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Four new species of Cyrtandra (Gesneriaceae) from the South Pacific islands of Fiji

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  • DKI US Pacific Basin Agricultural Research Center USDA-ARS

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During fieldwork in Fiji, four new species of Cyrtandra (Gesneriaceae) were discovered and are described herein: C.gregoryi M.A.Johnson, sp. nov., C.hispida M.A.Johnson, sp. nov., C.longifructosa M.A.Johnson, sp. nov., and C.waisaliensis M.A.Johnson, sp. nov. The addition of four new species brings the current number of Fijian Cyrtandra to 41 endemic species. Two of the four species are known from only a single locality, and all of the new species are likely endangered or critically endangered. Continued fieldwork in the islands of Fiji is warranted in order to better understand current species distributions and population demographics of Cyrtandra in this species-rich and still poorly explored region of the South Pacific.
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Four new species of Cyrtandra (Gesneriaceae) from the South Pacic islands of Fiji 85
Four new species of Cyrtandra (Gesneriaceae)
from the South Pacific islands of Fiji
Melissa A. Johnson1
1 Rancho Santa Ana Botanic Garden, Claremont Graduate University, 1500 N. College Ave., Claremont, CA
91711, USA
Corresponding author: Melissa A. Johnson (mjohnson@rsabg.org; melissa.johnson@cgu.edu)
Academic editor: Eric Roalson|Received 13 October 2017|Accepted 1 December 2017|Published 15 December2017
Citation: Johnson MA (2017) Four new species of Cyrtandra (Gesneriaceae) from the South Pacic islands of Fiji.
PhytoKeys 91: 85–104. https://doi.org/10.3897/phytokeys.91.21623
Abstract
During eldwork in Fiji, four new species of Cyrtandra (Gesneriaceae) were discovered and are described
herein: C. gregoryi M.A.Johnson, sp. nov., C. hispida M.A.Johnson, sp. nov., C. longifructosa M.A.Johnson,
sp. nov., and C. waisaliensis M.A.Johnson, sp. nov. e addition of four new species brings the current num-
ber of Fijian Cyrtandra to 41 endemic species. Two of the four species are known from only a single locality,
and all of the new species are likely endangered or critically endangered. Continued eldwork in the islands
of Fiji is warranted in order to better understand current species distributions and population demographics
of Cyrtandra in this species-rich and still poorly explored region of the South Pacic.
Keywords
Cyrtandra, Gesneriaceae, Fiji, South Pacic, islands, new species, taxonomy, conservation
Introduction
e Southeast Asian-Pacic genus Cyrtandra J.R. Forster & G. Forster (Gesneriaceae)
comprises ca. 800 species, with centers of diversity in Borneo, New Guinea, the Phil-
ippines, and the Pacic islands (Atkins et al. 2013). Species of Cyrtandra are restricted
to the understory of rainforests and exhibit high diversity in habit (shrubs, small trees,
herbs, or vines), ower color (white, yellow, purple, pink, red), and fruit morphology
(indehiscent capsules or berries). In the Pacic, Cyrtandra is one of the largest and most
PhytoKeys 91: 85–104 (2017)
doi: 10.3897/phytokeys.91.21623
http://phytokeys.pensoft.net
Copyright Melissa A. Johnson. This is an open access article distributed under the terms of the Creative Commons Attribution License (CC
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Melissa A. Johnson / PhytoKeys 91: 85–104 (2017)
86
widely distributed genera of owering plants, with ca. 175 species occurring across a
region that extends from the Solomon Islands, east to the Marquesas, and north to
the Hawaiian Islands. e vast majority of species are single island endemics, with the
entire range of a species often being restricted to a single valley or mountain region.
Recent phylogenetic studies suggest that Cyrtandra evolved in Southeast Asia, fol-
lowed by dispersal to the Pacic islands (Clark et al. 2008, 2009; Johnson et al. 2017),
likely via frugivorous birds. e Pacic clade appears to have originated in Fiji ca. 9
mya, with subsequent founder events from Fiji to archipelagos both near (e.g., Samoa)
and far (e.g., the Hawaiian Islands) resulting in the current distribution of Cyrtandra
across the Pacic (Johnson et al. 2017). While Fiji hosts the second highest number
of endemic Cyrtandra species in the Pacic (second only to the Hawaiian Islands, with
60 spp.), only limited research has been conducted on the genus in this diverse region.
Gillett (1967) conducted the only thorough taxonomic review of Fijian Cyrtandra to
date, with his treatment dividing 35 species among six informal groups. However, upon
further study of Cyrtandra across the South Pacic, Gillett (1973) acknowledged that
these groupings were largely inadequate and that a more accurate treatment of Fijian
Cyrtandra would require considerably more eldwork. Smith’s treatment of Cyrtandra
in the Flora of Fiji (1991) was largely drawn from Gillett (1967), although two species
were restored from synonymy bringing the number of recognized species to 37.
