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PhytoKeys 140: 107–123 (2020)

Pinguicula rosmarieae – a new butterwort from Peru 107
doi: 10.3897/phytokeys.140.49529 RESEARCH ARTICLE
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Pinguicula rosmarieae Casper, Bussmann & T.Henning

(Lentibulariaceae), a new butterwort from the
Amotape-Huancabamba Zone (northern Peru)

S. Jost Casper1, Rainer W. Bussmann2, Tilo Henning3

1 Waldpark Seniorenpflegeheim, Prellerstraße 16, D-01309 Dresden, Germany 2 Department of Ethnobotany.

Institute of Botanu, Ilia State University, 1 Botanical Street, 0105 Tbilisi, Georgia 3 Freie Universität Berlin,
Botanischer Garten Botanisches Museum, Königin-Luise-Str. 6-8, D-14195 Berlin, Germany

Corresponding author: Tilo Henning (HenningTilo@web.de)

Academic editor: E. Fischer  |  Received 19 December 2019  |  Accepted 26 January 2020  |  Published 4 March 2020

Citation: Casper SJ, Bussmann RW, Henning T (2020) Pinguicula rosmarieae Casper, Bussmann & T.Henning
(Lentibulariaceae), a new butterwort from the Amotape-Huancabamba Zone (northern Peru). PhytoKeys 140: 107–123.
https://doi.org/10.3897/phytokeys.140.49529

Abstract
The insectivorous genus Pinguicula occurs along the whole Andean mountain chain from Colombia-
Venezuela in the north to Tierra del Fuego in the south with a short interruption in the Peruvian-Chilean
desert range. This paper describes a new and striking species of Pinguicula that occurs in the south-eastern
part of the Amotape-Huancabamba Zone in north Peru. It grows either as a lithophyte on moist rocks
or as an epiphyte on Polylepis multijuga Pilg. in the wet highlands of the Cordillera Central. Pinguicula
rosmarieae Casper, Bussmann & T.Henning, sp. nov. is clearly distinguished by a basal rosette of ovate-
obovate leaves spread out flat on the ground and especially by a two-partite corolla with a straight uniform
tube-spur complex, two features unknown from other Andean Pinguicula species. The morphological sim-
ilarity to P. calyptrata Kunth is discussed and the habitat and distribution of P. rosmarieae are characterised.

Resumen
El género insectívoro Pinguicula se encuentra a lo largo Andes desde Colombia y Venezuela en el norte
hasta Tierra Fuego en el sur, con una breve interrupción en el los desiertos peruano-chilenos. Este artículo
describe una nueva y distintiva especie de Pinguicula que se encuentra en la parte sur de la zona Amotape-
Huancabamba en el norte del Perú. Puede crecer tanto como litófita sobre rocas húmedas o como epífita
sobre Polylepis multijuga Pilg. en las tierras altas y húmedas de la Cordillera Central. Pinguicula rosmarieae
Casper, Bussmann & T.Henning, sp. nov. se distingue claramente por tener una roseta basal de hojas
ovadas-obovadas, postradas sobre el suelo y, especialmente, por la corola bipartida con un espolón uniforme
recto, una combinación de características desconocidas de otras especies andinas de Pinguicula. Se discute la
similitud morfológica con P. calyptrata Kunth y se caracterizan el hábitat y la distribución de P. rosmarieae.

Copyright S. Jost Casper et al. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
108 S. Jost Casper et al. / PhytoKeys 140: 107–123 (2020)

Keywords
Lentibulariaceae, Pinguicula, Pinguicula rosmarieae, Peru, Amotape-Huancabamba Zone, Cordillera Cen-
tral, endemic, taxonomy, new species, distribution

Introduction

In South America, the insectivorous genus Pinguicula (Lentibulariaceae) is represented

by seven at first sight ± closely related, although not well-known taxa: P. antarctica
Vahl, P. calyptrata Kunth, P. elongata Benj., P. involuta Ruiz & Pav., P. jarmilae Halda
& Malina, P. nahuelbutensis Gluch and the here newly described P. rosmarieae Casper,
Bussmann & T.Henning. Forming a nearly continuous chain of ± vicarious species, the
taxa occur over a distance of 8,500 km in the Andes from the north (Colombia, Santa
Marta) via Venezuela, Ecuador, Peru, Bolivia and Chile to the extreme south (Chile,
Argentina: Tierra del Fuego), with a gap between ~30°S and ~36°S in the Peruvian-
Chilean arid region. This “Atacama”-gap has been bypassed (perhaps in former cooler
and wetter periods). In northern Peru (~ 05°30'–06°30'S), the ‘páramo-butterwort’ P.
calyptrata is more or less replaced by the litho-/epiphyte P. rosmarieae and, further to
the south, the ‘Jalca-Puna’-butterworts P. involuta and P. jarmilae close the gap.

