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Phytotaxa 533 (3): 173–180
https://www.mapress.com/pt/
Copyright © 2022 Magnolia Press Article PHYTOTAXA
ISSN 1179-3155 (print edition)
ISSN 1179-3163 (online edition)
Accepted by Yi-Hua Tong: 20 Jan. 2022; published: 11 Feb. 2022
https://doi.org/10.11646/phytotaxa.533.3.3
173
Licensed under Creative Commons Attribution-N.C. 4.0 International https://creativecommons.org/licenses/by-nc/4.0/
Vaccinium carmesinum (Ericaceae), a new species of blueberry from Mt. Tago
Range, Mindanao Island, Philippines
MAVERICK N. TAMAYO1,11*, FULGENT P. CORITICO2,3,12, VICTOR B. AMOROSO2,3,13, DARIN S. PENNEYS4,14,
DANILO N. TANDANG5,6,7,8,9,15 & PETER W. FRITSCH10,16
1 Department of Biology, College of Science and Engineering, Texas Christian University, 2800 South University Drive, Fort Worth,
Texas, 76129 USA
2 Center for Biodiversity Research and Extension in Mindanao (CEBREM), Central Mindanao University, Musuan, Bukidnon 8710
Philippines
3 Department of Biology, College of Arts and Sciences, Central Mindanao University, Musuan, Bukidnon 8710 Philippines
4 Department of Biology and Marine Biology, University of North Carolina Wilmington, 601 South College Road, Wilmington, North
Carolina 28403 USA
5 Herbarium (HAST), Biodiversity Research Center, Academia Sinica, Taipei, 115 Taiwan
6 Philippine National Herbarium, Botany and National Herbarium Division, National Museum of Natural History, National Museum of
the Philippines, T.M. Kalaw St., Ermita, Manila, 1000 Philippines
7 Biodiversity Program, Taiwan International Graduate Program, Academia Sinica and National Taiwan Normal University, Taipei,
11529 Taiwan
8 Department of Life Science, National Taiwan Normal University, Taipei, 11677 Taiwan
9 Biodiversity Research Center, Academia Sinica, Taipei, 11529 Taiwan
10 Botanical Research Institute of Texas, 1700 University Drive, Fort Worth, Texas, 76107 USA
11
�
m.n.tamayo@tcu.edu; https://orcid.org/0000-0003-0157-5116
12
�
cfulgent@cmu.edu.ph; https://orcid.org/0000-0003-3876-6610
13
�
victorbamoroso@gmail.com; https://orcid.org/0000-0001-8865-5551
14
�
penneysd@uncw.edu; https://orcid.org/0000-0003-0727-2829
15
�
sue93653@yahoo.com; https://orcid.org/0000-0003-2708-661X
16
�
pfritsch@brit.org; https://orcid.org/0000-0002-3606-663X
* Author for correspondence
Abstract
Vaccinium carmesinum is described as a new species of Ericaceae from Mt. Tago Range, Mindanao Island, Philippines. It is
similar to V. platyphyllum Merrill and V. luzoniense S.Vidal but is distinct from the former by longer and wider leaves, longer
racemes, longer bracts, glabrous corollas, and glabrescent fruits, and from the latter by longer petioles, leaf glands distributed
along the blade margin, glabrous rachis, and lanate filaments. Vaccinium carmesinum bears the widest leaves among Philip-
pine Vaccinium. Its discovery increases the number of Vaccinium species recognized in the Philippines to 37.
Key words: Mindanao, Pantaron Mountain Range, Vaccinieae, volcanic-igneous
Introduction
Vaccinium Linnaeus (1753: 349) (Vaccinieae: Vaccinioideae) is one of the most species-rich genera in the family
Ericaceae, estimated to comprise ca. 450–500 species distributed around the world except in Antarctica and Australia
(Sleumer 1966–1967; Argent 2014). The floristic region of Malesia is considered to be the center of diversity of the
genus (Sleumer 1966–1967) with species generally inhabiting montane forests (Argent 2018), particularly mossy
forest, on exposed mountain ridges or summits. Although the genus is diverse, its taxonomic limits are still uncertain
(Argent 2008; Fritsch et al. 2020). Evidence of its monophyly has not been resolved (Vander Kloet & Dickinson 2009)
with the current phylogenetic data based on DNA sequences strongly suggesting nonmonophyly (Kron et al. 2002).
The island of Mindanao is the largest southern island in the Philippines and is known to harbor great biodiversity and
species endemicity (Amoroso et al. 2019). Within this island lies Mt. Tago Range—a component of the central cordillera
of Mindanao Island separating the provinces of Bukidnon in the west and Agusan del Sur in the east (Gronemeyer et al.
