. 4 mm 
high, composed of ca. 21 linear scales arranged in 2 series. Receptacle 
subconical, naked, glabrous, ca 1 mm across, 0.5 mm high. Florets 20- 
30 per head; corollas white, 1.5-2.0 mm long; throats ca. 1.2 mm long, 
grading into a weakly defined throat, atomiferous-glandular 
throughout, the lobes pubescent. Achenes ca. 1 mm long, glabrous; 
pappus a small saucer-shaped crown of united scales ca. 0.25 mm high. 

TYPE: MEXICO. NAYARIT: Mpio. de Huajcori, Rancho de Los 
Sauces, mouth of canyon in "Bosque Templado (encinos chaporros y 
otros arboles)." 28 Sep 1985, /. Solis 577 (HOLOTYPE: TEX). 

ADDITIONAL COLLECTION EXAMINED: MEXICO. 
NAYARIT: same locality as type, 30 Sep 1985, /. Solis 527 (TEX). 

The species is named for Ignacio Solis Cumplido ("Don Nacho") 
who is a Tepahuan Indian and collected for CIIDR-Durango for several 
years in extreme southwestern Durango and nearby Nayarit, in the 
canyon country. 

ACKNOWLEDGEMENTS 

I am gratefiil to Gayle Turner for the Latin diagnosis and to Tom 
Wendt for obtaining information as to the credentials of the collector. 
M. Dias de Moraes and H. S. Berg provided the digital photos. 

LITERATURE CITED 

Turner, B.L. 1997. Ageratum, in Comps of Mexico 1, Phytologia 
Memoirs 11: 51-55. 



Phytologia (Mar 2005) 87(1) 47 



BIDENS MELCHERTII (ASTERACEAE), A NEW SPECIES 
FROM JALISCO, MEXICO 

B. L. Turner 

Plant Resources Center, University of Texas, Austin 78712-0471, USA 

ABSTRACT 

A new species, Bidens melchertii sp. nov., is described from the 
state of Jalisco, Mexico. It is closely related to the square-stemmed, 
white-rayed, perennial, B. pringlei, but is readily distinguished from the 
latter by characters of the achene. 

KEY WORX)S: Bidens, Asteraceae, Mexico, Jalisco. 

Preparation of a taxonomic treatment of the genus Bidens for 
Mexico by Prof Thomas Melchert of the University of Iowa has 
necessitated description of the following novelty: 

Bidens melchertii B. L. Turner, sp. nov. Fig. 1. 

Similis Bidens pringlei S. Wats, sed acheniis sine pappis (non 
biaristatis) et brevioribus, corporibus tetragonis et non recurvatis. 

Erect rhizomatous herbs to 80 cm high. Stems suffruticose, leafy 
throughout, the lower portions multinerved, the upper essentially 
square in cross section, glabrous or nearly so. Leaves deeply (l)-2 
pinnatisect, 4.0-7.5 cm long; petioles 1-2 cm long; ultimate divisions 
linear to lance-linear, 0.5-2.0 mm wide, their apices somewhat pungent. 
Heads showy, 2-3 cm across the expanded rays. Ray florets 8; ligules 
creamy white, 6-12 mm long, ca. 5 mm wide. Disk florets numerous, 
yellow. Outer phyllaries ca 10, linear with sharply acute apices, 
recurving with age. Inner phyllaries yellow-margined. Receptacular 
bracts linear-lanceolate, their apices extending beyond the 



48 Phytologia (Mar 2005) 87(1) 



w 



r 



#// 



45yz] w. 






U3g 



H E t 8 A R t O 



Fig. 1. Bidens melchertii, holotype (TEX). 



Phytologia (Mar 2005) 87(1) 49 



disk florets. Achenes epappose, black, glabrous, ca. as long as the 
chaff, linear, tetragonal in cross section, the outer series notably 
incurved and somewhat obcompressed, 3.7-6.0 mm long (the very 
outermost nearly linear-clavate, the innermost nearly straight, to 8.5 
mm long). 