Collection history
In 2017, searching for material of P. involuta during his study of the South American
Pinguicula taxa, the first author came across the Catalogue of the Flowering Plants
and Gymnosperms of Peru (Peru Checklist) (https://www.tropicos.org/Project/
PEC). At first sight, he believed that the specimen MO 6607881 (Paniagua Zam-
brana, Bussmann & Vega Ocaña 8586), designated as Pinguicula, could represent a
new species, but more information, especially photos and, ideally, additional material
was needed to confirm this initial assumption. The plants were gathered in November
2012 in the eastern Andes of north Peru in the area surrounding the Laguna Huaya-
bamba (“Huayllabamba”) and described as growing on a vertical rock wall in the spray
of a small waterfall. The collectors located the sampled population in the Department
La Libertad, but, instead, they were already in the adjacent Department San Martín.
Further investigations drew the attention to another, earlier collection (May 2001:
T. Henning and C. Schneider 275) deposited in the Berlin herbarium (B) at the
BGBM. They collected a violet-white flowering Pinguicula growing in large stands
on moist rock surfaces in the cliffs forming the southern limit of the Laguna de los
Cóndores (~06°51'S, ~77°42'W), ca. 20 km east of Leymebamba (B 100136109, B
100136110; duplicate specimens in HUT). The original label indicates “Departamen-
to Amazonas, Province Chachapoyas”. However, the Laguna and the adjacent area
are part of the Department San Martín and situated some 4–5 km east of the border
to the Department Amazonas. These specimens enabled the first author to conduct
 Pinguicula rosmarieae – a new butterwort from Peru 109

morphological comparisons with the material at MO for a thorough investigation. The

collections showed the same taxon from a different population some 15 km further
north in the same mountain range. The morphological differences to the other Peru-
vian Pinguicula species proved to be stable, at least in these two populations sampled
independently and, together with the reported peculiar habitat preferences, enough to
justify the description of a new species.
Finally, just very recently (2019), a third population has been reported and docu-
mented photographically by Lázaro Santa Cruz Cervera (USM) from the province of
Bongará in the Department Amazonas, some 100 km further north.

Taxonomic background
During the process of describing the new species, the most important Peruvian her-
baria in Lima (USM) and Trujillo (HUT) have been contacted in order to locate the
duplicates and gather information about potential additional collections and records.
After a review of all herbarium material that we could get access to, it became appar-
ent that most of the Pinguicula-collections from north Peru have been misidentified as
Pinguicula involuta. Instead, all collections made north of the Department Huánuco
in central Peru belong to P. calyptrata, a species traditionally referred to as a Colombi-
an-Ecuadorean taxon. This common misconception is due to the inadequate original
description and the therein indicated distribution of these two Andean butterworts.
Whereas P. calyptrata was collected by Humboldt and Bonpland (Humboldt et al.
1817) in Saraguro in the southern Ecuadorean province Loja, P. involuta was described
by Ruiz and Pavon (1798) from Huánuco in central Peru and especially the latter is
insufficiently characterised in the protologue (Casper and Hellwig 2019).
However, the distributional patterns revealed from the two previously described
and the undescribed species draw a much clearer picture corresponding to the phyto-
geographic characteristics of the so-called Amotape-Huancabamba Zone (hereinafter:
AHZ) (Weigend 2002). Instead of the (simplified) subdivision into a northern (P.
calyptrata) and southern (P. involuta) taxon whose limits roughly correspond to the
border between Peru and Ecuador, Pinguicula is present in the AHZ with two taxa, one
occurring in the west stretching northwards (P. calyptrata) and one endemic to the east-
ern slope of the Peruvian Andes (P. rosmarieae). South of the AHZ, they are replaced
by the widespread P. involuta whose distribution extends into Bolivia. The new taxon,
Pinguicula rosmarieae Casper, Bussmann & T.Henning spec. nov. is here described as
new to science. The morphology of the new species and its affinities to related taxa are
illustrated and discussed. The coarse biogeographical patterns observed for the Peruvi-
an species are outlined and explained in the context of the characteristics of the AHZ.
A distribution map, based on collection data from revised herbarium material and a
key to the Peruvian Pinguicula-species, is provided to enable a reliable determination of
existing and future collections which is a crucial component of floristic studies as basis
for urgent conservation efforts.
110 S. Jost Casper et al. / PhytoKeys 140: 107–123 (2020)

The present study is the first of a series of contributions at different stages of com-
pletion (Casper; Casper et al. in prep.), each dealing with a certain taxonomic or no-
menclatural problem that became apparent in anticipation of a comprehensive synopsis
of the South-American Pinguiculas that is in preparation and will be published soon.
Therein, all questions regarding the historical biogeography as well as nomenclatural
and taxonomic issues will be discussed extensively.

Materials and methods

The first author examined herbarium specimens of Pinguicula L. from South America
in preparation for a revision of the Andean Pinguiculas at HUT, USM, MO, B and
M (Thiers 2019) and from specimen scans using online databases (www.tropicos.org).
This study combines the results of the herbarium studies, the experiences of the col-
lectors (R.W. Bussmann and T. Henning) in the natural habitat and observations and
reports kindly received from the Peruvian colleagues.