TAMAYO ET AL.
174 • Phytotaxa 533 (3) © 2022 Magnolia Press
2014). This mountain range is part of the ancestral land of the Higaonon tribe and is considered a terrestrial Philippine
Key Biodiversity Area (KBA) that supports the Tagoloan River watershed spanning the provinces of Bukidnon and
Misamis Oriental (BWPDC 2012). Mt. Tago Range harbors one of the remaining intact forests in Northern Mindanao.
However, it is currently not legally protected (Coritico et al. 2020a) and is relatively unexplored because of political
instability (Lagunday et al. 2017; Lagunday & Amoroso 2019). Nonetheless, documenting and collecting the flora of
this range has been successful (Coritico et al. 2020a, b). Preliminary data from this work demonstrate that the range
boasts a high degree of biodiversity and endemicity. For example, many of the new Philippine Nepenthes Linnaeus
(1753: 955) species discovered and described during the past decade were discovered in this range (Gronemeyer et al.
2011a, b, 2014; Lagunday et al. 2017; Lagunday & Amoroso 2019).
Previous fieldwork aimed at collecting specimens in botanically unexplored areas of the Philippines has led to
the discovery of several new Vaccinium species (e.g. Salares et al. 2018; Fritsch et al. 2020; Tamayo et al. 2021).
These discoveries have demonstrated that Philippine Vaccinium contains a higher species diversity than previously
understood. Presently, 36 species of Vaccinium are recognized for the Philippines (Pelser et al. 2011 onwards; Tamayo
et al. 2021). However, many of these need further taxonomic study, as half of the Philippine Vaccinium species are rare
and often known only from the type locality (Vander Kloet 1996).
During fieldwork in Mt. Tago Range (2014–2019) as part of botanical survey projects, a species of Vaccinium
with a scrambling habit, notably wide leaves, and lustrous white flowers was vouchered. The specimen was compared
with other similar species and based on detailed study of its vegetative and floral characters it is concluded that the
specimen represents an undescribed taxon. Here we describe this new species under a morphological species concept
(Cronquist 1978) and provide an illustration and photographic images of the living plants. This discovery raises the
number of species of Vaccinium documented in the Philippines to 37.
Materials and Methods
Dried herbarium specimens and in situ photographs of the plant were used as the basis for the description. Microscopic
details were described with the use of an AmScope stereomicroscope of up to 64× magnification. Herbarium specimens
from A, BRIT, CAS, and K were also studied for comparison. The taxonomic works of Copeland (1930), Sleumer
(1966–1967), Argent (2008), and recent Philippine Vaccinium literature (Co et al. 2002; Salares et al. 2018; Fritsch et
al. 2020; Tamayo et al. 2021) were consulted to ensure the uniqueness of the species and to compare it with the other
known species of Vaccinium in the region.
Taxonomic Treatment
Vaccinium carmesinum M.N.Tamayo & P.W.Fritsch, sp. nov. (Figs. 1–2)
Type:—PHILIPPINES. Mindanao Island, Bukidnon Province, Municipality [City] of Malaybalay, Barangay Kibalabag, Mt. Limbawon,
[Mt. Tago Range,] accessory trail to peak, 8.26217°N, 125.18055°E, 1546 m elevation, 10 June 2019, Plants and Lichens of the
Southern Philippines Survey 611 (holotype PNH!, isotypes A!, BRIT BRIT572077!, CAS!, CMUH!, NY!).
Paratypes:—PHILIPPINES. Mindanao Island: Province of Bukidnon, Municipality [City] of Malaybalay, Barangay
Kibalabag, Mt. Limbawon, [Mt. Tago Range,] open area with Pandanus, 8.27577°N, 125.18333°E, 1832 m elevation,
30 June 2015, Peter W. Fritsch 2081 (BRIT BRIT554025!, CAS 490415!); Mt. Kiamo summit, [Mt. Tago Range,]
on ridge of heathland scrub, 8.2563°N, 125.14799°E, 1760 m elevation, 7 May 2014, Darin S. Penneys 2377 (BRIT
BRIT554030!, CAS 490401!).
Diagnosis:—Vaccinium carmesinum resembles V. platyphyllum Merrill (1917: 294) and V. luzoniense S.Vidal
(1886: 168), but differs from the former by longer and wider leaves, longer racemes, longer bracts, glabrous corollas,
and glabrescent fruits, and from the latter by longer petioles, leaf glands distributed along the length of the blade
margin, a glabrous inflorescence rachis, and lanate filaments.