TYPE: MEXICO. JALISCO: Mpio. De Talpa, "Km. 15 Camino 
El Tuito a Mina de Zimapan," pine forests, 1650 m, 14 Oct 1989, 
Carlos Diaz Luna & Jose A. Lomali S. 20968 (HOLOTYPE: TEX; 
ISOTYPE: MEXU). 

ADDITIONAL SPECIMEN EXAMINED: MEXICO. JALISCO: 

Mpio. De Talpa, Cerros de Las Minas de Oro, pine-oak forests, 2080 
m, 30 Oct 1971, 7?. Gonzales T. 562 (MICH). 

Bidens talpana superficially resembles B. pringlei, both being 
square-stemmed perennials with deeply pinnatisect leaves having 
nearly filiform ultimate segments, and possessing showy white-rayed 
heads. Their achenes, however, are quite distinct, those of B. pringlei 
having flatter, long-attenuate achenes with slightly recurved biaristate 
apices (vs. pappus absent). Bidens talpana also resembles B. mollifolia 
Sherff, the two taxa having short, clavate to linear- clavate achenes. 

The species is named for my sixth academic son, Thomas 
Melchert, currently Prof Emer. at the Univ. of Iowa, Ames, and 
longtime scholar of the genera Bidens, Cosmos, and Thelesperma. He 
called this novelty to my attention in his submitted manuscript for the 
upcoming treatment of Bidens for Mexico (cf Turner, Comps of 
Mexico, Vol. 1. Eupatorieae. Phytologia Memoirs 11. 1997). Without 
his knowledge, I proudly provided the eponym concerned. 

ACKNOWLEDGEMENTS 

I am grateful to Gayle Turner for the Latin diagnosis; Marta Dias 
de Moraes and Heidi S. Berg prepared the digital photos. 



50 Phytologia (Mar 2005) 87(1) 



A NEW SPECIES OF TRIXIS (ASTERACEAE: MUTISIEAE) 
FROM OAXACA, MEXICO 

B. L. Turner 

Plant Resources Center, University of Texas, Austin 78712-0471, USA 

ABSTRACT 

Trixis hintoniorum B. L, Turner, sp. nov., is described from 
Oaxaca, Mexico. So far as known, it is a prostrate shrub with winged 
stems confined to the rocky beaches of southern Oaxaca, Mpio. 
Pachutla. A photograph of the holotype is provided. 

KEY WORDS: Asteraceae, Mutisieae, Trixis, Mexico, Oaxaca. 

Routine identification of Mexican Asteraceae has revealed the 
following novelty: 

Trixis hintoniorum B. L. Turner, sp. nov. Fig. 1. 

Similis T. silvaticae sed foliis crassioribus, caulibus sine alis, 
flosculis omnibus similaribus, labiis exterioribus 1-2 mm longis. 

Shrub, reportedly (from label data) "common" and forming 
"procumbent" colonies. Stems puberulous, wingless. Larger leaves 
(ca. 10 cm below capitulescence) 6-8 cm long, 3.5-4.0 cm wide, sessile 
or nearly so, relatively thick, pinnately venose, somewhat bicolored, 
moderately appressed-pilose beneath, the margins serrulate. Heads 
subtended by 2-4 ovate bracts ca. as long as the involucres, mostly 
arranged 3-5 in terminal congested cymes scarcely exceeding the 
leaves. Involucral bracts 11, linear-lanceolate, appressed-pubescent, 
the apices gradually acuminate. Receptacles densely pubescent, the 
hairs 0.5-1.0 mm long. Florets ca. 23, the corollas yellow, all more or 
less alike, those at the periphery with outer lips 1 -2 mm long. Achenes 
(immature) ca. 5 mm long, densely pubescent throughout; pappus of 
numerous tawny bristles 9-10 mm long. 



Phytologia (Mar 2005) 87(1) 5 1 



I 




Fig. 1 . Trixis hintoniorum (Holotype TEX). 



52 Phytologia (Mar 2005) 87(1) 



TYPE: MEXICO. OAXACA: Mpio. Pochutla, Playa La Tijera, 
sea level along rocky beaches, 27 Oct 1995, Hinton et al. 26474 
(Holotype TEX). 