Results
Key to the Peruvian species of Pinguicula

1 Foliage not star-like, leaf blades oblong-obovate-ovate, with margins slightly

(mostly ~2 mm) curled up; corolla bi-partite, i.e. divided in the lip and the
straight, more or less uniform tube-spur complex (the funnel-shaped tube
merges into the conical blunt spur, with little to no angle); living on water-
rinsed sandstone rocks or as epiphyte on Polylepis twigs; cloud forest (eastern
North-Peru)............................................................................. P. rosmarieae
– Foliage star-like (“stellate”), leaf blades ovate, with margins distinctly curled
up (appearing boat-shaped); corolla tri-partite, i.e. divided distinctly in lip,
tube and spur, i.e. the tube distinctly angled with the spur...........................2
2 Corolla with nearly equal-sized notched lobes (subisolobate, i.e. lobes of the
upper-lip only slightly smaller than those of the lower lip); ~2 mm behind the
lower lip middle lobe base, a prominent clapper-like palate covered by yellow
hairs inserted; the tube typical funnel-shaped; the spur short, conical, blunt;
páramo–jalca (North-Peru)........................................................P. calyptrata
– Corolla with distinctly unequal-sized notched lobes, (i.e. lobes of the upper-
lip lobes distinctly smaller than those of the lower-lip); the lower-lip middle
lobe dominating the lip, often distinctly bent down; no distinct palate in-
serted; the tube cylindrical, nearly as long as wide; the spur slender, sickle-
shaped, pointed; puna (Central- and South-Peru).........................P. involuta
 Pinguicula rosmarieae – a new butterwort from Peru 111

Taxonomic treatment
Pinguicula rosmarieae Casper, Bussmann & T.Henning, sp. nov.
urn:lsid:ipni.org:names:77206951-1
Figs 1 A–H, 3A

Type. Peru: Dept. San Martín, Prov. Huallaga [“Dept. Amazonas, Prov. Chachapoyas”
(sic!)], Lagunas de los Cóndores (06°50'40.5"S, 77°41'52.2"W), 3,000 m a.s.l. Violet-
white flowers, diameter up to 100 mm. Lithophyte, 24 May 2001, T. Henning & C.
Schneider 275. (Holotype: HUT 41126!: det. Pinguicula spec. – Figs 1H, 3A; Iso-
types: B: B 100136110 (Fig. 2A)! B 100136109!)
Diagnosis. Herba perennis rosulata, rosula ca. 100 mm in diametro; lamina folio-
rum circuito suborbiculata vel obovata, margine vix (~3 mm) involuta, ca. 30–40 mm
longa ac lata, solum plusminusve adpressa; scapus 1(–4), erectus, 20–40 mm altus, teres;
flores 1, parvi, ~8–10 mm longi (tubo-calcari incluso), bilabiati; calyx lobis oblongis,
lobis superis usque ad basin fere divisis, lobis inferis usque ad dimidium divisis, ad an-
gulum ~45° divaricatis; corolla lobis 5, oblongis, ~5–7 mm longis, subisolobatis, apice
valde emarginatis; corollae tubum infundibuliformi-cylindraceum cum calcari conico
± uniformem rectum coniunctionem formans, 5–6 mm longum apice obtusum.
Habitatio in locis apertis et humidis montium Andinensium regionis Peruviae sep-
tentrionalis usque ad 3.100 m supra mare, praesertim ad rupes et saxa. Habitu Pin-
guiculae Andinensium simili, praecipue differt tubo-calcari-coniunctio recto uniformi.
Description. Perennial rosette leaved herb with 1 (–4) flowered scapes. Rhizome ~
10 mm long, with numerous adventitious fibrous roots. Leaves (4–) 6–10, flat on the
ground, ± succulent (dried translucent-membranous), adult (20–) 30–40 (–50) mm
long, nearly as long as wide, the blades ovate-obovate-oblong in outline, rounded at
the tip attenuated to the base into a short petiole, the margins weakly (up to 3 mm)
curled up, yellowish-green, upper surface of lamina covered with sessile glandular
hairs. Hibernacula (winter buds, dormant buds) absent. Scapes 1–2 (–4), erect, (20–)
30 (–40) mm tall, terete, filiform (0.5–1 mm thick), one–flowered, green to reddish-
brown, scattered with glandular hairs, often becoming glabrous or nearly so. Flowers
small, ~8–10 (–11) mm long (including tube-spur-complex). Calyx two-lipped, green
to pale brown to purple, upper surface of sepals scattered with stalked glandular hairs;
upper lip divided deeply into three nearly equal-sized oblong lobes, at apex pointed;
lower lip up to ½ divided into two oblong lobes, at apex pointed. Corolla two-lipped,
bluish-magenta to white-violet; upper lip two-lobed, lobes obovate, ~5–6 mm long
and ~2–3.5 mm wide, shallowly notched at the apex; lower lip larger and longer than
the upper lip, with three oblong to obovate-oblong lobes (the median lobe somewhat
larger than the two lateral ones), 4.5–5 mm wide, each distinctly (~1/5 to 1/3 of its
length) notched. Tube (tube-spur-complex) at the throat funnel-shaped, on both sides
broader than the spur, on the back side higher than the spur, proximally cylindrical
112 S. Jost Casper et al. / PhytoKeys 140: 107–123 (2020)

Figure 1. Pinguicula rosmarieae (A–H) and P. calyptrata (I, J).

