STUDIES OF NEOTROPICAL COMPOSITAE-VIII.

THE NEW COMBINATION PSEUDONOSERIS GLANDULOSA

AND REVISION OF PSEUDONOSERIS (LIABEAE)

John F. Pruski

Missouri Botanical Garden

P.O. Box 299
St. Louis, Missouri 63166

ABSTRACT

The new combination Pseudonoseris glandulosa (Compositae: Liabeae: Paranepheliinae) is
proposed and Pseudonoseris revised with two species recognized, both found in Andean Peru and
Bolivia only in and south of the Huancabamba Depression. Both species of Pseudonoseris were
described originally in Onoseris (Mutisieae). Also included are images of holotypes, representative

specimens, and capitula of both species of Pseudonoseris, as w ? ell as a distribution map.

KEY WORDS: Andes, Asteraceae, Bolivia, Compositae, Huancabamba Depression, Peru, Liabeae,
Onoseris, Pleistocene climatic oscillations, Pseudonoseris discolor, Pseudonoseris glandulosa,
Mutisieae, Paranepheliinae

Liabeae was described by Rydberg (1927) to accommodate five genera, including Liabum
Adans. (Compositae), an opposite-leaved radiate-capitulate American genus having vernonioid styles
(Cassini 1823). The styles of Liabeae resemble technically those of Vernonieae and Cichorieae by
their adaxiai continuous stigmatic surfaces and abaxiai branch sweeping papillae extending
downward onto the distal portion of trunk (Robinson 1983; Pruski & Sancho 2004). Robinson and
Brettell (1974) recognized 14 genera of Liabeae with several segregates culled mostly from Liabum s.
lat, including Pseudonoseris H. Rob. & Brettell (Compositae: Liabeae: Paranepheliinae). Tribe
Liabeae now contains 19 genera, with 13 of these genera occurring in Peru, its center of diversity
(Robinson 1978, 1983; Pruski & Sancho 2004; Soejima et al. 2008; Robinson & Funk 2011).

Robinson (1983) placed only Pseudonoseris and Paranephelius Poepp. into his newly
described subtribe Paranepheliinae. Funk et al. (2012) expand Paranepheliinae to include six genera,
including Erato DC. which extends into Central America. Soejima et al. (2008) dated origins of the
Pseudonoseris-Paranephelius ciade to the Miocene (very approximately to 13 Ma). Thus,
Pseudonoseris and Paranephelius (or their immediate precursors) diversified well after the significant
central Andean uplifts (Oligocene, ca. 30 Ma.), perhaps concurrently with major northern Andean
orogeny (Pliocene-Pleistocene, ca. 3-5 Ma) and increased habitat heterogeneity in both the central
and northern Andes (Soejima et al. 2008).

The central and northern Andes are discontinuous roughly at the Huancabamba Depression
(approximately between 5° and 6 C S latitude in the deep upper Rio Maranon and Rio Chamaya basins,
continuing westward over the Abra de Porculla at ca. 2145 m, and down to Olmos), an oft-cited
biogeographic barrier foi some plant distributions as well as an endemism center' (Wurdack 1953;
Simpson 1975; Molau 1988; Funk et al. 1995; Avers 1999; Weigend 2002). It is perhaps noteworthy
that, although climatic fluctuations in the Pleistocene lowered (in glacial maxima) vegetational zones
facilitating some migrations across the Huancabamba Depression, core subtribal Paranepheliinae
members Pseudonoseris and Paranephelius remain known from only in and south of the
Huancabamba Depression (Robinson 1978, 1983; Soejima et al. 2008). While these two genera may
have radiated only after the onset of Pleistocene climatic oscillations, the Huancabamba Depression
may still have acted (either well before or during the Pleistocene) as a barrier to migration.

Pruski: Revision of Pseudonoseris {Uabeae) 2

Figure 1. Representative specimen of Pseudonoseris discolor (Muschl.) H. Rob. & Brettell {Araujo 4194, MO).