Botanical explorations in Fiji from 1840–1953 resulted in the description of all 37
currently recognized species of Fijian Cyrtandra. Since the 1960s, relatively few collec-
tions of Fijian Cyrtandra have been made. In the 1970s, a number of new roads were
built in Fiji to accommodate the expanding agricultural industry, vastly increasing ac-
cessibility into remote regions (e.g., central Vanua Levu, eastern Taveuni; Lin 2012).
In 2014 and 2015, I was able to undertake extensive eldwork across the four largest
Fijian islands (Viti Levu, Vanua Levu, Taveuni, and Kadavu), focusing on regions that
have been poorly explored in the past. is work resulted in the discovery of four new
species that are here described and illustrated.
Methods
Diagnoses of the new species are based on morphological traits and DNA sequence
variation in a phylogenetic context. Morphological measurements were taken from live
plants in the eld, as many characters essential to Cyrtandra identication are lost upon
drying (particularly oral characters). Information was also taken from liquid xative-
preserved owers and fruit, as well as from digital photographs. To ensure accurate
identication, comparisons were made with all existing species descriptions (Gillett
1967, Smith 1991) as well as with herbarium specimens housed at BISH, GH, K, NY,
PTBG, RSA, SUVA, UC, and US. Samples of all four new species were included in
a recent molecular phylogeny of the Pacic clade of Cyrtandra, which is based on ve
loci and a dense taxon sampling of 121 species (including 30 Fijian species; Johnson
et al. 2017). is study provided the information necessary to identify the closest
Four new species of Cyrtandra (Gesneriaceae) from the South Pacic islands of Fiji 87
relatives of each of the new species based on shared phylogenetic history. Conserva-
tion status was assessed in accordance with IUCN Red List Category criteria (IUCN
Standards and Petitions Subcommittee 2016).
Taxonomic treatment
Cyrtandra gregoryi M.A.Johnson, sp. nov.
urn:lsid:ipni.org:names:60475584-2
Figs 1, 2
Diagnosis. Cyrtandra gregoryi is closely related to C. ciliata Seem. (Fig. 2), but diers
in its elliptic to ovate leaves up to 39 × 17 cm (vs. lanceolate to ovate leaves up to 28
× 12 cm), axillary inorescences with indument of brown trichomes (vs. cauliorous
inorescences with indument of white trichomes), pale green calyces 6–13 mm long
that are cleft unequally into lanceolate coriaceous lobes (vs. calyces white, 7–10 mm
long, cleft nearly to the base into equal linear-lanceolate lobes), these splitting along
two to three sutures and recurving after anthesis (vs. remaining erect after anthesis),
and corollas with exserted style and stamens (vs. style and stamens included).
Type. FIJI. Taveuni: near the end of the Lavena coastal walk along the Wainibau
stream, ca. 3.5 km NW of Lavena Village, 16°52.10'S, 179°54.32'W, 32 m elev., 04
August 2014, M.A. Johnson 105 with G.J. Hora (holotype: RSA).
Description. Shrub 0.9–2.2 m tall; stems unbranched to few-branched, with a dense
indument of dark brown uniseriate multicellular trichomes ca. 0.5 mm long. Leaves op-
posite, internodes 2–7 cm long, blades elliptic to ovate to obovate, 22–39 cm long, 7–17
cm wide, upper surface sparsely strigillose, lower surface glabrate except for the densely
pubescent 9–13 secondary veins on each side, margins serrulate to subentire, apex acute,
base cuneate to rounded, petioles 5–11 cm long, densely pubescent with short brown
trichomes; inorescence an axillary cyme with dense brown pubescence on the peduncles
and pedicels, 3–22 owers, cymules 1–4 owered, peduncle to 31 mm long, terminated
by bracts to 6 mm long, narrowly lanceolate, deciduous after anthesis, pedicels to 31
mm long; calyx 6–13 mm long, outer surface pale green and moderately pubescent with
appressed dark brown uniseriate trichomes, inner surface white and papillate, unequally
cleft into 5 lanceolate coriaceous lobes, 3–10 mm long, apex acuminate, upper lobes oc-
casionally connate, often continuing to split along two or three sutures and curl back after
anthesis, deciduous; corolla white, tube funnelform, slightly curved near mid point, 23–27
mm long, 6–9 mm wide, outer surface glabrate to densely pubescent with short glandular
trichomes, inner surface glabrous, upper lobes 5–7 mm long and 5–6 mm wide, lower
lobe 7–8 mm long and 5–6 mm wide, lobes recurving after anthesis; stamens 2, ca. 9 mm
long, becoming exserted from the corolla tube during ovulate phase, base of the laments
bright orange, anthers apically connate, staminodes 3; nectary disc cupulate, annular; gy-
noecium (ovary, style, and stigma) 16–24 mm long, ovary glabrous, style pubescent with
capitate glandular trichomes, exserted, stigma shallowly bilobed; fruit not seen.
Melissa A. Johnson / PhytoKeys 91: 85–104 (2017)
88
Figure 1. Cyrtandra gregoryi M.A.Johnson A Habit B Inorescence C Corolla, anterior view D Calyx
EFlower, lateral view F Gynoecium. Drawn from Johnson 105 (RSA) and eld images.