 Pinguicula rosmarieae – a new butterwort from Peru 113

(nearly as long as wide), on the ventral side merging without any sharp angle into the
cylindrical to cone-like stubby, at apex rounded, yellow-greenish spur; tube and spur
forming a more or less uniform funnel- to cone-like straight, from the ventral side
appearing as a box-like ‘tube-spur-complex’, ~6 mm long; the tube-spur-complex exter-
nally dark blue to purple lengthwise-striped by parallel veins. Palate bipartite, weakly
developed (not clapper-like), inserted immediately behind (~1–2 mm) the corollas´
lower-lip middle lobe, yellow, set with short-stalked glandular hairs, proximally elon-
gated into a short ventral hair strip; each of the two lateral corolla lobes with a small

Figure 2. Specimen scans of Pinguicula rosmarieae and P. calyptrata. A Isotype of P. rosmarieae – Scan
(Henning & Schneider 275, B100136110), with permission B paratype, from vertical sandstone cliff east
of the Laguna Huayllabamba, 3,250 m a.s.l., the three flowers (left) showing corollas with their significant
straight uniform tube-spur-complex. The rosette is composed of flat outspread leaves. – Scan of MO
6607881, with permission C P. calyptrata, the leaves have curled up margins and the corollas are typical
tri-partite in corolla, tube and spur. The corolla lobes are nearly equal-sized, the tube funnel-shaped and
the spur short and angled with the tube. – Scan MO 6589755; with permission.

Figure 1. Continued. A P. rosmarieae, 2-scaped; upper flower opened, in profile view, lower one in
bud; epiphytic on Polylepis multijuga Pilg., in Polylepis multijuga-Iochroma stenanthum S.Leiva, Quip. &
N.W.Sawyer – dominated cloud forest. Peru, Department San Martín, close to ‘El Jardín’(Inca-hut and sur-
rounding area east of the Laguna Huayabamba), 3,090 m a.s.l., 06°56'044"S, 077°41'54"W B ditto, flow-
er, profile view C ditto, flower, semi-ventral view D ditto, flower, dorsal view E P. rosmarieae rosette from
the northernmost known habitat, “Hatumpampa” Department Amazonas, Province Bongará (no voucher
specimen) F ditto, flower, semi-ventral view G ditto, flower ventral view H corolla in frontal view, lower-lip
lobes to ¼ of its length notched, throat without distinct palate. Peru, Department San Martín, Laguna de
los Cóndores (Henning & Schneider 275) I P. calyptrata, corolla in frontal view, lobes with lateral margins
slightly covering each other, lower-lip lobes to 1/6 of its length notched, throat with clapper-like yellow
palate. Peru, Department San Martín, Sphagnum-bog, 3,000 m a.s.l. above the Laguna de los Cóndores
(Bussmann, A. Glenn, G. Chait & C. Vega Ocaña 16447) J flowering stand of P. calyptrata near Pulan,
Cajamarca. (Credits: photographs A–D, I R. W. Bussmann E–G, J L. Santa Cruz Cervera H T. Henning).
114 S. Jost Casper et al. / PhytoKeys 140: 107–123 (2020)

yellow hair bubble at their base, stretching proximally along on each side of the inner
tube wall. Indumentum (apart from that visible in the photographs), stamens, pollen
grains, ovary, stigma, capsule, seeds not seen. Chromosome number unknown.
Pinguicula rosmarieae is distinguished by its notable uniform funnel-cone-shaped
straight tube-spur-complex, a feature unknown in any other Andean Pinguicula taxon.
It is an endemic species, restricted to the eastern slopes of the Cordillera Central within
the Amotape-Huancabamba-Zone of northern Peru (Departments Amazonas and San
Martín; Fig. 4). It grows in rocky habitats, either under moving water or in the spray
of small waterfalls and occasionally epiphytic on moss-covered twigs of Polylepis multi-
juga, at altitudes of about 2500 m–3100 m a.s.l. (Fig. 3).
Etymology. The new species is named after Dipl.-Biol. Rosmarie Casper, beloved wife
and steady companion of the scientific efforts of S. J. Casper and mother of their children.

Discussion
Affinities

Based on current floristic literature, Pinguicula involuta was considered as the only but-
terwort native in northern Peru. However, a revision of the available herbarium material
in this study has revealed that this simple assumption, according to the historical taxo-
nomic treatments, P. involuta is native to Peru whilst P. calyptrata is native to Ecuador, is
largely incorrect. Furthermore, the “northern” Pinguicula of Humboldt and Bonpland
(i.e. P. calyptrata), that was first discovered in the Saraguro range in southern Ecuador,
has repeatedly been collected in northern Peru. We have seen specimens of P. calyptrata
from the Peruvian Departments Piura, Amazonas, San Martín, Lambayeque and Ca-
jamarca (Fig. 4, cf. the list of specimens below). In turn, P. involuta, so far, has never
been collected in these Departments, although misidentified collections and erroneous
floristic literature suggest that. Based on our observations and fieldwork, the north-
ernmost locality of P. involuta is in the Department Huánuco in central Peru, some
350 km further south of the southernmost P. calyptrata localities (Province Pachitea,
Panao; road from Chaglia [Chaglla] to Rumichaca [Tambo de Vaca], km 81, 09°51'S,
75°53'W), leg. M. Weigend, K. Weigend, T. Henning, & Ch. Schneider 5426, Fig. 4).
At present, the exact distribution limits of P. involuta and P. calyptrata remain uncertain.
Morphological data indicate that the new taxon is distinct from P. involuta and P.
calyptrata. A thorough study of South American Pinguicula is in preparation and will
elucidate the morphological affinities amongst all relevant taxa.
Foliage: The leaf blades of P. rosmarieae are ovate-obovate-oblong, dried membranous-
translucent (Fig. 2A) and spread out, i.e. they are not boat-like and they lack heavily
curled-up margins (Fig. 1A, E): The whole rosette (up to 100 mm across) is not star-
shaped compared to P. calyptrata (Figs 1J, 2C) and P. involuta (field name of the latter in
the original collection was P. stellata, see Casper and Hellwig 2019). The foliage shape of P.
rosmarieae is similar to the geographically-distant P. albida Wright that occurs in Cuba and
 Pinguicula rosmarieae – a new butterwort from Peru 115