In Anderson's (1972) treatment of Trixis, because of its wingless 
stems, this taxon will key to or near T. megalophylla Greenm. and/or T. 
silvatica Robinson & Greenm., but it is markedly different from both. 
According to label data and appearance of pressed material, T. 
hintoniorum is a prostrate shrub occurring along the beach. In habit 
and leaf shape it resembles T. silvatica, but possesses thicker leaves 
and larger, more numerous-flowered heads, having 1 1 involucral bracts 
(versus 8). Trixis hintoniorum also has characteristics of T. parviflora, 
a coastal species of northern Oaxaca and Guerrero, but the latter 
possesses narrower thinner leaves, the heads with only 8 involucral 
bracts and 10-13 florets. 

It should be noted that Anderson, in her discussion of T. silvatica, 
mentioned a single "population" of the latter to have 11-13 involucral 
bracts and 27-29 florets per head (specimen not cited). This 
description fits the heads of T. hintoniorum, but corollas of the latter 
have markedly small lips (1-2 mm long vs 3.1-4.0 mm) and, as already 
noted, has markedly winged stems. 

ACKNOWLEDGEMENTS 

I am grateful to Gayle Turner for the Latin diagnosis and to her 
and Ms. Xochitl Munn-Estrada for reviewing the paper. 

LITERATURE CITED 

Anderson, C. 1972. A monograph of the Mexican and Central 
American species of Trixis (Compositae). Mem. N. Y. Bot. Gard. 
22: 1-68. 



Phytologia (Mar 2005) 87(1) 53 

ELEVATION OF STEVIA LEMMONIIYAR. HISPIDULA 
(ASTERACEAE: EUPATORIEAE) TO SPECIFIC RANK 

B. L. TURNER 

Plant Resources Center, University of Texas, Austin 78712-0471, USA 



Grashoff (1972, 1974) described Stevia lemmonii A. Gray var. 
hispidula Grashoff, distinguishing this from its sister taxon, S. I. var. 
lemmonii, by its glandular-pubescent involucres (vs. eglandular) and its 
more southern distribution. He commented further that "It would 
appear from the ranges of the two varieties [Fig.l] that var. hispidula is 
the more primitive of the two, var. lemmonii being a more northern, 
glandular taxon." Grashoff also noted that "neither Gray (1882) nor 
Robinson (1930) ever mentioned the glandular pubescence of the 
typical variety, although they often placed great emphasis on this 
character in other groups." 

In my recent account of Stevia for Mexico (Turner, 1 997) I largely 
followed Grashoff s treatment, this based upon relatively few 
specimens. Subsequent collections of this duo from northwestern 
Mexico suggest that they are worthy of specific rank, at least I have not 
found that these two allopatric sister taxa show any signs of 
intergradation. My views regarding the treatment of such infraspecific 
variation are discussed in more detail elsewhere (Turner and Nesom, 
2000). The necessary nomenclature follows: 

Stevia hispidula (Grashoff) B. L. Turner, stat. nov. 

BASIONYM: Stevia lemmonii var. hispidula Grashoff, Brittonia 
26:364. 1974. 

Turner (1997) noted that Grashoff cited a single specimen {Gentry 
5587) from Sinaloa, Mexico that he referred to var. lemmonii. At the 
time I took the collection concerned to be a "stipitate-glandular form" 
of var. hispidula. Recent collections from the area concerned now 
suggest that Grashoff was correct in his original observation. 



54 Phytologia (Mar 2005) 87(1) 




Fig 1. Distribution oiStevia lemmonii (closed circles) and S. hispidula 
(open circles). 

LITERATURE CITED 

Grashoff, J. 1972 A systematic study of the North and Central 
American species of Stevia. Doctoral Dissertation, the University 
of Texas, Austin. 



1974. Novelties in Stevia (Compositae: Eupatorieae). 

Brittonia 26: 347-384. 



Turner, B. L. 1997. , in Comps of Mexico 1, Phytologia Memoirs 11: 
170-197. . 



