Viola lilliputana sp. nov. (Viola sect. Andinium, Violaceae), one of the world's smallest violets, from the Andes of Peru

Abstract

A new violet species of Viola Sect. Andinium,
Viola lilliputana, is described from a single dry puna locality on an extensive intermontane plateau southeast of Cerro Palla Palla in the high Andes of Ayacucho Department in southern Peru. This diminutive rosulate violet is evidently among the smallest in the world and probably one of the smallest terrestrial dicots. It belongs to a distinctive species group with pinnatifid leaves that is endemic to central and southern Peru, including V. hillii, V. membranacea and V. weibelii. The new species is similar to V. weibelii in its large, strongly adnate stipules, elongate leaf lobes and dilated unappendaged style with ventral stigmatic orifice. It differs conspicuously from all other members of the pinnatifid-leaved group in its conduplicate leaf blades, straight, mostly nonoverlapping, oblong-lanceolate to broadly elliptical lobes with obtuse to rounded apices, and large basally fused pedicel bractlets. Despite many new collections of vascular plants from the high Andes of Peru and northern Bolivia in recent decades, this distinctive new species is still known only from its type locality, collected on the Iltis-Ugent expedition from November 1962 to January 1963.

Full text

Viola lilliputana sp. nov. (Viola sect. Andinium, Violaceae),
one of the world's smallest violets, from the Andes of Peru
H
ARVEY
E. B
ALLARD
,J
R
.
1
AND
H
UGH
H. I
LTIS
2
1
Department of Environmental and Plant Biology, Ohio University, 315 Porter Hall, Athens, OH
45701-2979, U.S.A.; e-mail: ballardh@ohio.edu
2
Department of Botany, University of Wisconsin-Madison, 132 Birge Hall, Madison, WI 53706,
U.S.A.; e-mail: swis@charter.net
Abstract. A new violet species of Viola Sect. Andinium, Viola lilliputana, is descr-
ibed from a single dry puna locality on an extensive intermontane plateau southeast of
Cerro Palla Palla in the high Andes of Ayacucho Department in southern Peru. This
diminutive rosulate violet is evidently among the smallest in the world and probably
one of the smallest terrestrial dicots. It belongs to a distinctive species group with
pinnatifid leaves that is endemic to central and southern Peru, including V. hillii,V.
membranacea and V. weibelii. The new species is similar to V. weibelii in its large,
strongly adnate stipules, elongate leaf lobes and dilated unappendaged style with
ventral stigmatic orifice. It differs conspicuously from all other members of the pin-
natifid-leaved group in its conduplicate leaf blades, straight, mostly nonoverlapping,
oblong-lanceolate to broadly elliptical lobes with obtuse to rounded apices, and large
basally fused pedicel bractlets. Despite many new collections of vascular plants from
the high Andes of Peru and northern Bolivia in recent decades, this distinctive new
species is still known only from its type locality, collected on the Iltis-Ugent exped-
ition from November 1962 to January 1963.
Key Words: Andes, new species, Peru, Sect. Andinium,Viola.
The genus Viola encompasses 525–600
species (Clausen, 1964; Ballard et al., 1998),
with centers of diversity in the Andes and
southern Patagonian cone of Latin America,
mountains of eastern Asia, Melanesia, and the
mountains of southern Europe. Taxonomic
knowledge in Europe and Asia is relatively
good; in that area, new investigations on
violets have gradually moved toward those
focused on phylogenetic relationships and
evolutionary origins. This is not true of
montane regions in central and southern
South America, where alpha taxonomy
remains a very active pursuit.
Four currently recognized sections of Viola
are distributed primarily or exclusively in
South America: sect. Andinium W. Becker,
sect. Leptidium Ging., sect. Rubellium W.
Becker and sect. Chilenium W. Becker. The
largest group of Viola in this region requiring
extensive systematic scrutiny is Sect. Andi-
nium, one of the most primitive in the genus
(Ballard, et al., 1998; Marcussen et al., 2011)
and the earliest to invade high-mountain
habitats, comprising probably more than 100
species. Various specialists and lay taxono-
mists have provided many names in an effort
to capture the taxonomic diversity in this
fascinating group, represented almost entirely
by rosulate or subterranean-stemmed and
often diminutive herbs thriving in some of
the most forbidding terrestrial vascular plant
habitats on earth. Due to extreme environ-
mental selection (e.g., divergent daytime and
nighttime temperatures, the latter commonly
reaching subzero values, and sporadic to
practically nonexistent precipitation), many
species take on a diminutive “pincushion”
growth form. The smaller taxa usually require
a hand lens if not a dissecting microscope to
Brittonia, 64(4), 2012, pp. 353–358 ISSUED: 1 December 2012
© 2012, by The New York Botanical Garden Press, Bronx, NY 10458-5126 U.S.A.