may represent an individual adaptation to the rocky, exposed habitats. Both habitats share
nutrient-poor and often acidic white-sand-soils (for Peru: M. Weigend, pers. comm.).
However, this observation is contradicted by the fact that the rosettes of P. calyptrata and
P. involuta retain their star-like appearance on relatively open stands. Shape and posture of
the foliage are, therefore, taxonomically meaningful for species delimitation.
Corolla: The violet-white to pale-bluish corolla is similar to that of most Andean
Pinguicula: its lips are nearly equal-lobed (subisolobate). The two lobes of the upper-lip
are (mostly only shallowly) notched at the distal margins (Fig. 1B–D, F–H), the three
lobes of the lower-lip are longer and at its distal margins deeply notched (up to ¼ –1/3
of its length; Fig. 1G–H). The most striking feature that separates P. rosmarieae from
other Andean Pinguicula taxa is the more or less uniform straight tube-spur-complex
that appears as almost entire. The typical tri-partite divided Pinguicula corolla (i.e. into
corolla lobes, tube and spur) appears nearly bi-partite here and is divided only into the
corolla lobes and the tube-spur complex. The funnel-shaped, cylindrical tube merges
ventrally into the short and comparatively wide cone-shaped spur, lacking a sharp
angle (Fig. 1A, B). The spur is stubby, rounded at the apex and light yellow-green.
The back of the tube only weakly protrudes from the spur: from this point, the stubby
spur slightly attenuates proximally and ends in an obtuse apex. Looking at the corolla
from a lateral (Fig. 1B) or semi-ventral (Fig. 1C) perspective, the tube and spur are not
markedly separated. The tube-spur-complex is lengthwise dark parallel-veined. A tri-
partite yellow to white palate is only weakly developed (Fig. 1H, i.e. non clapper-like
as, for example, in P. calyptrata – Fig. 1I) and placed immediately behind the corolla
lower-lip middle lobe (that appears as a shallow yellowish-greenish shimmering dent
on the ventral tube-side directly behind the base of the middle lobe).
To illustrate the differences that can be observed on herbarium specimens, we have
chosen Pinguicula calyptrata (Bussmann et al. 16447 MO, barcode: MO 6589755,
Fig. 2C) collected in the immediate neighbourhood of the P. rosmarieae-type popula-
tion (Fig. 2A). The sheet shows three well-preserved, single-scaped flowering specimens
with the flowers in lateral profile view. The leaf rosette with its curled-up leaf margins
measures up to ~30 mm in diameter. The flowering scapes are up to ~50 mm tall; the
flowers are up to ~12 mm long (spur included), the corolla is distinctly tri-partite into
the lip, tube and spur; its lobes are nearly equal-sized (subisolobate), the lower-lip mid-
dle lobe is only slightly larger than the lateral lobes, ~5 mm long; the corolla tube is
distinctly funnel-shaped (at throat widest, ~5 mm), about as long as the corolla lobes,
~5 mm long; the spur is distinctly separated from the tube, short, ~2.5 mm long, thin,
at apex rounded (sometimes almost imperceptibly thickened), angled at about 60°–90°
with the length axis of the tube. Overall, in P. calyptrata as in the other Andean Pin-
guicula-taxa, the corolla is distinctly divided into three parts: lip, tube and angled spur.
Contrarily, in P. rosmarieae, the corolla appears bi-partite: tube and spur form a
straight uniform funnel-cone-shaped tube-spur-complex, i.e. the spur is not angled
with the tube. These features are clearly visible on the specimen collected by Paníagua-
Zambrana, Bussmann & Vega Ocaña 8586 (MO 6607881; Fig. 2B), gathered in the
surroundings of the Laguna Huay(ll)abamba (Department San Martín). The plants
116 S. Jost Casper et al. / PhytoKeys 140: 107–123 (2020)

were found growing either as lithophytes on a vertical rock wall in the spray of a small
waterfall or as epiphytes on twigs of Polylepis multijuga. They are distinguished by
a spread-out 6–8 leaved rosette appressed to the ground. Its flower shows the strik-
ing straight uniform tube-spur-complex which we also observed in the Laguna de los
Cóndores-Pinguicula (see the three specimens on the left-hand side of the sheet). Pho-
tographs (Fig. 1H, I) of the corollas (frontal view) support the deep morphological
disparity between the two taxa.
In P. rosmarieae (Fig. 1H), the corolla is widely open and appears radially sym-
metrical at first sight. The corolla lobes are spread out, the two lobes of the upper lip
are smaller than those of the lower-lip, their distal margins are shallowly notched (to
~1/6 of their length). The lower-lip lobes are much larger, deeply notched (to ~¼ –1/3
of their length). The throat and the adjacent tube portion are not dark, a pronounced
palate is not developed; it is replaced by a weak yellowish shimmering patch at the base
of the corolla lower-lip middle lobe, continuing proximally (to the middle of the tube-
spur-complex) lengthwise in two white hairy stripes.
In P. calyptrata (Fig. 1I), the corolla is widely open and also appears radially sym-
metrical. The corolla lobes are spread out, covering each other slightly with their lateral
margins. They are nearly equal in size, except for the middle-lobe of the lower-lip that
dominates the corolla to a certain degree. The distal margins of all corolla lobes are only
shallowly notched, the throat being dark-purple coloured. Nearly 2 mm behind (proxi-
mally) the middle lobe, at the base of the lower-lip, a pronounced yellow clapper-like
palate is inserted, from which two white hairy stripes stretch out into the tube.