Figure 2. Close-sip of capitula of Pseudonoseris showing She linear-lanceolate phyllaries and the lor.g-stipitale-
glandular peduncles. A Pseudonoseris discolor (Muschl.) R Rob. & Brettell. B. Pseudonoseris glanduiosa
(Hieron.) Pruski. (AAraujo 4194, MO; B Smith & Cabaniilas 7303, MO). [The scale bar on the right has

Here, Pseudonoseris is revised, the new combination P. glanduiosa is proposed, and this
high-elevational Andean genus is recognized as containing only two species. Pseudonoseris striata
and P. szyszylowiczii were each usually recognized elsewhere (Robinson & Brettell 1974; Robinson
1983; Beltran et al. 2006; Soejima et al. 2008), but both are reduced here to synonymy of P.
glanduiosa. Pseudonoseris glanduiosa and P. discolor are disjunct from each other by about 1100
km (Fig. 3).

PSEUDONOSERIS H. Rob. & Brettell, Phytologia 28: 59. 1974. TYPE: Liabum striatum Cuatrec. [=
Pseudonoseris glanduiosa (Hieron.) Pruski].

Small perennial subscapose herbs, leaves in basal rosettes or stems leafy proximally, with
milky latex, scape lypically with a low arachnoid-pubescence and long s tip itate- glandular trichomes
held well above arachnoid pubescence. Leaves opposite, sometimes in basal or cauline rosettes,
sessile; blade subeotire to lyrate-pinnatilobed, lanceolate to obovate, chartaceous to stiffly so,
venation pinnate, surfaces discolorous, adaxial surface smooth to rugose, pubescent to glabrate,
abaxial surface cinereous -griseoiis-tonieiitose (sometimes drying fulvous), base slightly clasping.
Capitulesccnce subscapose, pauci capitulate, open cymose with alternate branching; peduncles
elongate, long-stipitate-glanduiar. Capitula radiate; involucre campanulate; phyllaries ca. 40, weakly
imbricate., graduate, 3-5-seriate, linear -lanceolate, iong-stipitate-glandu'ar, outer ones acute apically,
often spreading in iruit, inner ones long -attenuate apically; clinanthium (phoranthiurn or receptacle)
epaleate, iow~alveolate, subglabrous. Ray Berets 12-26. pistillate, 1 -seriate; corolla orangish-yellow
to scarlet, tube pilose-villous, limb 1 in ear- ob lanceolate, exseiled from involucre, abaxially glabrous
and not arachnoid-i.omentulose, usually 4-nerved and 3 -denticulate. Disk Burets 25-55, bisexual;
corolla gradually funnelform, 5-lobed, orangish-yellow or reddish- orange, hirsute near tube-throat
juncture, tube elongate, lobes glabrous or with a single distal gland; anther tliecae pale, base obtuse,
ecaudate, endothecial tissue polarized, apical appendage ovate to oblong, slightly longer than wide;
pollen tricoiporafe, echinate. tectum finely mi crop erf orate, coiumeliate layer well-defined (Robinson
& Marticorena 1986); style base very slightly dilated, glabrous, trunk slightly papillose distaliy.

brandies elongate and nearly filiform, 3-4 mm long, narrowing distally, abaxial surface rough-
papillose, adaxially with continuous stigmatic surface. Cypselae prismatic to obovoid, thickly 10-
costate (merely striate when immature), costae often tan and setose with nearly appressed trichomas,
furrows usually brown and glabrous, cells walls with elongate raphids (Robinson 1983); pappus
distinctly or indistinctly 2-seriate, mostly with an inner series of 15-30 elongate stramineous scabrid
capillary bristles, several mid-sized outer squamellae typically present, these sometimes absent
adaxially and represented instead by small outer bristles, x = 12. Two species found in Andean Peru
and Boh via.

Pseudonoseris is a central Andean genus (Fig. 3) of two species that Robinson (1983) treated
within Liabeae subtribe Paranepheliinae. Pseudonoseris is diagnosed by its subscapose habit, milky
latex, discolorous pinnately-veined sessile leaves, indumentum of long-stipitate-glandular trichomes
on the scape, peduncles, and phyllaries, erect alternate-branched capitulescences, brightly colored ray
corollas with limbs abaxially glabrous and not arachnoid-tomentulose, pale anthers, and filiform style
branches. Color photographs of each species in the field were provided by Soejima et al. (2008). The
two species of Pseudonoseris were each once treated within sometimes stipitate-glandular, brightly
colored, and large-capitulate Onoseris (Mutisieae), which differs most conspicuously by outer florets
with bilabiate corollas, tailed anthers, and short-ovate style branches.