view diagnostic features or even to confirm
correct assignment to the family. Whole
species groups, such as the Viola cotyledon
Ging. assemblage, often have several to many
species differing only by subtle floral charac-
ter-states, including morphological features of
the style, the latter commonly tipped by
lateral and/or terminal protuberances, crests,
horns or other processes differing in each
species. While some species exhibit a fairly
widespread distribution (often crossing coun-
try borders), the majority are quite narrow to
highly localized endemics, in some cases
known only from a single mountain range—
or the type locality. One unusual species
assemblage in the central Andes of Peru
encompasses a small number of pinnatifid-
leaved taxa, the only Andinium violets with
conspicuously lobed leaf blades. The present
report briefly characterizes this fascinating
group of tiny violets and describes a distinc-
tive new member confined to a single locality
in a remote high-elevation plateau of southern
Peru.
Circumstances of the Discovery
From November 1962 to January 1963
Hugh Iltis, his student Don Ugent and their
spouses made an expedition to southern Peru
to find potatoes in the Andes, to add new
germplasm to the potential variability of
cultivated potatoes (Ugent, 1970; Iltis,
1982), and to collect vouchers and study
specimens of cacti and other plants in the
region. The small group traveled from the
coast of Peru, near Nazca, east-northeastward
across the western range of the Andes to
Cuzco, by way of the then-poorly developed
"road" passing through Puquio and Chal-
huanca. Cross-referencing of journal entries
for habitat descriptions and settlements
against features on high-resolution topo-
graphic maps and aerial photographs have
identified the exact route taken by Iltis’s
group and each stopping point, as well as
the exact collecting locality of the new violet
species.
On 14 December, 1962, after camping out
above and passing eastward through the
bustling town of Puquio, they headed up on
to the road to Chalhuanca. This turned out to
be a very rough road, where progress was
extremely slow and sometimes treacherous,
with steep inclines on either or both sides.
Finally reaching Laguna Yaurihuiri at nearly
13,000 feet, they passed a restless night
suffering from sirroche (mountain sickness,
with headaches, extra heartbeats and fatigue),
in tents buffeted by strong winds. Early on
the morning of 15 December, they packed up
and left Laguna Yaurihuiri heading east on
the two-track, driving spellbound through
miles of treeless arid puna comprising a large
intramontane plateau situated in the center of
the western slope of the Andes. The puna was
dotted liberally by short bunchgrasses as well
as scattered small herbs of many families,
including Azorella diapensioides A. Gray and
Lilaeopsis andina A. W. Hill (Umbelliferae),
Gentiana and related genera (Gentianaceae)
and Astragalus species (Leguminosae); and
mounded cushions of Pycnophyllum (Caryo-
phyllaceae), Aretiastrum (Valerianaceae) and
white-fuzzy Mniodes coarctata Cuatrec.
(Asteraceae). Wherever there was seepage or
water, the species diversity was enriched by
the addition of enormous hard cushions
several yards across and half a yard high, of
Distichia muscoides Nees & Meyen or Oxy-
chloe andina Phil. (both Juncaceae), often
intermixed with various Boraginaceae and
Umbelliferae. The puna was a surprisingly
rich landscape on a lilliputian scale.
After passing Lagunas Pucacocha and
Islacocha, with their multitudes of brilliant
pink-orange flamingos, the road led them by
the tiny settlement of Negro Mayo, the only
one since Puquio. Approximately 12 kilo-
meters beyond Negro Mayo, and 6 kilometers
east of the boggy road crossing of a small
creek issuing from Laguna Parccoccocha at
the base of Cerro Palla Palla, the company
halted in the late afternoon to watch some
vicuñas cavorting in bare sand patches of the
puna. The intermontane plateau here con-
sisted of rocky alpine tundra strewn with
huge scattered basalt boulders (Fig. 1), dotted
with sporadic cushion plants and tiny single-
tufted plants growing amid much bare soil
and gravel.
It was here that serendipity brought Iltis
and the tiny new species of violet face to
face. Crouched to photograph two vicuñas
reveling in a dust bath, Iltis momentarily
dropped the haze filter off his camera (the
354 BRITTONIA [VOL 64