Habitat
The Departments Amazonas and San Martín in northern Peru partly occupy the Sierra
zone between the dry coastal region (Costa) and the upper Amazon river lowlands
(Selva) and are largely characterised by extensive and very species-rich, cloud forests
and wet subalpine grasslands (páramos). In contrast to Pinguicula calyptrata, which is
largely found on wet, often peaty, soils, in the páramo region, Pinguicula rosmarieae
occupies a completely different, even wetter, habitat. The species has been found either
growing on steep, often vertical, rock-walls, normally on sandstones, in the spray of
waterfalls (Fig. 3A, B) or rarely as an epiphyte in dense moss layers on Polylepis multi-
juga (Fig. 3C). Both represent equally extreme habitats, with extremely wet, nutrient-
poor and acidic conditions and considerable mechanical stress.
The population at the type location grew in full sunlight on a steep sandstone cliff
immediately above the famous tombs built by the Chachapoyas culture (AD ca. 800–
1500). The tombs were built underneath natural overhangs, thereby allowing dry stor-
age of the mummies. The type population (Henning & Schneider 275) grows above
these overhangs exposed to constant dripping water and the general high precipitation
typical for the eastern slopes of the Andes in this region (Fig. 3A).
 Pinguicula rosmarieae – a new butterwort from Peru 117

Figure 3. Habitats of Pinguicula rosmarieae in the Department San Martín. A Large stands at the type
locality above the Laguna de los Cóndores B sandstone rock walls with small waterfall near ‘El Jardín’
C ‘El Jardín’ Polylepis multijuga stands with P. rosmarieae growing as an epiphyte. (Credits: A T. Henning,
B, C R. W. Bussmann).
118 S. Jost Casper et al. / PhytoKeys 140: 107–123 (2020)

Distribution
Pinguicula rosmarieae is endemic to the northernmost foothills of the Cordillera Central
in northern Peru. The herbarium specimens, known so far, are from three nearby col-
lections in the same mountain range southeast of Leymebamba, stretching over some
15 km in a north-south direction. Both expedition teams have mistakenly located the
collection sites in the adjacent western Departments (Amazonas and La Libertad, re-
spectively), since the border runs along the pass of the Cordillera. According to current
online map-sources (Google maps), all populations, deposited in herbaria so far, were
found some 4–6 km east of the border to San Martín (Fig. 4). The presumed narrow
endemism has just recently been rebutted by the report of the new taxon from the
Province of Bongará in the Department Amazonas some 100 km further north. Photo-
graphs made by L. Santa Cruz Cervera clearly show the characteristic rosette and flower
patterns of P. rosmarieae, photographed in a similar open habitat near the famous Gocta
falls. However, the taxon seems restricted to the northern branches of the Cordillera
Central, but reaches further into the northernmost foothills. The collection data indi-
cate that P. calyptrata and P. rosmarieae show distributional overlap over the entire range
of the latter. True sympatry is nevertheless prevented by the different habitat require-
ments of the two taxa.
The area lies well in the so-called Amotape-Huancabamba Zone, an important
biodiversity-hotspot that spans from the Pacific coast over the cordilleras to the tropical
lowlands of southern Ecuador and large parts of northern Peru (Fig. 4, for details see:
Weigend 2002, 2004;). Many plant groups have a centre of diversity here (Weigend et
al. 2005; Struwe et al. 2009; Deanna et al. 2018) and show a concentrated occurrence
of narrow-endemic taxa in that region (Berry 1982; Ayers 1999; Weigend 2002; Hen-
ning and Weigend 2009, 2009a; Henning et al. 2019). While the vegetation and flora
of the inner-Andean valleys and the western slopes are relatively well-investigated, the
eastern flanks of the Cordillera Central facing Amazonia are still under-collected in
many areas. Especially, the areas east of Leymebamba (Dept. Amazonas) in the north
and Buldibuyo (Dept. La Libertad) in the south remain largely unexplored, since the
eastern slopes can only be reached by foot. Single collection trips have yielded un-
expected, supposedly narrowly endemic taxa in other plant groups, although only a
tiny fraction of the area could be sampled to date (e.g. Loasaceae: Nasa rugosa subspp.
gracilipes and pygmaea; Henning et al. 2011).
The whole region is characterised by great geological diversity, with large areas of dolo-
mitic karst, caused by the exceptionally high rainfall (the indigenous Chachapoya were
often called “warriors of the clouds”), interspersed with small areas of sandstone outcrops
and metamorphic rocks. The vegetation of the sandstone areas is particularly intriguing,
with many endemic species and unique vegetation types (e.g. Weinmannia sp. and Polylepis
multijuga-Iochroma stenanthum dominated forests). P. rosmarieae has been found exclusively
in the spray of waterfalls on sandstone cliffs and as an epiphyte on Polylepis – both systems
characterised by extreme moisture and almost permanent cloud cover, a fact that is recog-
nised in the local topographic maps, which often simply indicate “clouds” in these areas.
 Pinguicula rosmarieae – a new butterwort from Peru 119