„J

A Pseudonoseris discolor

• Pseudonoseris glandulosa

1 * <
V * (

\ * i_

N \

t ^

350 700 km ^^\^

III T

Figure 4. Photograph? of the destroyed Berlin holotypes of the two recog
Onoseris discolor Muschl [= Pseudanoseris discolor (Muschl.) I-I. Rob. i
Her od. [= Pseudorioseris glandubosa (Hieron ) Pruski]

Although in the Soejima ei. ai. (2008) study the relationships betweeo Pseudorioseris
glandulosa (as 'Pseudorioseris szyszylowiczif) and Paranephelius are unresolved, Pseudorioseris is
maintained here as diagnosed morphologically by Robinson (1983). Pseudorioseris differs from
Paranephelius most obviously by some stipitate- glandular (vs. eglandular) indumentum, erect
branched (vs. usisaliy sessile) capitulescences, and by abaxially glabrous (vs. arachnoid-tomentulose)
ray corolla limbs. Cytologically, the base number of Liabeae appears tobtx = 9, Paranephelius was
reported as x = 9 and x =14. and Pseudorioseris has been counted only once, being given as x = 12
(Dillon & Turner 1982; Robins on et al. 1985).

Typification and etymology. The genus, name for it:; vesemb lance to Onoseris, is typified by
Liohum striatum, the holotype of which is extant. The holotypes (Fig. J) of our two species were
destroyed, but neither species circums crip lion is in doubt. Thus, there is no pressing need
taxonomically to lecSoiypify or neotypify either species name, and I am content to wait for possible
holotype fragments or isotypes to be found The descriptive basionym epithets discolor.

Pruski: Revision of Pseudonoseris (Liabeae) 6

glandulosum, and striatum refer to the salient characters of Pseudonoseris of discolorous leaves, the
distal 'tipitate-Ma.iduLn m.liinientutn and the striate-costate cypselae, respectively.

1. Leaves simple, never lyrate-pinnatilobed, adaxial surface rugose to rugulose; ray corollas orange or

sometimes orangish-yellow; cypselae with pappus indistinctly biseriate Pseudoiioseris discolor

1. Leaves lyrate-pinnatilobed, adaxial surface smooth; ray corollas scarlet or reddish-orange, at least
some cypselae with an obviously biseriate pappus adaxially Pseudonoseris glandulosa

PSEUDONOSERIS DISCOLOR (Muschl.) H. Rob. & Brettell, Phytologia 28: 60. 1974. Onoseris
discolor Muschl., Bot. Jahrb. Syst. 50, Beibl. Ill: 94. 1913. Liabum lanatum Ferreyra, Bol.
Soc. Peruana Bot. 1: 17. 1948 (non Liabum discolor (Hook & Arn.) Berth. & Hook, f ex
Hemsl.). TYPE: PERU. Puno. Inter Sandia et Cuyocuyo, 2600-2800 m, 1 May 1902,
Weberbauer 883 (holotype: B|, photograph in F, MO, Macbride neg. 15889). Figs. 1, 2A,
4A