Distribution and ecology. Cyrtandra gregoryi is only known from two locations
in eastern Taveuni, Fiji, where it grows in lowland forests and along stream banks from
ca. 30–50 m (Fig. 3).
Phenology. Individuals of this species were in ower when collected in August and
November, with fruits likely becoming mature ca. 5–6 months later.
Etymology. I am pleased to name this new species after my husband, Gregory
Hora, to whom I am most grateful for his assistance in collecting this and other species
across Fiji.
Four new species of Cyrtandra (Gesneriaceae) from the South Pacic islands of Fiji 89
Figure 2. Cyrtandra gregoryi M.A.Johnson and closest relative (C. ciliata) based on a molecular phylog-
eny by Johnson et al. (2017). A C. gregoryi shrub habit B C. gregoryi corolla, anterior view C C. gregoryi
ower, lateral view D C. gregoryi axillary cyme inorescence E C. gregoryi adaxial leaf surface F C. ciliata
corolla, anterior view G C. ciliata ower, lateral view H C. ciliata cauliorous cyme inorescence. All
photos taken in the eld by M. Johnson.
Phylogenetic placement. A recent phylogenetic study by Johnson et al. (2017)
placed Cyrtandra gregoryi sister to C. ciliata with strong support (Fig. 4). Cyrtandra cilia-
ta is endemic to the Fijian islands of Vanua Levu, Taveuni, and Koro from 300–1100m
Melissa A. Johnson / PhytoKeys 91: 85–104 (2017)
90
Figure 3. Distribution of Cyrtandra gregoryi and C. hispida on Taveuni, Fiji.
elevation. ese species share a cymose inorescence and glabrous to glabrate leaves. e
key provided in the taxonomic treatment by Gillett (1967) would place C. gregoryi in
species Group 5 based on the branching cyme inorescence, non-woody inorescence
axis, deciduous calyx, styles and/or stamens being exserted from the corolla tube, and
stamens 8–12 mm long.
Conservation status. Proposed IUCN Red List Category: Endangered (EN)
based on an estimated area of occupancy of < 500 km2 (criterion B2), known to exist
at no more than ve locations (B2a), inferred decline in area of occupancy (B2bii),
decline in area, extent, and/or quality of habitat (B2biii), decline in number of mature
individuals (B2bv), and population size estimated to number fewer than 250 mature
individuals (D).
Although Bouma National Heritage Park protects ca. 15,000 hectares of intact rain-
forest on eastern Taveuni, indigenous Fijians are permitted to clear land near villages
for agriculture. As a result, large areas of coastal forest are increasingly being cleared for
dalo (taro, Colocasia esculenta (L.) Schott) and yaqona (kava, Piper methysticum L.f.),
the two main export crops of Taveuni. Given that C. gregoryi appears to be restricted to
low-elevation forests, it is highly likely that individuals of this species were extirpated
during clearing for human settlements and agriculture. Invasive plants are also a major
threat to native plants in the area; mile-a-minute vine (kudzu, Pueraria lobata (Willd.)
Ohwi) may be particularly problematic as it rapidly grows over trees and shrubs and can
Four new species of Cyrtandra (Gesneriaceae) from the South Pacic islands of Fiji 91
Figure 4. A portion of the Maximum Likelihood phylogram from Johnson et al. (2017) based on three
nuclear (ITS, ETS, Cyrt1) and two chloroplast (psbA-trnH, rpl32-trnL) loci. Support values shown for
each branch are bootstrap and posterior probabilities when ≥ 50% and ≥ 0.50, respectively. An asterisk
indicates 100% BS or 1.0 PP; a dash indicates that the branch was not supported.
kill other plants with heavy shading. Lastly, anthropogenic-induced climate change is a
threat to island plant communities. Tropical cyclones are expected to increase in intensity
and severity in the coming years (Emmanuel 2005, Knutson 2010), and can have dev-
astating eects on island vegetation due to high winds, ooding, and storm surges (e.g.,
Cyclone Pam in 2015 caused extensive damage to Vanuatu’s forests). Most recently, Cy-
clone Winston, the strongest tropical cyclone to make landfall in the South Pacic Basin
in recorded history, ravaged the islands of Fiji in February of 2016. e coastal regions
of eastern Taveuni were inundated by massive storm surges, and much of the vegetation
was damaged by winds of up to 185 mph. Although forested regions can often regenerate
after a natural disaster if given sucient time, recent research in the South Pacic sug-
gests that a cyclone can be a catalyst for human-coping strategies that increase pressure
on forest ecosystems and exposes them to invasive plant species (Goulding et al. 2016).
Melissa A. Johnson / PhytoKeys 91: 85–104 (2017)
92
Additional specimens examined. FIJI. Taveuni: Tavoro Falls Trail in Bouma Na-
tional Heritage Park, 19 November 2016, J.C. Game 16/235 with S. Fawcett (PTBG).