VACCINIUM CARMESINUM Phytotaxa 533 (3) © 2022 Magnolia Press • 175
FIGURE 1. Vaccinium carmesinum. A. Leafy branchlet. B. Inflorescences showing flower buds and foliaceous bracts. C. Corolla. D.
Dissected flower showing stamens, style, ovary, hypanthium, and calyx. E. Dorsal view of stamen. F. Ventral view of stamen. G. Fruit
cross section. Illustration by MNT.
TAMAYO ET AL.
176 • Phytotaxa 533 (3) © 2022 Magnolia Press
FIGURE 2. Vaccinium carmesinum. A, B. Flowering branchlet. C. Leafy branchlet. D. Inflorescences showing flower buds and foliaceous
bracts. E. Longitudinal section of flower showing corolla, style and stamens. F. Immature fruits. Photographs: A, D by DSP; B, E, F by
MAKP; C by DNT.
VACCINIUM CARMESINUM Phytotaxa 533 (3) © 2022 Magnolia Press • 177
Description:—Terrestrial leaning shrub or tree, evergreen, 2–5 m tall, sparsely branched. Branchlets glabrous,
red when young, grayish brown at maturity, terete, 3–8 mm wide, lenticellate; perennating buds compressed-ovoid,
1.5–2.5 mm long; bud scales overlapping with minutely ciliate margins. Leaves persistent on older branchlets, spirally
and evenly arranged, slightly overlapping, internodes 1–5 cm long; petiole crimson red, in cross section rounded
abaxially and slightly raised adaxially, 10–18 × 1–5 mm, glabrous; lamina broadly elliptic, ovate, or rarely subrounded,
with the larger leaves on each branchlet 7–15 × 0.4–9 cm, coriaceous, both surfaces reddish when young turning pale
green abaxially and glossy adaxially, in sicco both surfaces light brown to ferrugineous, without punctae, glabrous;
midvein flattened or sunken adaxially, strongly raised abaxially, secondary veins 3 or 4 on each side of midvein with
first pair arising from base and remainder along midvein, arc-ascending, abaxially raised, adaxially sunken, tertiary
veins faintly evident or obscure, base cuneate to truncate, margin entire, weakly revolute, apex slightly acuminate,
marginal glands sunken, 10–18 per side, scattered along length of margin but more concentrated towards the apex, 0.3–
0.5 mm wide. Inflorescence pseudo-terminal or terminal, racemose, developing beyond confines of perennating bud, 1
per axil, 6–8 cm long at anthesis, densely 10- to 12-flowered; peduncle and rachis crimson red, slightly ridged, terete,
glabrous; flower bracts caducous, crimson red, dark brown in sicco, foliaceous, ovate to elliptic, planar or occasionally
cucullate, 6–15 × 2.5–3.5 mm, coriaceous, glabrous, margin entire, ciliolate, with several yellowish or reddish globose
glands, 0.15–0.20 mm diameter mainly on basal half and with cilia ca. 0.1 mm mainly on apical third, apex acute
to obtuse with a terminal gland. Pedicel 3.5–15 × 0.5–0.9 mm at anthesis, terete, spreading, glabrous, occasionally
with 1 or 2 globose glands near base or occasionally on apical half, ebracteolate. Flowers articulated at junction with
pedicel, 3.5–12 mm long. Hypanthium crimson red, cupuliform, 1.5–2 × 2–2.5 mm, white-hirsutulous with trichomes
0.10–0.15 mm long; calyx limb 1.0–1.2 mm long, white-hirsutulous; calyx lobes broadly triangular, 0.8–1.2 mm long,
white hirsutulous, margin entire, often ciliolate, apex acute, with a prominent greenish (reddish in sicco) globose
sessile terminal gland ca. 0.25 mm diameter. Corolla broadly acute, lustrous white, conical-urceolate, 7–12 × 2.5–6
mm, outside glabrous, inside white-lanate especially on upper and lower third, trichomes 0.5–1 mm long; corolla lobes
5 or 6, 1–2 × 1–1.5 mm, apex acute to obtuse. Stamens 8 to 10, monomorphic, distinct, 5.5–7.2 mm long; filaments
white, straight, gradually dilated at base, 3.5–4.8 mm long, pink towards base, densely white-lanate with trichomes
0.5–1 mm long; anthers 2–2.4 mm long, cells 1.4–1.6 mm long, echinulate, tubules parallel, narrowly cylindrical,
distinctly narrower than cells, opening by oblique ventrally oriented apical pores, 0.6–0.8 mm long, pore apex rounded,
spurs absent. Ovary 5- or 6-locular but appearing pseudo-10- to 12-locular with false partitions extending ca. 1.5 mm
from inner wall; ovules in two columns per locule. Disk disciform, ca. 2 mm in diameter, puberulent, margin shallowly
ridged. Style reddish, not exserted from corolla, 10–12 mm long, glabrous. Fruit on pedicels 1.4–2.1 cm long, deep
purple, dark brown or reddish in sicco, globose, slightly ridged, 4–6 × 4–6 mm, glabrescent except for minute cilia on
calyx lobe margins; persistent calyx lobes erect; disk ca. 4.5 mm in diameter. Seeds numerous, minute, brown, ca. 0.8
mm long.
