Phytologia

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 \
REO SA np Re REPL OCIA He CH ee is*:

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

B. L. TURNER, A new species of Seymeria (Scrophulariaceae) from Oaxaca,
eI he al 7 eas Rees Fay eae aie ete 40 ~~

B. L. TURNER, A new species of Ageratum (Asteraceae; Eupatorieae) from
PREC MIC ere ee 8 peg ay PES HAD iheincer apy AME Grae aA. 44

B. L. TURNER, Bidens melchertii (Asteraceae), a new species from Jalisco, a
NOTES SD RE RS POE RR RG RECN 5 OC) ADE yal eed Coe 47

B. L. TURNER, A new species of Trixis (Asteraceae: Mutisieae) from Oaxaca, ”
2 EE SGD: eg SES Oe RE Re Nae MEO eIy Tape ct rat NI ye URN 50

B. L. TURNER, Elevation of Stevia lemmonii var. hispidula (Asteraceae:
Eupatorieae) to specific rank..................0.4. ida gibt ped URE Ie tafe hee ATR aS 53

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

RTE” SORT MANS Dad en siete Me TRB ct eae eRe eins ara | 56
oy a TE Heo De SS EMA REA SE ROOM, DAE UB EAN gradi ere CR po Conk 58
Index to new names and new combinations in volume OTL is 5 sacha ta aes peek Gag ie Cam ee 59

_ Subscription Information & Information for Authors...........¢.¢00eecccccccceeeeceece. Inside back Jacket

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: 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 \
EES SCE aa Oa OIE EAL PL en HR OOM Nag ise:

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

B. L. TURNER, A new species of Seymeria (Scrophulariaceae) from Oaxaca,
11 RSE SOR Se aU anes ORM ERO SN ct p nyt AML LEM tah UN aa ce 40 ~

B. L. TURNER, A new species of Ageratum (Asteraceae; Eupatorieae) from -

CULTS Tg aed SRR SEE Fol ae a eae Siete) eR ORIRS Lone Dy Jil era) PWR DE MEG RC. 2, voids, 44
B. L. TURNER, Bidens melchertii (Asteraceae), a new species from Jalisco, _-
oe EGS 9 Se Pe Apna GRE PaaS OR RIM aLene rh N ph Ne SARE RIE tt 47
B. L. TURNER, A new species of Trixis (Asteraceae: Mutisieae) from Oaxaca, om
MME RN hgh PCF dai cro ait has ov Aur Gicg sANee S dees SDRS IR) earn eee MARS aR aah aa tod am iad 50
B. L. TURNER, Elevation of Stevia lemmonii var. hispidula (Asteraceae:
Eupatorieae) to specific rank...................... ia i dk Shad De Oe Fa Cop aee a Se tea eas a3
Botanical Anecdotes: Bajacalia moranii (Asteraceae: Tageteaé), comments regarding its
WMI ehh eee TTL dy Sits b cine Labo vae he ots Ddda UTNE PRON TTA Oued rt Re ENN 56
Tye TOT Al a UC Giga eee hg aie OY A a COP ENO DL BPE RE ipa may Peri Saad get Ale, a ein te 58
Index to new names and new combinations in volume 87(1)................ccceceeeeenee geen eeeeeeeeanneees 59

_ Subscription Information & Information for Authors...........6......:cesceeeneee seen es Inside back Jacket

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
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Phytologia (Mar 2005) 87(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 alterniis, 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 alterniis vel fasciculatis, capitulis discoiformis;

phyllariis absentibus vel 1-seriatis, receptaculis paleatis; flosculorum
peripherialum pistillatis, corollis 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 = 10

The subtribe contains three genera including Athroisma (12 spp.),
Blepharispermum Wight ex DC. (15 spp.) and Leucoblepharis Arn. (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 alterniis, 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.
& Arn.

A Athroismeae ceteris, differt characteribus conjuncte: habitu
herbaceo; foliis alterniis; 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 alterniis, petiolatis, laminis integris vel lobatis/runcinatis;
capitulis ligulatis; phyllariis 2-3 seriatis, dimorphis; receptaculis
paleatis; flosculis radiorum pistillatis; flosculis discorum bisexualibus,
corollis 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 dimorphic involucre of Chromolepis are characteristics not
seen elsewhere in Zaluzaniinae.

Phytologia (Mar 2005) 87(1) 7

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

A Heliantheae ceteris, differt characteribus conjuncte: habitu
herbaceo; foliis alterniis, 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 alterniis, 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,

10 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 alterniis, 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, corollis actinomorphis, lobis 5; antheris

Phytologia (Mar 2005) 87(1) 11

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) &7(1)

vel sessilis, laminis integris; capitulis ligulatis; phyllariis 1- seriatis;
receptaculis paleatis; flosculis radiorum pistillatis; flosculis discorum
bisexualibus, corollis 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) 13

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

Euphrosyne dealbata (A. Gray) Panero, comb. nov. [va dealbata A.
Gray, Pl. Wright. 1:104. 1852.