Notes. Eight individuals of C. gregoryi were recorded during eld surveys along the
Lavena coastal trail, with all of these being reproductive. Additional eld surveys in the
area are likely to reveal more individuals. No other Cyrtandra species were observed
growing sympatrically with C. gregoryi in the Lavena region, although C. tempestii
Horne ex. C.B. Clarke was collected 0.64 km to the SE. An additional collection was
made of a single individual of C. gregoryi near the Tavoro Falls in Bouma National
Heritage Park, an area that also hosts C. ciliata.
Cyrtandra hispida M.A.Johnson, sp. nov.
urn:lsid:ipni.org:names:77174061-1
Figs 5, 6
Diagnosis. Cyrtandra hispida is morphologically similar to C. waisaliensis sp. nov., but
diers in its axillary cyme inorescence of 2–4 owers (vs. cauliorous cyme inores-
cence of 2–8 owers), green bracts and bracteoles 5–9 mm long (vs. white bracts and
bracteoles 3–10 mm long), calyx pale green and 29–31 mm long (vs. calyx white and
23–37 mm long), corolla tube 31–34 mm long (vs. corolla tube 23–27 mm long), and
staminodes 2 (vs. staminodes 3).
Type. FIJI. Taveuni: Des Voeux Peak, 16°50.48'S, 179°57.97'W, 1109 m elev., 13
July 2014, M.A. Johnson 91 with G.J. Hora (holotype: RSA; isotype: SUVA).
Description. Shrub 0.6–1.2 m tall; stems unbranched or few-branched, with light
brown hispid uniseriate multicellular trichomes ca. 5 mm long. Leaves opposite, in-
ternodes 3–9 cm long, the blades oval to ovate to obovate, 15–22 cm long, 8–10 cm
wide, upper and lower surfaces densely strigose with light brown uniseriate trichomes to
2mm long, 5–7 secondary veins on each side, margins serrate to biserrate, apex acute to
acuminate, base oblique or aequilateral and rounded to cuneate, petioles 4–9 cm long,
densely pubescent with hispid trichomes ca. 6 mm long; inorescence an axillary cyme,
2–4 owered, densely hispid throughout, peduncle 10–14 mm long, terminated by
green bracts 5–9 mm long, ovate to narrowly lanceolate, densely hispid, pedicels 27–28
mm long, often subtended by bracteoles similar to the outer bracts; calyx pale green,
cylindrical, 29–31 mm long, unequally cleft into ve triangular lobes 4–11 mm long,
outer and inner surfaces densely hispid with uniseriate trichomes, persistent on develop-
ing fruits; corolla white, bilabiate, tube narrowly funnelform, slightly curved near mid
point, outer and inner surface glabrous, 31–34 mm long and 8–9 mm wide, upper
lobes 12–15 mm long and 10–14 mm wide, lower lobe 13–20 mm long and 10–15
mm wide; stamens 2, ca. 7 mm long, anthers apically connate, staminodes 2; nectary
disc cupulate, annular, deciduous from the fruit; gynoecium (ovary, style, and stigma) ca.
22mm long, ovary and style glabrous, stigma applanate, bilobed; berries green when
immature, ellipsoid, glabrous, up to 18 mm long and 11 mm wide, tipped by the basal
3 mm of the persistent style, enclosed by the persistent calyx, mature fruit not seen.
Four new species of Cyrtandra (Gesneriaceae) from the South Pacic islands of Fiji 93
Figure 5. Cyrtandra hispida M.A.Johnson. A Habit B Inorescence C Corolla, anterior view D Co-
rolla, longitudinal section E Calyx, longitudinal section and young fruit. Drawn from Johnson 91 (RSA,
SUVA), Johnson 212 (SUVA), Johnson 215 (RSA), and eld images.
Distribution and ecology. Cyrtandra hispida is only known from two populations in
the upland rainforests of Taveuni, Fiji, where plants occur on exposed hillsides composed
of volcanic cinders, and on rocky stream banks from 697–1126 m elevation (Fig. 3).
Melissa A. Johnson / PhytoKeys 91: 85–104 (2017)
94
Figure 6. Cyrtandra hispida M.A.Johnson and closest relative (C. cephalophora) based on molecular phy-
logeny by Johnson et al. (2017). A C. hispida corolla, anterior view B C. hispida ower, lateral view C
C. hispida shrub habit and axillary cyme inorescence D C. cephalophora shrub habit E C. cephalophora
capitate-cylindrical cauliorous inorescence and young fruits F C. cephalophora corolla, anterior view.
All photos taken in the eld by M. Johnson, with photos of C. hispida taken from the type collection.
Phenology. Individuals of this species were in ower when collected in July, with
fruits likely becoming mature ca. 5–6 months later (December–January).
Etymology. is species is named for the sti trichomes that cover the stems,
leaves, and inorescences.