filter being nearly clear, it immediately dis-
appeared from view after rolling a few feet
away). Hunting for the filter on hands and
knees, he soon found it—atop a tiny patch of
plants appearing as seedlings of some type of
Caryophyllaceae. Iltis pulled up a few tiny
tufts from their droughty lodging and, under
the hand lens, saw that they were in flower.
Upon evening-time inspection in the tent, he
realized that the diminutive plants must be
violets—the smallest he had ever seen or
heard of. Shortly after returning to the
University of Wisconsin-Madison in early
1963, he examined the handful of violet
specimens more thoroughly and investigated
Andean violet names that might match the
identity of these fascinating little plants,
settling on V. hillii W. Becker or V. weibelii
W. Becker with pinnatifid leaves as close but
not wholly satisfactory matches.
The Violet's Status is Clarified
The matter was left unresolved until nearly
30 years later in 1992, when Ballard, who
was then a new Ph.D. student at UW-
Madison Botany Department, met Iltis while
conducting revisionary studies on Latin
American Viola. Together, the authors delved
anew into species descriptions of Wilhelm
Becker (1906,1909,1922,1928), violet
treatments for Peru by Baehni and Weibel
(1941) and Macbride (1941), and protologues
of previously published names for other New
World violets. Over the next decade the
first author located and examined type
material and herbarium collections of all
of the potentially relevant names for South
American violets and evaluated species
circumscriptions of described species
against Iltis's tiny species. He determined
that the latter was both utterly distinct and
undescribed.
Viola lilliputana H. H. Iltis & H. E. Ballard,
sp. nov. Type: Peru. Dept. Ayacucho:
Along 'road' between Puquio (Dept. Aya-
cucho) and Chalhuanca (Dept. Apurimac),
Laguna Yauriviri, 12,000–14,000 feet or
more, top of Altiplano, 15 Dec 1962, H. H.
Iltis & D. Ugent et al. 518 (holotype: MO;
isotypes: K, USM, WIS). (Fig. 2)
F
IG
.1. Xeric puna in vicinity of Viola lilliputana population, showing characteristic scattered rubble and
diminutive cushion plant flora of the high Andean "alpine tundra." Photo by H. H. Iltis.
355BALLARD
&
ILTIS
:
VIOLA
(
VIOLACEAE
)
2012]