Figure 4. Distribution of Pinguicula rosmarieae and related taxa based on geo-referenced collections.
Green stars: P. rosmarieae, red dots: P. calyptrata, yellow square: P. involuta, dotted line: northern and
southern limits of the Amotape-Huancabamba-Zone.
120 S. Jost Casper et al. / PhytoKeys 140: 107–123 (2020)

Pinguicula is yet another example of a plant group that is present in the Amotape-
Huancabamba Zone with an endemic taxon, whereas its widespread relatives have
their distribution limits at the zonal boundary, either seen from within (P. calyptrata)
or outside (P. involuta). Since Pinguicula rosmarieae has been collected in those areas
of the overall region that are comparatively easy to access, it cannot be ruled out that
there are additional taxa awaiting discovery. However, the nondescript shape of the
insectivorous butterwort, represented by only a few, widespread taxa outside of that
region (Fig. 4), does not encourage gathering.
On a broader scale, the endemism of P. rosmarieae is by no means a unique phe-
nomenon amongst the South American Pinguicula. P. jarmilae Halda & Malina from
southeast Bolivia (Department Chuquisaca, municipio Villa Serrano, village Nuevo
Mundo), for example, is only known from the type locality (Halda et al. 2007). Al-
though it grows on vertical sandstone (laterite) rock faces under flowing/moving water,
its morphology is quite different from that of P. rosmarieae: the superficial similarity is
likely coincidental and does not reflect a close relationship between the two taxa.

Preliminary conservation status

Due to the aforementioned lack of a continuous botanical exploration of the region,
we have to consider P. rosmarieae as Data Deficient (DD), according to the IUCN
threatened species assessment guidelines (2001, 2017). However, within its distribu-
tional range, it seems to be limited to the easternmost ridge of the Cordillera Central.
As an insectivorous plant, it seems to be adapted to special habitat types, characterised
by ‘open’ vegetation, high precipitation and nitrogen-poor soils. These habitats might
only be available in a narrow altitudinal band along the mountain chain. As our ob-
servations suggest, such sites are not present at similar altitudes in the west and it is
very likely that suitable habitats are only available a few kilometres further east up to
the lower altitudes of the Andes. This might indicate that P. rosmarieae is generally a
rare taxon and might, thus, be vulnerable to habitat destruction. The lack of additional
collections emphasises this assumption. This delicate butterwort might be in danger of
extinction, but more data is needed to better assess the entire distribution and abun-
dance, especially to the east of the known populations.

Annotated list of P. rosmarieae specimens (paratypes)

Peru. “Dept. Amazonas: Distr. Chachapoyas” (sic!) = Dept. San Martín, Prov. Hual-
laga, Lei[y]mebamba, Oseres [10 km east of Leymebamba]. Bosque Montano, 2542
m a.s.l. (~06°58'S, ~77°40'W), 22 May 2015. C. Vega Ocaña, L. Cotrina P., J. Valle, R.
W. Bussmann, & N. Paniagua Zambrana 247 – HAO, MO 2852736. dp! [Det. Pin-
guicula RBU 2015; involuta E. Feltz (Ma), 2016]. – Duplicate: MO 6726506. “Dept.
La Libertad, Distr. Uchumarca” (sic!) = Dept. San Martín, Prov. Huallaga, páramo and
 Pinguicula rosmarieae – a new butterwort from Peru 121

sandstone cliffs east of Laguna Huay(ll)abamba (06°58'53"S, 77°43'09"W) 3250 m

a.s.l., 02 Nov 2012. – N. Paníagua Zambrana, R. W. Bussmann & C. Vega Ocaña 8586
[det. Pinguicula involuta RBU 2015]. – MO 6607881. dp!

List of P. calyptrata specimens in the research region (used for Fig. 4)