Phylogenetic placement. A recent phylogenetic study by Johnson et al. (2017)
placed Cyrtandra hispida in a weakly supported clade with four other species (C. cepha-
lophora Gillespie, C. waisaliensis sp. nov., C. dolichocarpa A. Gray, C. longifructosa sp.
nov.) that are recorded from the Fijian Islands of Viti Levu (C. cephalophora) and/or
Vanua Levu (C. cephalophora, C. waisaliensis, C. dolichocarpa, C. longifructosa) (Fig.
4). Within this clade, C. hispida is most similar morphologically to C. waisaliensis (sp.
nov., described below). Both species have large bilabiate corollas, persistent cylindrical
calyces, ovate to obovate leaves, and a dense indument of long sti trichomes covering
the stems, leaves, and inorescences. Additional sampling of species and of nuclear
genic regions may be required to condently place C. hispida with its closest relatives.
e key provided in the taxonomic treatment by Gillett (1967) would place C. hispida
in species Group 2 based on the branching cyme inorescence and the persistent calyx.
Conservation status. Proposed IUCN Red List Category: Endangered (EN) based
on an estimated area of occupancy of < 500 km2 (criterion B2), known to exist at no
more than ve locations (B2a), projected decline in extent of occurrence (B2bi), area
of occupancy (B2bii), and area, extent, and/or quality of habitat (B2biii). Although
the two areas where this species has been collected are within the Taveuni Forest Re-
serve, the forest above Somosomo Village is currently being cleared for a hydropower
Four new species of Cyrtandra (Gesneriaceae) from the South Pacic islands of Fiji 95
dam (M. Johnson, pers. obs.). Additional threats include mining for gold and copper,
invasion by plant species such as Clidemia hirta (L.) D. Don (Koster’s curse; M. John-
son, pers. obs), and damage from tropical cyclones. Further surveys are needed in the
upland forests of Taveuni (which remain relatively unexplored, exceptions being the
area surrounding Lake Tagimoucia and the road to Des Voeux Peak) to determine the
extent of occurrence and population demographics of C. hispida.
Additional specimens examined. FIJI. Taveuni: mountains above Somosomo,
16°47.67'S, 179°56.10'W, 693 m elev., 24 August 2015, M.A. Johnson 212 (SUVA),
M.A. Johnson 215 (RSA).
Notes. Cyrtandra hispida was observed to grow sympatrically with three species on
Des Voeux Peak (C. leucantha A.C. Sm., C. ciliata, and Cyrtandra sp.) and three species
in the mountains above Somosomo (C. leucantha, C. ciliata, C. taviunensis Gillespie).
Several individuals were observed that appeared to be of hybrid origin in these popula-
tions, with the widespread and common C. ciliata inferred as one of the parents based
on similar oral morphology. While the observation of ongoing hybridization in these
populations suggests the possibility of C. hispida being of hybrid origin, none of the
sympatric species have morphological characters similar to C. hispida. Furthermore,
C. hispida is placed in a clade of species that are endemic to the neighboring islands of
Vanua Levu and Viti Levu, and does not appear to be closely related to species endemic
to Taveuni.
Cyrtandra longifructosa M.A.Johnson, sp. nov.
urn:lsid:ipni.org:names:77174062-1
Figs 7, 8
Diagnosis. is species is closely related to C. dolichocarpa (Fig. 8), but diers in its
glabrous elliptic-ovate leaves (vs. moderately pubescent lanceolate-ovate leaves), blades
up to 22 × 9 cm (vs. blades up to 17 × 7 cm), petioles 3–9 cm long (vs. petioles 1–4
cm long), deciduous lanceolate bracts (vs. persistent ovate bracts), peduncles 2–4 mm
long (vs. 5–10 mm long), pedicels 11–18 mm long (vs. 21–27 mm long), deciduous
beaked calyx (vs. persistent cylindrical calyx), and corolla tube 23–29 mm long (vs.
corolla tube 36–55 mm long).
Type. FIJI. Vanua Levu: ca. 0.8 km NE of Waisali Village along the Waisali Creek,
16°38.51'S, 179°14.54'E, 110 m elev., 09 July 2014, M.A. Johnson 65 with G.J. Hora
(holotype: SUVA; isotype: RSA).
Description. Shrub 0.9–1.7 m tall; stems unbranched to few branched. Leaves op-
posite, internodes 1–5 cm long, the blades narrowly elliptic to elliptic-ovate, 17–22 cm
long, 7–9 cm wide, upper and lower surface glabrous, 8–10 secondary veins on each
side, these slightly impressed, margins serrulate, apex acuminate, base oblique to aequi-
lateral and attenuate to cuneate, petioles 3–9 cm long, glabrous to glabrate; inorescence
an axillary cyme, 1– 2(3–4) owers, densely pilose with black trichomes ca. 1 mm
long throughout, peduncle 2–4 mm long, terminated by green bracts, 3–8 mm long,
Melissa A. Johnson / PhytoKeys 91: 85–104 (2017)
96
Figure 7. Cyrtandra longifructosa M.A.Johnson. A Habit B Corolla, staminate phase, anterior view
CMature elongate cylindrical fruit D Inorescence and young fruit E Corolla, ovulate phase, anterior
view. Drawn from Johnson 65 (SUVA, RSA), Johnson 59 (RSA), and eld images.