Inter species Sectionis Andinii W. Becker habitu
rosulato similis atque ad Violam weibelii J. F. Macb.
foliis pinnatifidis, lobis lateralibus elongatis, margin-
ibus setosis, stipulis grandibus adnatis, stylo dilatato
non appendiculato, orificio stigmatico ventrali acce-
dens, sed laminis conduplicatis, lobis lateralibus
plerumque oblonge lanceolatis vel ellipticis rectis,
apice rotundato, bracteolis pedicellatis grandibus
basaliter connatis differt.
Acaulescent perennial herbs, up to 1.1 cm
tall. Rootstock slender, vertical, unbranched
with single crown or branched near top and
cushion-forming with 2–3 coherent crowns.
Leaves spiral, in a rosette, long-petiolate.
Stipules persistent, adnate to petiole ½–3/4 of
their length, scarious, lanceolate, the apex
acuminate, the margin entire, 6–9×0.4–
1.5 mm, the outer surface glabrous, the margin
ciliolate or glabrous. Petiole 3–4 mm long,
glabrous. Entire leaf 8–15×2.1–3.5 mm; blade
length:width ratio 3.6–4.7 (when flattened), the
leaf blade not decurrent, the lamina herbaceous,
conduplicate, smooth or rugulose, lanceolate in
broad outline, deeply pinnately lobed, the upper
and lower surfaces glabrate to glabrous, the
margin along lobes entire, involute or flat,
sparsely to moderately setose; terminal lobe
2–3×1–1.5 mm, conspicuously larger than the
lateral ones, ovate to rhombic, the apex
narrowly rounded, the base narrowly cuneate
and symmetrical; lateral lobes mostly nonover-
lapping, 6–7 on each side, 0.8–1.5 ×0.25–
0.7 mm, mostly oblong-lanceolate to narrowly
elliptical or slightly asymmetrically rhombic,
occasionally broadly elliptical to suborbicular,
with obtuse to rounded apices. Pedicels 5–
10 mm long, glabrous; bractlets persistent,
fused together at base, attached near middle of
pedicel, scarious, lanceolate to linear-lanceo-
late, acuminate, entire, 3–4×0.5–1 mm, gla-
brous, the margin ciliolate. Flowers 1.6–2mm
long; sepals subequal in size and shape, the
surface smooth, veins none, lanceolate, acumi-
nate, 1.15–1.7×0.35–0.6 mm, glabrous
throughout, the auricles rounded, 0.1–
0.15 mm long, glabrous. Corolla:calyx length
ratio 0.6–0.8; corolla color unknown; upper
petals spreading, broadly oblong to broadly
elliptic, the apex acutish to rounded, 1.5–2×
0.6–0.8 mm, glabrous throughout; lateral petals
spreading, broadly oblong to broadly elliptic,
the apex acutish to rounded, 1.5–2×0.6–
0.8 mm, glabrous throughout; bottom petal
porrect, elliptic, the apex rounded with margins
somewhat conduplicate, 1.7–2×0.9–1.2 mm,
the outer surface glabrous throughout, the inner
surface papillate basally near throat; spur
laterally thickened, distinctly broader than tall,
deflexed downward, 0.8–1.2× 0.5–0.7 mm;
F
IG
.2. Viola lilliputana.A. Individual resting on a U.S. penny. B. Style, ventral view (top) and lateral view
(bottom). C. Flower in profile. D. Leaf with broader lateral lobes spread out, showing setose (and often involute)
margins. E. Leaf with more typical oblong-lanceolate lateral lobes. (From the type.) Photos and illustration by H. E.
Ballard, Jr.
356 BRITTONIA [VOL 64

bottom petal blade:lateral petal blade length
ratio ca. 1, bottom petal blade:spur length ratio
1.83–2.22. Stamens 0.9–1 mm long; anthers
sessile, ca. 0.45× ca. 0.4 mm, the outer surface
glabrous, the glands on bottom pair of stamens
sessile, very slender, long-conical. Dorsal con-
nective scales broadly oblong to orbicular, the
apex rounded or truncate, 0.45–0.5 × 0.25–
0.5 mm, glabrous, the margin entire, scale:
anther length ratio 1–1.05. Ovary glabrous.
Style base straight, the body dilated toward
apex, straight, unappendaged laterally, gla-
brous; stigmatic orifice apico-ventral in a
longitudinal furrow; style ca. 0.75 mm long.
Capsules and seeds unknown.
Distribution and ecology.—Details in the
second author's collection notebook, as described
in the introductory narrative, concur with site-by-
site post hoc georeferencing to place the location
of the new violet in the general vicinity of
14.6365°S, 73.6740°W. This corresponds to
6 km distance by road east of the small stream
intersecting the Puquio-Chalhuanca truck trail
north of Laguna Parccoccocha and appearing to
issue from that lake, on an extensive intermontane
plateau stretching from the base of (and southeast
of) the peak of Cerro Palla Palla, in Dept.
Ayacucho. This is a slightly different location
than suggested by the "mass produced" labels.
The violet is known only from this type locality,
where it inhabits dry gravel amid scattered
basaltic rubble and boulders, as shown in Fig. 1.
Phenology.—Flowering in December, pos-
sibly some time before and afterward; fruiting
likely in January and later. (No flowers had
yet shown sufficient post-pollination enlarge-
ment to indicate imminent fruiting.)
Etymology.—The specific epithet makes a
fanciful reference to the tiny growth form of
the new species as “lilliputian”. The term
represents the race of tiny people much
smaller than Gulliver, the main character in
Book 1 of Gulliver’s Travels authored by
Jonathan Swift, in which Gulliver became
shipwrecked on the isle of Lilliput and
experienced various (mis)adventures.
Viola lilliputana is morphologically similar
to a small group of other Andinium violets with
deeply pinnatifid leaves, all distributed in
central and southern Peru and immediately
adjacent Bolivia: V. hillii W. Becker (poly-
morphic species complex), V. membranacea W.
Becker and V. w e i b e l ii J. F. Macbr. Its glabrate
to glabrous leaf laminas (but sparsely to
moderately setose margins), very large mem-
branous linear-lanceolate stipules and dilated
unappendaged style with ventral stigmatic
orifice (Fig. 2)allyittoV. weibelii. However,
its conduplicate leaf blades, more or less
straight (or slightly asymmetrically rhombic)
nonoverlapping leaf lobes with rounded apices,
and very large basally fused pedicel bractlets
(often hiding the associated flower buds) set it
apart from all described species in the pinnat-
ifid-leaved group. In broader comparisons with
the world’sknownvioletflora the new species
also appears to be one of the world’s smallest
violets (if not the smallest) and surely stands as
one of the smallest terrestrial dicots, with the
entire aboveground plant body scarcely topping
1 cm in height—rivaled in stature only by
sporadic and particularly diminutive individu-
als of certain species in the annual V. parvula
Tineo complex (wild pansies of Sect. Melanium
W. Becker) of southeastern Europe and adja-
cent western Asia. This distinctive and highly
localized little violet is hereby named and
distinguished from other members of the
pinnatifid-leaved Andinium group.
Key to species in the pinnatifid-leaved group of Viola Sect. Andinium
1. Leaf lobes irregular in distribution and size, quadrate, shorter than broad, not distinctly narrowed at base, the
apices truncate; a number of sites in central Peru (Depts. Ancash and Junín)………………V. membranacea
1. Leaf lobes regular in distribution and size, oblong-lanceolate, asymmetrically rhombic or elliptical, at least
the middle and lower longer than broad, sometimes distinctly narrowed ("subpetiolate") at base, the lobe
apices attenuate, acute, obtuse or rounded; central and eastern Peru, and northernmost Bolivia.
2. Foliage, pedicel and calyx moderately to densely pubescent; at least the upper leaf lobes no more than
twice as long as broad, incurved distally with apex sharply acute, or with lobes incurved and
asymmetrically trapezoidal with an apparent medial or basal auricle on inner margin (this actually
representing the apex of a strongly hooked lobe); stipules small and inconspicuous, free, ca. 1 mm long;
style various……………………………………………………………………V. hillii (species complex)
2. Plant glabrous or with pubescence restricted to conspicuous setae along leaf blade margins; all leaf lobes
elongate, commonly 2 or more times as long as broad, straight to medially incurved-falcate, the apex
357BALLARD
&
ILTIS
:
VIOLA
(
VIOLACEAE
)
2012]