Peru: Dept. Piura: [Prov.] Huancabamba, Lomas Redonda (Sapalache-Chinguelas)
2400 m (05°09'S, 79°26'W) a.s.l., 15 Ago 1981. A. Sagástegui Alva & al. 10187 – MO
2940293 [Barcode 348636], MO 4025844 [Barcode 348635] (dp!), HUT (dp!) – (det.
P. involuta P. Taylor & M. Cheek). Dept. Lambayeque: [Prov.] Ferreñafe, [Distr.] Inca-
huasi cerca a la laguna Tembladera. Vegetación de jalca, zonas humedas. 3300 m a.s.l.
(~06°09'S, ~79°19'W). 08 Oct 1989. S. Llatas Quiroz 2606 – F 2051195 (dp!). (det. P.
involuta P. Taylor 1992). Dept. Cajamarca: Prov. San Ignacio, Distr. Tabaconas, Local.
Santuario Nac. Tabaconas-Namballe, alrededores de las lagunas Coyona (Arrebiatadas),
05°14'S, 79°16'W (Arriba Laguna Lagartocha), 3140 m–3180 m a.s.l.. S. M. Baldéon
Malpartida 5108 & L. Adriazon Ocupa. – USM 00266675 dp! (det. P. involuta). – Ca-
jamarca, km 30 de la carretera Cajamarca-Bambamarca Jalca, estepa de gramineas, so-
bre un afloramiento rocoso. 3600 m a.s.l., 23 Mar 1985. – Sánchez-Vega, I. M., U. Mo-
lau & L. Ohmann 3756. – F 2216084 (Barcode V0469923F - dupl. CPUN; dp! det.
Pinguicula L.) – [Prov.] Santa Cruz, [Distr.] Pulán, [caserio] El Molino, 2,500 m a.s.l.
(06°46'S, 78°55'W), 12 Feb 2007. L. Santa Cruz, M. Chocce & M. Beltrán 996– USM
241552 (dp!), HUT 50771 (dp!). – [Prov.] Santa Cruz, [Distr.] Pulán, Pampa el suro,
2500 m a.s.l. (06°50'S, ~78°54'W), 31 Ene 2008. L. Santa Cruz 2098 – HAO, USM
240752 (dp!). – Prov. San Miguel, distrito Tongod, Bosque San Pedro Norte (06°45'S,
78°49'W), 03 Nov 2001. I. M. Sánchez Vega & M. Sánchez M. 11122 – F 2245015 [V
0410057F] (dp! det. P. involuta). Dept. Amazonas: [Prov.] Bagua, [Santuario Nacion-
al] Cordillera [de] Colan-La Peca, 9600 ft a.s.l. (05°35'S, 78°14'W), 29 Aug 1978. Ph.
J. Barbour 3222 – USM 53585 (pd!), MO 2798203 (dp! – Barcode MO 348631, det.
P. involuta P. Taylor 1992). – Cordillera Colan-La Peca, 9600–11075 ft a.s.l. (05°35'S,
78°16'W), 08 Sep 1978. Ph. J. Barbour 3425 – F 1909251 (V0469936F; dp!), MO
2789874 (pd! det. P. involuta Taylor). – Amazonas: 3000–4000 m a.s.l. (~06°07'S,
~77°39'W), 09 Nov 2012. H. van der Werff, L. Valenzuela, G. Shareva & A. Reyes Bar-
rantes 25401 – MO. – Cerro de Fraijaca (Huaui-Huni) n. e. Tambo de Ventilla. Jul 07
1948. A. W. Pennell 15875 USM 90946 (det. Pinguicula - dp!). – Prov. Chachapoyas,
declives superiors de [Cerro] Puma-Urcú, [~2 km] este-sureste de [ciudad] Chacha-
poyas (~06°14'S, ~77°52'W), alt. 2700 m. – 3000 m., Jun 01 1962. J. J. Wurdack 679.
– USM 90948 (dp!), COL (dupl. dp! det. P. antarctica, by Fernandes-Pérez 1964), NY.
– Prov. Chachapoyas, bosque bajo y húmedo al lado de (moist scrub forest on south
side of ) Molinopampa-Diosan pass, alt. 2700 m – Balsa road to Leymebamba, just
below Abra Callacalla (= Alba Barro Negro) on the slope towards Leymebamba, 3559
m a.s.l. (06°44'S, 77°53'W), 19.10.2000 – M. Weigend, E. Rodríguez R., H. Förther
& N. Dostert 867 – USM 166766 (dp! det. Pinguicula spec.) – [Prov.] Chachapoyas,
122 S. Jost Casper et al. / PhytoKeys 140: 107–123 (2020)

[Distr.] Balsas, En el Paso de Calla Calla (06°48'S, 77°53'W), 07 Oct 2001. I. M.

Sãnchez-Vega, M. Sãnchez, M. 11061 – F. – Cerros [Cordillera] Calla Calla. 26 km
above Leimebamba, road to Balsas. Km 403, 3360 m a.s.l. (~06°48'S, ~77°53'W), 16
Oct 1964. P. C. Hutchison & J. K. Wright 6993. – MO 2233627 (GBIF: as P. antarc-
tica); USM 90947 (dp!). – Prov. Chachapoyas, Distr. Chachapoyas. Trail to Laguna de
Los Cóndores, surroundings of Laguna Esperanza/Siete Lagunas (06°49'S, 77°43'W)
3275 m–3500 m a.s.l., Jun 26 2010. R. W. Bussmann, A. Glenn, G. Chait & C. Vega
Ocaña 16447. Dept. La Libertad: Distr. Uchumarca, páramo in the surroundings of
Vira Vira/Lagunas La Quinuas (07°00'S, 77°45'W), 3050 m a.s.l. – Photograph: R. W.
Bussmann (det P. involuta = P. calyptrata).

Acknowledgements
Thanks are due to the curators and staff of the herbaria listed above, in particular Asunción
Cano (USM), Eric F. Rodríguez-Rodríguez (HUT) and Dr Robert Vogt (B). Especially, we
are indebted to Prof Dr Maximilian Weigend (Bonn), Prof Dr Frank Hellwig (Jena), Dr
Hermann Manitz (Jena) and Dr Katy E. Jones (Prague) for additional pieces of informa-
tion, for reading and criticising our manuscript and for the supply of literature. Further-
more, we would also thank Dr Rafael Acuña Castillo for help with the Spanish translation
of the abstract. Photos and habitat data, kindly provided by Lázaro Santa Cruz Cervera
(Lima), significantly added to the understanding of the new species´ distributional range.
Eric F. Rodríguez R. and a second, anonymous referee are thanked for their profound
review of the manuscript. We finally acknowledge the support by the German Research
Foundation and the Open Access Publication Fund of the Freie Universität Berlin.

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