lanceolate, deciduous after anthesis, pedicels 11–18 mm long; calyx pale green, outer
and inner surfaces pubescent with appressed uniseriate trichomes, narrowly fusiform in
bud, 28–32 mm long, beaked, the 5 lobes often remaining connivent, splitting along
one suture 14–19 mm long, deciduous; corolla white, becoming strongly bilabiate in the
Four new species of Cyrtandra (Gesneriaceae) from the South Pacic islands of Fiji 97
Figure 8. Cyrtandra longifructosa M.A.Johnson and closest relative (C. dolichocarpa) based on molecular
phylogeny by Johnson et al. (2017). A C. longifructosa corolla, anterior view B C. longifructosa ower, lat-
eral view C C. longifructosa elongate cylindrical fruits D C. longifructosa axillary inorescence and young
fruits E C. longifructosa shrub habit F C. dolichocarpa corolla, anterior view G C. dolichocarpa ower,
lateral view H C. dolichocarpa shrub habit. All photos taken in the eld by M. Johnson, with photos of C.
longifructosa taken from the type collection.
ovulate phase, tube cylindrical, curved near the mid point, outer surface glabrous, inner
surface with uniseriate trichomes throughout and short glandular trichomes near the
mouth of the tube, the tube 23–29 mm long, 5–7 mm wide, upper lobes 8–12 mm long
and 8–11 mm wide, lower lobe 12–14 mm long and 11–14 mm wide; stamens 2, ca. 6
mm long, base of the laments reddish orange, anthers apically connate, staminodes3;
Melissa A. Johnson / PhytoKeys 91: 85–104 (2017)
98
nectary disc cupulate, annular, deciduous from mature fruit; gynoecium (ovary, style, and
stigma) ca. 24 mm long, ovary glabrous, style pubescent with capitate glandular tri-
chomes along distal ⅓ of its length, stigma bilobed; berries cylindrical, elongate, mature
fruit to 40 mm long and 5 mm wide, glabrous, turning white at maturity.
Distribution and ecology. Cyrtandra longifructosa is known only from one popu-
lation in the Waisali region of central Vanua Levu, Fiji, where it occurs in the wet forest
understory along a small creek at ca. 110 m elevation (Fig. 9).
Phenology. Individuals of this species had owers, immature fruits, and mature
fruits when collected in July.
Etymology. Named for the elongate cylindrical fruits, one of the diagnostic char-
acteristics of this species.
Phylogenetic placement. e phylogenetic study by Johnson et al. (2017) placed
Cyrtandra longifructosa as sister to C. dolichocarpa (endemic to Vanua Levu and Rabi,
Fiji) with strong support (Fig. 4). ese species both have large bilabiate corollas and
elongate cylindrical berries. e key provided in the taxonomic treatment by Gillett
(1967) would place C. longifructosa in species Group 3 based on the branching cyme
inorescence, non-woody inorescence axis, deciduous calyx, inserted anthers and
styles, and calyx lobes about the same length as the calyx tube.
Conservation status. Proposed IUCN Red List Category: Critically Endangered
(CR): based on an estimated area of occupancy of < 10 km2 (criterion B2), known to
Figure 9. Distribution of Cyrtandra waisaliensis and C. longifructosa on Vanua Levu, Fiji.
Four new species of Cyrtandra (Gesneriaceae) from the South Pacic islands of Fiji 99
exist only at a single location (B2a), projected decline in extent of occurrence (B2bi),
area of occupancy (B2bii), and area, extent, and/or quality of habitat (B2biii). is
species is only known from one locality in the central mountains of Vanua Levu, war-
ranting additional surveys in areas of Vanua Levu with intact rainforest (e.g., Waisali,
the Natewa Peninsula) to determine the full extent of occurrence and population de-
mographics of C. longifructosa. Regions with suitable rainforest habitat on Vanua Levu
are threatened by logging, mining for bauxite and gold, invasive plant species such as
Clidemia hirta (Koster’s curse; M. Johnson, pers. obs.), and tropical cyclones.
Additional specimens examined. FIJI. Vanua Levu: NE of Waisali Village along
Waisali Creek, 16°38.51'S, 179°14.54'E, 110 m elev., 09 July 2014, M.A. Johnson 59
with G.J. Hora (RSA).
Notes. e population of C. longifructosa was observed to contain ca. 20 individu-
als, many of which were reproductive. No other Cyrtandra species were observed grow-
ing in the immediate vicinity, although the closely related species C. dolichocarpa, C.
waisaliensis, and C. cephalophora were all collected 2.25 km W of the C. longifructosa
population described here.