Rosulate or less commonly leafy-stemmed herbs 10-70 cm tall; stems arachnoid-
pubescent, also moderately dense long-stipitate-glandular (trichomes to 1.8 mm long and sometimes
longer than stem diam.) to near base, intemodes not hirsute and without simple patent trichomes, all
leaves basal or proximal-cauline, when cauline with leaves paired with internodes 1^1 cm long but
always much shorter than the leaves. Leaves 4-24 x 1-6 cm, lanceolate to oblong, broadest at mid-
blade or slightly above mid-blade, simple, never lyrate-pinnatilobed, adaxial surface rugose to
rugulose, usually lingering arachnoid-pubescent especially along midrib and secondary veins,
sometimes glabrate, broad-based or more commonly base narrowly acute, margins unequally sinuous-
dentate to less commonly crenate or subentire, apex acute to sometimes obtuse. Capitulescence 1-2
per plant, held 9-50 cm above leaves, loosely cymose, 2-7-capitulate, branched in distal half;
peduncles usually 2-10 cm long modoiaUl\ deme long-ihpikite-jhndul i sometimes uith 2-3
linear bracteoies 1-3 mm long. Capitula 14-16(-18) mm long (excluding rays); involucre 10-13 x
8-18 mm; phyllaries 3-13 x 1-2 mm, with a purplish mid-zone and scarious margins. Ray florets
12-26; corolla orange or sometimes orangish-yellow, tube 10-13 mm long, limb 20-24 x 2.5-3 mm.
Disk florets: corolla 11-13 mm long, orangish-yellow, lobes 2-3 mm long. Cypselae 2-3.2 mm
long; pappus indistinctly biseriate, inner pappus bristles 15-25, 8-10 mm long, outer pappus bristles
few, 0. 1-0.3 mm long, not obviously much broader than inner series of bristles.

Representative exsiccatae examined. BOLIVIA, La Paz. Franz Tamayo, Parque Nacional
Madidi, Keara-Moxos, Kellutoro, 3000 m, 13 May 2008, Araujo 4143 (BM, LPB, MO), Araujo 4194
(B, GH, LPB, MO); Franz Tamayo, ANMI Apolobamba, sector Laitiki hacia Piara. entre Pelechuco y
Apolo, 2650 m, 20 Apr 2006, Fuentes et al. 10422 (LBP, MO, NY). PERU. Puno. Sandia, Limbani,
3300 m, 7 .Tun 1974, Chavez 2382 (MO); Sandia, dry open hillside near Limbani, 3200-3450 m, 14-
16 May l942,Metcalf 30531 (MO, UC).

Distribution and ecology. Pseudonoseris discolor occurs in puna from 2600-3450 meters
elevation, north of Lake Titicaca in the Andes of Puno, Peru, and adjacent La Paz, Bolivia (Fig. 3).
The species is saxicolous and flowers from April to June.

Illustrations of Pseudonoseris discolor w~ere provided by Ferreyra (1948) and Robinson
(1983). Pseudonoseris discolor was given by Robinson and Brettell (1974), Robinson (1983),
Beltran et al. (2006), and Soejima et al. (2008) as endemic to Peru, but this species is now known
from several collections in adjacent Bolivia. The protologue described it as similar to Onoseris
glandulosa |= P. glandulosa]. The few inner pappus bristles and the indistinctly biseriate pappus are
distinctive features of P. discolor. Some color photographs of P. discolor appear to show the ray
corollas as orangish-yellow, but the plants are usually described as orange-flowered.

Pruski: Revision of Pseudonoseris (Liabeae) 7

PSEUDONOSERIS GLANDULOSA (Hieron.) Praski, comb. nov. Onoseris glandulosa Hieron., Bot.
Jahrb. Syst. 21: 366. 1895. TYPE: PERU. Cajamarca. Prope La Cruz de Celendin, inter
Pacasmayo et Moyobamba, 3100 m, Apr-Jun 1868-1877, Stubel 35h (holotype: B|,
photograph in F, MO, Macbride neg. 15890). Figs. 2B, 4B, 5.

Liabum szyszylowiczii Hieron., Bot. Jahrb. Syst. 36: 503. 1905. Pseudonoseris szyszylowiczii
(Hieron.) H. Rob. & Brettell, Phytologia 28: 60. 1974. TYPE: PERU. Cajamarca. Prope
Callacate, May 1879, Jelski 718 (holotype: B1% photograph in F, MO, Macbride neg. 18133).

Liabum striatum Ciiatrec, Collect. Bot. (Barcelona) 3: 306. 1953. Pseudonoseris striata (Cuatrec.)
H. Rob. & Brettell, Phytologia 28: 60. 1974. TYPE: PERU, [presumably near the
Lambayeque-Piura border near Abra de Porculla], Above Olmos, 1800-1900 m, May 1915,
Weberbauer 7707(holotype: F, photograph in MO, Field neg. 49222).