Phytologia (Mar 2005) 87(1) 55 



and G. Nesom. 2000. Use of variety and subspecies and 

new varietal combinations for Styrax platanifolius (Styracaceae) 
Sida 19: 257-262. 



I 



I 



I 



56 Phytologia (Mar 2005) 87(1) 



BOTANICAL ANECDOTES: Bajacalia moranii (Asteraceae: 
Tageteae), comments of James Reveal regarding its discovery 

B. L. Turner 

Plant Resources Center, University of Texas, Austin 78712-0471, USA 

Recent description (Loockermann et al. 2003) of the new species, 
Bajacalia moranii B. L. Turner elicited the following email from James 
L. Reveal, who participated in its discovery, type material having been 
collected with Reid Moran on a joint collecting trip to Baja California 
during February of 1973. 

Morning Billie: 

A tale about the discovery of Bajacalia moranii. 

We drove down Arroyo de Portezuelo looking for Eriogomum 
encelioides, a plant found in 1947 by Scott Gentry. Upon finding a few 
shrubs, we stopped the jeep and I got out my stuff to write a description 
based on what I could see in the field. 

Reid said he had to visit a bush. 

I said I would look further down the arroyo to see what the 
variation might be. He went up the draw; I went down the draw. 

Just around the comer I found what proved to be Eriogonum 
preclarum and yelled back to Reid that we would be a while as I found 
a new buckwheat. 

He yelled back that it was okay. He was taking a dump and a new 
Porophyllum was by his side. 

Ahhh, how one finds new species.... 

-Jim 



\ 



4 



Phytologia (Mar 2005) 87(1) 57 



The new species of Porophyllum, declared to be so by Reid and 
confirmed in the afore mentioned study with the description of B. 
moranii, was never pubHshed of course, but had I known the details of 
its discovery I might have provided a very different Latin name! 

I am grateful to my fried Jim who permitted my use of his email 
communication dated 14 Mar 2003. 

LITERATURE CITED 

Loockerman, D. J., B. L. Turner, and R. K. Jansen. 2003. 
Phylogenetic relationships within the Tageteae (Asteraceae) based 
on nuclear ribosomal ITS and chloroplast ndhF gene sequences. 
Systematic Botany 28: 191-207. 



I 



58 Phytologia (Mar 2005) 87(1) 



BOTANICAL HUMOR 







"Unyk, Professor, I fm^ hi The Iml mUiml Tfrmimlm ithimtii^:" 



In June of 1994 my family and I traveled through Scotland for two weeks. 
It was the second summer after I had discovered Botany and I was determined 
to learn every plant in Scotland by feverishly collecting every species and 
keying them out in Stace (1991). One plant, Circea lutetiana L., was described 
as rare in Northern Scotland. Not quite understanding if "rare" meant 
endangered or simply not common I fretted that I had extirpated the last 
population of C. lutetiana in Scotland. My father related this story to a friend 
of the family, Molly Sadler. Molly, a teacher of ESL at Amarillo College, 
illustrates as a hobby and has had works published in the Wall Street Journal 
and Forbes. With my father's (much embellished) story as inspiration she drew 
the above picture. The illustration is proudly displayed in my office and it is 
with much joy that I present it in Phytologia. -Justin Williams, Editor 

Stace, C, 1991. New Flora of the British Isles. University of Cambridge Press. Pp 1226. 



Phytologia (Mar 2005) 87(1) 59 



INDEX TO NEW NAMES AND COMBINATIONS IN 
PHYTOLOGIA 87(1) 

Ageratum solisii B. L. Turner, sp. nov. 44 

Anisopappinae Panero, subtrib nov. 5 

Athroisminae Panero, subtrib. nov. 2 

Bidens melchertii B. L. Turner, sp. nov. 47 

Centipedinae Panero, subtrib nov. 3 

Chromolepidinae Panero, subtribus nov. 5 

Doroniceae Panero, tribus nov. 1 

Dugesiinae Panero, subtribus nov. 7 

Dyscritothamninae Panero, subtribus nov. 10 

Enceliinae Panero, subtribus nov. 8 

Euphrosyne acerosa (Nutt.) Panero, comb. nov. 13 

Euphrosyne dealbata (A. Gray) Panero, comb. nov. 13 

Euphrosyne nevadensis (M. E. Jones) Panero, comb. nov. 13 
Haplodontium macrocarpum (Hook, ex Drumm.) J.R. Spence, comb. nov. 26 