Cyrtandra waisaliensis M.A.Johnson, sp. nov.
urn:lsid:ipni.org:names:77174063-1
Figs 10, 11
Diagnosis. e new species is closely related to C. dolichocarpa and C. longifructosa
(Fig. 8), but diers in its dense bristly pubescence on the young stems, leaves, petioles,
and inorescences (vs. moderate appressed pubescence on C. dolichocarpa; vs. glabrous
on C. longifructosa), cauliorous inorescences (vs. axillary inorescences in both C.
dolichocarpa and C. longifructosa), persistent foliaceous ovate white bracts to 10 mm
long (vs. non-foliaceous green bracts to 5 mm long in C. dolichocarpa; vs. deciduous
non-foliaceous lanceolate green bracts to 8 mm long in C. longifructosa), and multiple
persistent foliaceous white bracteoles (vs. single deciduous non-foliaceous green brac-
teoles in C. dolichocarpa; vs. bracteoles absent in C. longifructosa).
Type. FIJI. Vanua Levu: Waisali Forest Reserve along the lower portion of the Sa-
vuqoro Creek, 16°38.39'S, 179°13.25'E, 338 m elev., 07 July 2014, M.A. Johnson 50
with G.J. Hora (holotype: SUVA; isotype: RSA).
Description. Shrub 0.7–2.6 m tall; stems few- to many-branched, young stems cov-
ered in hirsute trichomes. Leaves opposite, those at a node unequal, internodes 2–8 cm
long, blades ovate to obovate, 13–24 cm long, 6–12 cm wide, upper and lower surfaces
densely hirsute with uniseriate multicellular trichomes, 6–7 secondary veins on each side,
margins serrate, apex acuminate, base oblique to aequilateral and attenuate to cuneate,
petioles 3–11 cm long, densely hirsute, trichomes ca. 5 mm long; inorescence of cauli-
orous cymes, 2–8 owered, cymules 1–4 owered, densely hirsute throughout, pedun-
cle 3–8 mm long, terminated by persistent foliaceous white bracts, ovate to lanceolate,
3–10 mm long, outer surface glabrous to glabrate, inner surface densely hirsute, pedicels
Melissa A. Johnson / PhytoKeys 91: 85–104 (2017)
100
Figure 10. Cyrtandra waisaliensis M.A.Johnson. A Habit B Cauliorous inorescence C Corolla, an-
terior view D Corolla, longitudinal section E Gynoecium F Calyx, longitudinal section and young fruit.
Drawn from Johnson 48 (RSA), Johnson 50 (SUVA, RSA), and eld images.
21–40 mm long, subtended by multiple persistent bracteoles similar to and closely sub-
tended by the outer bracts; calyx white, cylindrical, densely hirsute, 23–37 mm long,
unequally cleft into 5 triangular lobes, 4–14 mm long, persistent; corolla white, bilabiate,
tube funnelform, curved near mid point, outer surface glabrous to glabrate, inner surface
with capitate glandular trichomes near the mouth of the tube, 23–27 mm long, 6–9 mm
wide, upper lobes 11–12 mm long and 8–11 mm wide, lower lobe 13–17 mm long and
10–15 mm wide; stamens 2, 3–6 mm long, base of the laments reddish orange, anthers
Four new species of Cyrtandra (Gesneriaceae) from the South Pacic islands of Fiji 101
Figure 11. Cyrtandra waisaliensis M.A.Johnson. A Corolla, anterior view B Flower, lateral view CCau-
liorous cyme inorescence D Shrub habit E Rainforest understory habitat. All photos from the type
collection, taken in the eld by M. Johnson.
apically connate, staminodes 3; nectary disc prominent, annular, 2 mm high; gynoecium
(ovary, style, and stigma) 16–26 mm long, ovary glabrous, style pubescent with capi-
tate glandular trichomes along distal ⅓ of length, stigma bilobed; berries ellipsoid, green
when immature, enclosed by the persistent calyx, mature fruit not seen.
Distribution and ecology. Cyrtandra waisaliensis is known only from one popula-
tion in the Waisali Forest Reserve on Vanua Levu, Fiji at 300–360 m elevation, occur-
ring in the dense forest understory alongside a creek (Fig. 9).
Phenology. Flowers and immature fruits were observed in July, with fruits likely
becoming mature ca. 5–6 months later (December–January).
Etymology. e new species is named after the area of Vanua Levu where it was
collected, Waisali Forest Reserve.
Phylogenetic placement. A recent phylogenetic study by Johnson et al. (2017)
supported the placement of Cyrtandra waisaliensis as sister to C. longifructosa (endemic
to Vanua Levu) and C. dolichocarpa (endemic to Vanua Levu and Rabi; Fig. 4). ese
species all have large bilabiate corollas, and both C. dolichocarpa and C. longifructosa
have elongate cylindrical white fruits. However, C. waisaliensis is also morphologically
similar to C. hispida; these species share bilabiate corollas, persistent cylindrical caly-
Melissa A. Johnson / PhytoKeys 91: 85–104 (2017)
102
ces, and a dense indument of sti uniseriate trichomes. Cyrtandra hispida is currently
placed in a polytomy with C. cephalophora and the clade comprising C. waisaliensis,
C. longifructosa, and C. dolichocarpa. e key provided in the taxonomic treatment by
Gillett (1967) would place C. waisaliensis in species Group 2, based on the branching
cyme inorescence and the persistent calyx.