Rosulate or leafy-stemmed herbs 20-90 cm tall; stems arachnoid-pubescent, also long-
stipitate-glandular (trichomes to 1 mm long and sometimes longer than stem diam.) distally, grading
to hirsute with simple patent trichomes proximally, all leaves basal or proximal-cauline, when cauline
then in cauline-rosettes or with leaves paired with internodes 1-5 cm long but always much shorter
than the leaves. Leaves 6-18 x 2.5-10 cm, lyrate-pinnatilobed with terminal lobe the largest and
usually with 2-3 pairs of lateral lobes, oblong to obovate in outline, adaxial surface smooth,
hirsutulous to substrigillose with patent or subappressed trichomes, also sometimes arachnoid-
pubescent, broad-based or less commonly base attenuate, lobes nearly lateral, triangular-ovate with
sinuous-denticulate margins, sinuses rounded, apex obtuse to sometimes acute. Capitulescence 1-
2(-3) per plant, usually held 15-30 cm above leaves, loosely cymose, usually 2-9-capitulate,
branched only in distal half; peduncles usually 2-15 cm long, more densely pubescent than stem and
proximal capitulescence axis. Capitula 12-19 mm long (excluding rays); involucre 10-15 x 10-20
mm; phyllaries 3-15 x 1-2 mm, linear-lanceolate, appearing 2-costate at least near mid-phyllary.
Ray florets 13-21; corolla scarlet or reddish-orange, tube 10-11 mm long, limb 15-20 x 2-3 mm.
Disk florets: corolla 9-12 mm long, reddish-orange, lobes 2-2.5 mm long. Cypselae 2^1 mm long;
with pappus obviously (at least adaxially on some cypselae) biseriate, inner pappus bristles 25-30, 6-
8 mm long, outer squamellae 0.5-1.5 mm long, at least the adaxial ones on some cypselae noticeably
broad-based and obscuring inner bristle bases, the abaxial ones especially of the ray florets sometimes
represented by very small bristles. 2n = 24 (Dillon & Turner 1982).

Representative exsiccatae examined. PERU. Amazonas. Chachapoyas, encima de
'Leimebamba', 2600-2700 m, 16 Apr 1964, Ferreyra 15465 (MO, USM); 6 kms along road W of
Chachapoyas, 6600 ft, 13 Jan 1983. King & Bishop 9198 (MO, US); 15 km from Chachapoyas
towards Mendoza, 2200 m, 13 Mar 1998, van der Werff et at. 14833 (MO, US); 1 km SW of
Chachapoyas, 2300 m, 22 May 1962, Wurdack 467 (MO-2, US); Chachapoyas, km 422-417 on
'Leymebamba'-Balsas road, 2400-2700 m, 21 Feb 1984, Smith 6077 (MO). Ancash. Huari, 2500 m,
2 May 1962, Ames 7 (MO). Cajamarca. Jelij, grassland, 3035 m, 16 June 2009, Bussmann et al.
15526 (MO, NY); Jaen, Sallique, de Catala a Piquijaca, 1940-2195 m, 29 Jul 1998, Campos & Diaz
5396 (MO, US); Carretera entre Socota y Cutervo, 2000-2200 m, 20 Apr 1988, Diaz & Baideon 2844
(F, MO); 26 km NW [on maps this appears instead to be NE] of Celendin on road to Balsas, 2300 m,
5 Jan 1979, Dillon & Turner 1699 (F, MO, US); Yamaluc, entre Cochabamba y Huambos, 2300-
2500 [annotated in pencil as 1900-2000] m, 1 Aug 1946 (post fruit), Ferreyra 828 (MO); Km 156 de
la carretera Pacasmayo-Cajamarca, 2650 m, 5 Apr 1982, Sanchez-Vega 2758 (MO); Chachapoyas-
Celendin road, above Celendin, 3000 m, 28 May 1984, Smith & Cabanillas 7303 (MO).
Lambayeque. The type of synonymous Liabum striatum, presumably near the Lambayeque-Piura
border near Abra de Porculla. Piura. Huancabamba, Porculla. 2200 m, 10 May 1992, Llatas & Cruz
3 1 06 (F, MO).