Haplodontium tehamense (Showers) J.R. Spence, comb. nov. 26 

Jaegeriinae Panero, subtribus nov. 1 1 

Leptostomopsis (C. Muell.) J.R. Spence & H.P. Ramsay, stat. nov. 23 

Plagiobryoides JR. Spence, gen. nov. 24 

Plagiobryoides incrassatolimbata (Card.) J.R. Spence, comb. nov. 25 
Ptychostomum angustifolium (Brid.) JR. Spence & HP. Ramsay, comb. nov. 23 

Ptychostomum archangelicum (B.S.G.) J.R. Spence, comb. nov. 20 

Ptychostomum arcticum (R. Br.) J.R. Spence, comb. nov. 20 

Ptychostomum badium (Brid.) JR. Spence, comb. nov. 20 

Ptychostomum bimum (Schreb.) J.R. Spence, comb. nov. 20 

Ptychostomum calophyllum (R. Br.) J.R. Spence, comb. nov. 20 
Ptychostomum creberrimum (Taylor) JR. Spence & HP. Ramsay, comb. nov. 23 

Ptychostomum cryophilum (Mart.) J.R. Spence, comb. nov. 20 

Ptychostomum curvatum (Kaur. & Am.) J.R. Spence, comb. nov. 20 

Ptychostomum cyclophyllum (Schwaegr.) J.R. Spence, comb. nov. 20 

Ptychostomum inclinatum (C. Muell.) J.R. Spence, comb. nov. 21 

Ptychostomum intermedium (Brid.) J.R. Spence, comb. nov. 21 

Ptychostomum knowltonii (Barnes) JR. Spence, comb. nov. 21 

Ptychostomum lonchocaulon (C. Muell.) JR. Spence, comb. nov. 21 
Ptychostomum longisetum (Bland, ex Schwaegr.) J.R. Spence, comb. nov. 21 

Ptychostomum marratii (Hook. & Wils.) J.R. Spence, comb. nov. 21 

Ptychostomum meesioides (Kindb.) J.R. Spence, comb. nov. 21 

Ptychostomum neodamense (Itzigs.) J.R. Spence, comb. nov. 21 

Ptychostomum pallens (Sw.) J.R. Spence, comb. nov. 21 



60 Phytologia (Mar 2005) 87(1) 



Ptychostomum pallescens (Schleich.) J.R. Spence, comb. nov. 21 
Ptychostomum pseudotriquetrum (Hedw.) JR. Spence & HP. Ramsay, comb. nov. 23 

Ptychostomum purpurascens (R. Br.) JR. Spence, comb. nov. 22 

Ptychostomum reedii (Robins.) JR. Spence, comb. nov. 22 

Ptychostomum rutilans (Brid.) J.R. Spence, comb. nov. 22 

Ptychostomum salinum (Hag. ex Limpr.) J.R. Spence, comb. nov. 22 

Ptychostomum schleicheri (Schwaegr.) J.R. Spence, comb. nov. 22 

Ptychostomum turbinatum (Hedw.) JR. Spence, comb. nov. 22 

Ptychostomum warneum (Rohl.) J.R. Spence, comb. nov. 22 

Ptychostomum weigelii (Spreng.) J.R. Spence, comb. nov. 22 

Ptychostomum wrightii (Sull. & Lesq.) J.R. Spence, comb. nov. 22 

Seymeria mazatecana B. L. Turner, sp. nov. 40 

Spilanthinae Panero, subtribus nov. 9 

Stevia hispidula (Grashoff) B. L. Turner, stat. nov. 53 

Trixis hintoniorum B. L. Turner, sp. nov. 50 



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