Conservation status. Proposed IUCN Red List Category: Critically Endangered
(CR) based on an estimated area of occupancy of < 10 km2 (criterion B2), known to
exist at only a single location (B2a), projected decline in extent of occurrence (B2bi),
area of occupancy (B2bii), and area, extent, and/or quality of habitat (B2biii). is
species is only known from one locality in the central mountains of Vanua Levu, war-
ranting additional surveys in areas of Vanua Levu with intact rainforest (e.g., Waisali,
the Natewa Peninsula) to determine the full extent of occurrence and population de-
mographics of C. waisaliensis. Regions with suitable rainforest habitat on Vanua Levu
are threatened by logging, mining for bauxite and gold, invasive plant species such as
Clidemia hirta (Koster’s curse; M. Johnson, pers. obs.), and tropical cyclones.
Additional specimens examined. FIJI. Vanua Levu: Waisali Forest Reserve along
the lower portion of the Savuqoro Creek, 16°38.34'S, 179°13.18'E, 355 m elev., 07
July 2014, M.A. Johnson 48 with G.J. Hora (RSA).
Notes. e observed population of C. waisaliensis was comprised of ca. 20 indi-
viduals, many of which were reproductive. A single individual appeared to be of hybrid
origin, with the putative parents being C. waisaliensis and C. cephalophora based on
morphological characters intermediate between these two species.
Conclusions
e recognition of C. gregoryi, C. hispida, C. longifructosa, and C. waisaliensis brings
the new total of Fijian Cyrtandra species to 41. e four new species of Cyrtandra
described here demonstrate that the islands of Fiji remain poorly explored botani-
cally, at least in some regions. Of the 28 Fijian Cyrtandra species collected during eld
expeditions in 2014 and 2015, four of these were new to science. An additional four
Cyrtandra species could not be keyed out to any of the existing species due to a lack of
reproductive material. However, with further eld study it is possible that these, along
with other future collections, may be identied as new species.
Acknowledgements
I am grateful to Marika Tuiwawa, Alivereti Naikatini, Manoa Maiwaqa, Mereia Tabua,
and Sarah Pene (South Pacic Regional Herbarium, e University of the South Pacic)
for assistance with eldwork logistics in Fiji. Collecting permits were granted by e Fiji
Ministry of Education, export permits by Fiji Biosecurity, and import permits by the
United States Department of Agriculture. I also thank my husband Gregory Hora for
Four new species of Cyrtandra (Gesneriaceae) from the South Pacic islands of Fiji 103
being an excellent eld assistant and for providing endless encouragement and moral
support. is work would not have been possible without the kind peoples of Fiji, who
oered their assistance, expertise and hospitality. e following herbaria graciously al-
lowed the use of their collections for study: BISH, GH, K, NY, RSA, SUVA, UC, US,
and WU. John Game provided collection information and photographs of Cyrtandra
gregoryi for study. Funding for this project was provided by the following: Rancho Santa
Ana Botanic Garden; Garden Club of America Award in Tropical Botany; American
Philosophical Society Lewis and Clark Fund for Exploration and Field Research; e
Gesneriad Society Nelly D. Sleeth Scholarship Endowment Fund; e Gesneriad Soci-
ety Elvin McDonald Fund; Society of Systematic Biologists Graduate Student Research
Award; Sigma Xi Grants-in-Aid of Research; American Society of Plant Taxonomists
Graduate Student Research Grant; and Claremont Graduate University. Lastly, I am
grateful to my sister, Liv Johnson, for the beautiful line drawings, and to Lucinda Mc-
Dade, John R. Clark, J. Travis Columbus, J. Mark Porter, David Lorence, and one
anonymous reviewer for comments that greatly improved this manuscript.
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... In recent years, several studies (Wagner et al. 2001;Bramley et al. 2003;Atkins 2004;Bramley 2005;Lorence & Perlman 2007;Bone & Atkins 2013;Johnson 2017;Kartonegoro et al. 2018;Atkins et al. 2019;Nishii et al. 2019) have led to an increase in numbers of species of Cyrtandra. These studies emphasized the urgency to document and understand the biodiversity of cyrtandras before they succumb to anthropogenic pressures. ...
... Herbaria acronyms follow Index Herbariorum (Thiers, continuously updated). Descriptions follow schemes of recently published accounts of new species Johnson 2017;Kartonegoro et al. 2018;Atkins et al. 2019). Assessment of Conservation Status was implemented using GeoCAT (Bachman et al. 2011), following the IUCN Red List Category criteria (IUCN Standards and Petitions Subcommittee 2017). ...
... Atkins et al. (2013) estimated 800 species of Cyrtandra. Since then, several authors (Bone & Atkins 2013;Johnson 2017;Kartonegoro et al. 2018;Atkins et al. 2019) have described additional species. The number of species is expected to increase further as more alpha-taxonomic work and field collection are carried out. ...
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