(Keren.) Pruski. (Smith & Cabanillas 7303,

Pruski: Revision of Pseudonoseris {tiabeae) 9

Distribution and ecology. Pseudonoseris glcmdulosa is endemic to Andean Peru in and south
of the Huancabamba Depression, where it occurs in both Cordilleras, these separated by the Rio
Maranon (Fig. 3). Pseudonoseris glandulosa occurs mostly in the montane zone and in jalca
formations (sometimes it is saxicolous) from 1800-3400 meters elevation in the Departments of
Amazonas, Ancash, Cajamarca, Lambayeque, [expected in La Libertad], and Piura and flowers from
January to July. Although P. glandulosa presumably did not migrate across Huancabamba
Depression during Pleistocene climatic oscillations, it is moderately successful ecologically as it
occurs on the Cordilleras both west and east of Rio Maranon valley. Dorobaea laciniata B. Nord. &
Pruski and Talamancalia putcalensis (Hieron.) B. Nord. & Pruski (Nordenstam & Pruski 1995;
Beltran & Pruski 2000) are other regional subscapose lobe-leaved large-capitulate orange-flowered
taxa (both are Senecioneae) resembling superficially P. glandulosa.

The protologue noted specifically that this taxon lacked the inner corolla lips typical of
Onoseris (Mutisieae). Pseudonoseris glandulosa was excluded from Onoseris by Ferreyra (1944).
who referred provisionally this species to Liabum. Pseudonoseris glandulosa was not treated
subsequently in the Mutisieae in the Flora of Peru (Ferreyra 1995) nor elsewhere in die Mutisieae
(Katinas et al. 2008). By opposite leaves, radiate capitula, moderately long-lobed disk corollas,
ecaudate anthers, style trunks distally papillose, and style branches with continuous stigmatic
surfaces, P. glandulosa keys to tribe Liabeae in ftuski and Sancho (2004), where it matches
Pseudonoseris by its subscapose habit, indumentum of some long-stipitate- glandular trichomes, erect
alternate-branched capitulescences, brightly colored ray corollas, pale anthers, and filiform style
branches. Soejima et al. (2008) gave this species as having arisen through hybridization between
Pseudonoseris and Paranephelius. Cytologically, P. glandulosa was reported by Dillon and Turner
(1982) as In = 24, the only count published for the genus.

No significant consistent morphological differences are found between plants of (1) the
Pacific slopes of the Cordillera Occidental -the coastal range- (e.g., the presumed type locality of P.
striata), (2) those of the interior slopes Cordillera Occidental just west of the Rio Maranon (generally
between 6° and 7° S latitude) of Depto. Cajamarca (e.g., the types of P. glandulosa and P.
szyszylowiczii), (3) those east of the Rio Maranon in the Cordillera Oriental in Depto. Amazonas near
Chachapoyas, or (4) with those much further south in Depto. Ancash. Weigend (2002) noted that
although species are often restricted distribution lly to one side of the Andes, this generalization may
beak down in the upper Rio Maranon valley where several groups are found in the coastal as well as
interior mountains. Although plants of P. glandulosa from the Cordillera Occidental on average tend
to have a more obviously biseriate pappus man, for example, do plants of the Cordillera Oriental from
near Chachapoyas, this pappus variation is never as great as thai found within some individual florets
and within some individual capitula. Accordingly, both P, striata and P, szyszylowiczii are treated in
synonymy of P. glandulosa, lowering from three to two the number of species recognized in
Pseudonoseris.

ACKNOWLEDGEMENTS

I would like to thank Guy Nesom and Rosa Ortiz (MO) for reading the manuscript, Stephanie
Keil (MO) for taking the photographs of the MO sheets, Christine Niezgoda (F) for permission to use
the photographs of the Berlin holotypes, and Rosa Ortiz for help in preparing the map.

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