Euphrosyne nevadensis (M. E. Jones) Panero, comb. nov. /va
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. Harborne, 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.

a eee eee

Phytologia (Mar 2005) 87(1) 15

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 Hornsch. 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 alternate-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./. 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 (cf 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 “Cernuibryum” 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) 17

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 19" 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 Hornschuch
(1822) and Bridel (1826) described new genera long since
synonymized under Bryum. In Hornschuch’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.

18 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), B. 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
“Cermuibryum” 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 Hornschuch for those species of Bryum in the
informal group “Cernuibryum” of Spence (1987). Since all three
species first cited by both Bridel and Hornschuch 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.) Hornsch. 1822 Bryum uliginosum
P. compactum Hornsch. 1822 Bryum algovicum
P. pendulum Hornsch. 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
Hornsch., 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. Hornschuch’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 Hornschuch 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 Hornsch., Flora 5, 2: syll. 62, 1822. Lectotype
species: Ptychostomum cernuum Hornsch., Flora 5, 2: syll. 64, 1822.
In addition, Hornschuch included the species currently known as
Bryum algovicum (Brid.) B.S.G. in the genus as Ptychostomum
pendulum Hornsch., 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: Pohilia 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 B.
obtusifolium Brid., Muscol. Recent 2(3): 52. 1803.

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

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

Phytologia (Mar 2005) 87(1) 21

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, pl.
Bete 120/1803.

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. Musc. Suppl. 1, 2: 105, pl. 74, 1816.

Ptychostomum marratii (Hook. & Wils.) J.R. Spence, comb. nov.
Basionym: Bryum marratii Hook. & Wils., Bryol. Brit., p. x1
(add.), pl. XX XIIb, 1855.

Ptychostomum meesioides (Kindb.) J.R. Spence, comb. nov.
Basionym: Bryum meesioides Kindb., Bull. Torrey Bot. Club 16:
95, 1389.

Ptychostomum neodamense (Itzigs.) J.R. Spence, comb. nov.
Basionym: Bryum neodamense Itzigs. in C. Muell., Syn. Musc.
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.
Musc. Suppl. 1, 2: 107, pl. 75. 1816.

22 Phytologia (Mar 2005) &7(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.
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. Musc. Frond. Suppl.
by2e013.phs7B pips 18 ho:

Ptychostomum turbinatum (Hedw.) J.R. Spence, comb. nov.
Basionym: Mnium turbinatum Hedw., Sp. Musc. Frond. 191. 1801.

Ptychostomum warneum (RO6hl.) J.R. Spence, comb. nov.
Basionym: Mnium caespiticium var. warneum ROohl., Deutsch. FI.
(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 (Sull. & 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., Musc. 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. Musc. 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. Musc.
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) &7(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 aequis
exostomio, ciliis imperfectis vel nullis. 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 um x 12-30 um wide, margins with strongly differentiated border,
bi- to miultistratose, 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 Méielichhoferia Hornsch.
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 of Haplodontium and Plagiobryum, and very similar to

26 Phytologia (Mar 2005) 87(1)

the type of the genus, H. megalocarpum Arnott. 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. 1826. 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: 119-139.

. 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-alternate 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. Hedends. 2002. Phylogenetic relationships
between Bryum and supposedly related genera. J. Bryol. 24: 277-
289.

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

Pedersen, N., C.J. Cox, and L. Hedends. 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: 107-118.

Spence, J.R. and H.P. Ramsay. 1999. Proposal to conserve
the name Bryum (Musci, Bryaceae) with a conserved
type. Taxon 48: 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. 51. 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 32611, 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 15 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
usefully 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 multiplicity 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 19th 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.
Fernald, 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) 31

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. Witha
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
Herbarum 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 [V/2A:274-411. 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) &7(1)

species. In the most comprehensive modern 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...it 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 11 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) 87(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
prooisi::.”

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) &7(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
useful 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 modern 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 mirus Bailey and Rubus okeechobeus Bailey,
respectively. Rubus flagellaris Willd. 'Almus' and R. flagellaris 'Foster
Thornless' 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) &7(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

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; RB.
tallahasseanus Bailey; R. ucetanus Bailey; R. zoae
Bailey]

HIGHBUSH BLACKBERRY. Rubus argutus Link

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 6-12 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) 41

SIRE REET Y CHF

fe FEE & 84533 28

Le eens bk harm, KS |

peewee é :
Puuteion. 2. Sixt > tax h
Ps 2:
Dm Poe 5,

ees Bick Gate :

RE RSHRES AREACE AZ

i Sats Poke,

Fig.1. 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 Pennell (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 of Seymeria (Scrophulariaceae) from
Mexico. Phytologia 78: 28-35.

44 Phytologia (Mar 2005) 87(1)

A NEW SPECIES OF AGERATUM (ASTERACEAE;
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-corner
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

The Uobarsity of Txxax Herbarluen
O esha 3:
@. L. Torner

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. | 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, J. Solis 517 (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 CHIDR-Durango for several
years in extreme southwestern Durango and nearby Nayarit, in the
canyon country.