Distribution and Habitat:—Vaccinium carmesinum is endemic to two mountains (Mt. Kiamo and Mt. Limbawon)
in Mt. Tago Range, Mindanao, growing in tropical lower montane rainforest to upper montane rainforest. Populations
of V. carmesinum were mostly found near summits where they thrive on volcanic-igneous or clay substrate with
abundant humus. They also occur in areas of open shaded mossy forests, or on ridges covered in heathland scrub.
Paratypes of the new species were collected on ca. 10–30% west-facing slopes.
Etymology:—The epithet carmesinum is derived from the Greek word for crimson (blood red), as depicted by its
crimson red petioles, floral bracts, peduncle, rachis, pedicels, hypanthium, and calyces. Moreover, a crimson red stain
is extracted in notable quantity when the plants are soaked in a denatured alcohol solution.
Phenology:—Flowering in June. Fruiting from January to May.
Proposed Conservation Status:—Mt. Tago Range has not been extensively explored botanically, which results in
uncertainty as to the conservation status of the species. This range is a non-protected area; thus, the extent of occurrence
and area of occupancy for the species cannot be assessed. There are only two populations currently known. Hence, we
recommend a conservation status of data deficient (DD) (IUCN Standards and Petitions Committee 2019).
Discussion:—In its combination of morphological characters, Vaccinium carmesinum matches no other species
treated in relevant taxonomic treatments. In the artificial key to the species of Philippine Vaccinium (Copeland 1930),
V. carmesinum keys to V. platyphyllum. The new species differs from V. platyphyllum by having longer and wider
leaves (7–15 × 0.4–9 cm vs. 11–14 cm × 5–7 cm), longer racemes (6–8 cm vs. 4–6 cm), longer bracts (6–15 mm vs.
ca. 8 mm), longer pedicels (3.5–15 mm vs. ca. 8 mm) that are glabrous (vs. slightly pubescent) and ebracteolate (vs.
bracteolate), a glabrous (vs. sparsely pubescent) corolla outside, longer anthers (2.0–2.3 mm vs. ca. 1.5 mm), and
longer (4.0–6.0 mm vs. ca. 3 mm) and glabrescent (vs. pubescent) fruits (Merrill 1917).
TAMAYO ET AL.
178 • Phytotaxa 533 (3) © 2022 Magnolia Press
Vaccinium carmesinum can be distinguished from all other species of Philippine Vaccinium by its leaves, which
are the widest of any Vaccinium in the Philippines. The pedicels are also notably ebracteolate and have 0 to 2 globose
glands near the base or occasionally on the apical half. These glands might be homologous with bracteoles (typically
two per pedicel in Vaccinium) with a reduction in size and/or number. Copeland (1930) mentioned pedicel glands in
V. luzoniense. Unfortunately, this character was not thoroughly described for the other Philippine species in former
publication where the absence of bracteoles in a specimen is noted as “unobserved” (i.e. Sleumer 1966–1967).
Vaccinium carmesinum is a member of V. section Bracteata Nakai in Nakai & Koidzumi (1927: 234) sensu
Sleumer (Sleumer 1966–1967) as based on the combination of many-flowered racemose inflorescences, caducous
foliaceous bracts, absence of a membranaceous wing at the sinuses of the corolla, and anthers that open by short
introrse slits or terminal pores (Sleumer 1966–1967; Co et al. 2002; Salares et al. 2018). In Sleumer’s (1966–1967)
key to the Malesian V. section Bracteata, V. carmesinum keys to V. luzoniense. Vaccinium carmesinum differs from
V. luzoniense, however, by having longer petioles (10–18 mm vs. ca. 10 mm), longer and wider leaves (7–15 × 0.4–9
cm vs. 7–9 cm × 3–4.5 cm), with leaf glands distributed along the length of the leaf margin (vs. with merely a pair
of glands near the base), glabrous rachis (vs. with capitate-glandular trichomes), white (vs. red) corollas, and densely
lanate (vs. sparsely pubescent) filaments (Vidal 1886; Copeland 1930).