attenuate or narrowly to broadly rounded; stipules 6–9 mm long, large and conspicuous, adnate to petiole
for 1/2–3/4 of their length; style dilated, unappendaged, with ventral stigmatic orifice.
3. Leaves glabrous; lateral leaf lobes medially incurved-falcate, the distal portion of each overlapping the
base of the one above it, the apices attenuate into a prolonged hair-like tip; bractlets of the pedicels
small, distinct; known from a number of sites in central Peru (Dept. Junín) ………………V. weibelii
3. Leaves with margins sparsely to moderately setose (laminas essentially glabrous); lateral leaf lobes
straight, commonly oblong-lanceolate or slightly asymmetrically rhombic to narrowly elliptical,
occasionally broadly elliptical to suborbicular, predominately nonoverlapping, the apices obtuse to
narrowly (or broadly) rounded; bractlets of the pedicels very large, basally fused; known only from
type locality southeast of Cerro Palla Palla in southern Peru (Dept. Ayacucho)…………V. lilliputana
Acknowledgments
The authors thank Tom Wendt and an
anonymous reviewer for improving the quality
of the manuscript prior to publication, and
curatorial staff of the following herbaria for
facilitating our investigations through loan
administration or assistance during visits: B,
BM, BR, F, G/G-DC, GH, HAL, HBG, K, LZ,
M/BSM, MO, NY, P/P-HUMB/P-JUSS, PR, U,
US, W, and Z. The first author thanks Ohio
University undergraduates Jill Brown, David
May and Anya Porter for research assistance
that contributed to investigations on Central
Andean Andinium violets. He also gratefully
acknowledges financial support from Ohio
University’s Baker Fund, OU's Honors Tuto-
rial College, and Research Experiences for
Undergraduates supplements to National Sci-
ence Foundation grant DEB-9973958, "Test-
ing Systematic and Evolutionary Hypotheses
in a Primitive Neotropical Groups of Violets
(Vio la Sect. Leptidium)."
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