ACKNOWLEDGEMENTS
I am grateful 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 MELCHERTI (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 WORDS: 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 (1)-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)

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, R. 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, Mpoio.
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 7. 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) 51

Wave erry oF

» Sie
Ri orem Lend Le% Elawsts ;

ale bans, :
Revakates ox. vs Dhailaviex

Fig. 1. Trixis hintoniorum (Holotype TEX).

52 Phytologia (Mar 2005) 87(1)

TYPE: MEXICO. OAXACA: Mpio. Pochutla, Playa La Tyera,
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 7. 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 7. silvatica, but possesses thicker leaves
and larger, more numerous-flowered heads, having 11 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 7. 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 7. 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 LEMMONII VAR. 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. /. 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.1] that var. hispidula is
the more primitive of the two, var. Jemmonii 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, 1997) 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. Jemmonii. 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) &7(1)

2! hes Pe

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is , bn.

Sraare 7
200 Km ae
fy
a
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Fig 1. Distribution of Stevia 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

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

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

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 corner 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

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 published of course, but had I known the details of
its discovery I might have provided a very different Latin name!

[ 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.

58 Phytologia (Mar 2005) 87(1)

BOTANICAL HUMOR

CO a aa ache nt TT ee
all Ree ha,

*Look, Professor, | found it! The last existent erminalia uitianzal”

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. /utetiana 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)

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

Ageratum solisii B. L. Turner, sp. nov.
Anisopappinae Panero, subtrib nov.
Athroisminae Panero, subtrib. nov.

Bidens melchertii B. L. Turner, sp. nov.
Centipedinae Panero, subtrib nov.
Chromolepidinae Panero, subtribus nov.
Doroniceae Panero, tribus nov.

Dugesiinae Panero, subtribus nov.
Dyscritothamninae Panero, subtribus nov.
Enceliinae Panero, subtribus nov.

Euphrosyne acerosa (Nutt.) Panero, comb. nov.
Euphrosyne dealbata (A. Gray) Panero, comb. nov.
Euphrosyne nevadensis (M. E. Jones) Panero, comb. nov.

Haplodontium macrocarpum (Hook. ex Drumm.) J.R. Spence, comb. nov.

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

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

Plagiobryoides incrassatolimbata (Card.) J.R. Spence, comb. nov.

Ptychostomum angustifolium (Brid.) J.R. Spence & H.P. Ramsay, comb. nov.

Ptychostomum archangelicum (B.S.G.) J.R. Spence, comb. nov.
Ptychostomum arcticum (R. Br.) J.R. Spence, comb. nov.
Ptychostomum badium (Brid.) J.R. Spence, comb. nov.
Ptychostomum bimum (Schreb.) J.R. Spence, comb. nov.
Ptychostomum calophyllum (R. Br.) J.R. Spence, comb. nov.

Ptychostomum creberrimum (Taylor) J.R. Spence & H.P. Ramsay, comb. nov.

Ptychostomum cryophilum (Mart.) J.R. Spence, comb. nov.
Ptychostomum curvatum (Kaur. & Arn.) J.R. Spence, comb. nov.
Ptychostomum cyclophyllum (Schwaegr.) J.R. Spence, comb. nov.
Ptychostomum inclinatum (C. Muell.) J.R. Spence, comb. nov.
Ptychostomum intermedium (Brid.) J.R. Spence, comb. nov.
Ptychostomum knowltonii (Barnes) J.R. Spence, comb. nov.
Ptychostomum lonchocaulon (C. Muell.) J.R. Spence, comb. nov.

Ptychostomum longisetum (Bland. ex Schwaegr.) J.R. Spence, comb. nov.

Ptychostomum marratii (Hook. & Wils.) J.R. Spence, comb. nov.
Ptychostomum meesioides (Kindb.) J.R. Spence, comb. nov.
Ptychostomum neodamense (Itzigs.) J.R. Spence, comb. nov.
Ptychostomum pallens (Sw.) J.R. Spence, comb. nov.

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60 Phytologia (Mar 2005) 87(1)

Ptychostomum pallescens (Schleich.) J.R. Spence, comb. nov.
Ptychostomum pseudotriquetrum (Hedw.) J.R. Spence & H.P. Ramsay, comb. nov.
Ptychostomum purpurascens (R. Br.) J.R. Spence, comb. nov.
Ptychostomum reedii (Robins.) J.R. Spence, comb. nov.
Ptychostomum rutilans (Brid.) J.R. Spence, comb. nov.
Ptychostomum salinum (Hag. ex Limpr.) J.R. Spence, comb. nov.
Ptychostomum schleicheri (Schwaegr.) J.R. Spence, comb. nov.
Ptychostomum turbinatum (Hedw.) J.R. Spence, comb. nov.
Ptychostomum warneum (RO6hl.) J.R. Spence, comb. nov.
Ptychostomum weigelii (Spreng.) J.R. Spence, comb. nov.
Ptychostomum wrightii (Sull. & Lesq.) J.R. Spence, comb. nov.
Seymeria mazatecana B. L. Turner, sp. nov.

Spilanthinae Panero, subtribus nov.

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

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

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