In the key to the Bornean species of Vaccinium (Argent 2018), V. carmesinum keys to V. sarawakense subsp.
montanum Argent (2018: 108) but differs from it by having an inflorescence with fewer flowers (10- to 12-flowered vs.
7- to 20-flowered), glabrous rachis (vs. densely covered by short brown curved glandular trichomes), calyx lobes with
a sessile terminal gland (vs. absent), white (vs. pale pink) corollas, and the absence of anther spurs (vs. presence).
In the sectional treatment of Vaccinium (Vander Kloet and Dickinson 2009), V. carmesinum can be treated as a
member of V. section Euepigynium Schlechter (1919: 174) by its evergreen habit, monomorphic perennating buds, each
with more than two scales, one perennating bud per leaf axil, plinerved leaf blade venation, entire leaf blade margin,
peduncle longer than pedicels, calyx tube completely fused to the ovary, and pseudo-10-locular ovary. However,
the boundaries of V. section Euepigynium and other sections of Malesian Vaccinium delimited by Vander Kloet and
Dickinson (2009) were vaguely defined (i.e. the species included in each section are not provided). Hence, the sectional
limits of Vaccinium in Malesia need further study.
During the process of diagnosing Vaccinium carmesinum as distinct from other Philippine species, we have
become cognizant of problems in the taxonomy of the Philippine species. For example, V. ilocanum Merrill (1919:
441) and V. rizalense Merrill (1925: 43) were synonymized under V. platyphyllum by Copeland (1930) but characters
seem divergent among these species and the justification relied mainly on macroscopic characters. A detailed study
of this complex is currently in progress with emphasis on, e.g., ovary indumentum, corolla surfaces, and stamen
characters.
Acknowledgments
We thank the members of the Mindanao expedition team for helping to make this study possible: Florfe M. Acma,
Daniel L. Nickrent, Alice Gerlach, Vanessa Handley, Gordon McPherson, Lydia Marie Hicks, April Joie D. Lagumbay,
Jennifer G. Opiso, Jorgen Abellera, Joevina Nobleza, John Michael M. Galindon, Jeffrey P. Mancera, Noel E. Lagunday,
Aimanuelzon Yorong, Aldrin L. Hongco, J. Peter Quakenbush, Mescel S. Acola, Yvonne Love L. Cariño, Noe P.
Mendez, and Mc Andrew K. Pranada. We also thank Barangay Captain Shemuel Lagunday and local researchers of
Barangay Kibalabag, Malaybalay, Bukidnon for assistance during fieldwork. We also thank the Higaonon indigenous
tribe of Bukidnon and the Department of Environment and Natural Resources (DENR) for the issuance of gratuitous
permit; National Museum of the Philippines and Central Mindanao University (CMU) for logistical support. Financial
support for this project was provided by the Department of Science and Technology-Grants in Aid (DOST-GIA) to
CMU, and U.S. National Science Foundation grants DEB-1754667 to DSP and DEB-1754697 to PWF. MNT thanks
the College of Science and Engineering at Texas Christian University, the U.S. National Science Foundation, and the
Botanical Research Institute of Texas for additional financial support.
VACCINIUM CARMESINUM Phytotaxa 533 (3) © 2022 Magnolia Press • 179
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characteristics and diversity of East Asian plants. China Higher Education Press, Beijing, pp. 114–119.
Vander Kloet, S.P. & Dickinson, T.A. (2009) A subgeneric classification of the genus Vaccinium and the metamorphosis of V. section
Bracteata Nakai: more terrestrial and less epiphytic in habit, more continental and less insular in distribution. Journal of Plant
Research 122 (3): 253–268.
https://doi.org/10.1007/s10265-008-0211-7
Vidal, S. (1886) Revision de plantas vasculares Filipinas, Fam. 67 - Vacciniaceas. Establecimiento Tipo-Litográfico de M. Perez, Hijo,
Binondo, Manila. pp. 1–455.
APPENDIX 1. Additional Vaccinium specimens examined for morphological comparison.
Vaccinium luzoniense S.Vidal. PHILIPPINES. Luzon Island: Province of Benguet (District of Lepanto), S. Vidal 1535
(holotype K K000780748!).
Vaccinium platyphyllum Merrill. PHILIPPINES. Mindoro Island: Lagating (Mt. Halcon), Oriental Mindoro, 15 April
1991, B.C. Stone et al. PPI 758 (BRIT BRIT26871!); Barangay Lantuyan (Mt. Halcon), Oriental Mindoro, ca. 1500 m
elevation, 15 March 1997, G. Argent et al. PPI 20095 (BRIT BRIT26880!).
