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J. Bot. Res. Inst. Texas 14(2): 281 – 287. 2020
https://doi.org/10.17348/jbrit.v14.i2.1009
VACCINIUM HAMIGUITANENSE (ERIC ACE AE),
A NEW SPECIES FROM THE PHILIPPINES
Peter W. Fritsch Victor B. Amoroso
Botanical Research Institute of Texas Center for Biodiversity Research and
1700 University Dr. Extension in Mindanao (CEBREM) and
Fort Worth, Texas 76107, U.S.A. Depar tment of Biology, College of Arts and Sciences
pfritsch@brit.org Central Mindanao University
Musuan, Bukidnon 8710 PHILIPPINES
victorbamoroso@gmail.com
Fulgent P. Coritico Darin S. Penneys
Center for Biodiversity Research and Department of Biology and Marine Biology
Extension in Mindanao (CEBREM) and University of Nor th Carolina Wilmington
Department of Biology, College of Arts and Sciences 601 S. College Road
Central Mindanao University Wilmington, North Carolina 28403, U.S.A.
Musuan, Bukidnon 8710 PHILIPPINES penneysd@uncw.edu
cfulgent@gmail.com
aBsTr acT
Vaccinium hamiguitanense, a new species from the Philippines, is described and illustrated. The new species is most si milar to V. g it in‑
gense Ho ok. f. but differs b y having sma ller leaf blades, lea f blade margi ns with 2 to 4 impresse d more or less evenl y distributed crenations
(gla nds) per side, in flores cences wit h fewer flow ers, shorter p edicels t hat are pube rulent and mu riculate , and a glabrou s floral d isk. The new
specie s is endemic t o Mt. Hamigu itan Ran ge Wildlife S anctuar y in Davao Or iental Prov ince of Mind anao Isla nd in Tropical Uppe r Montane
Rai n Forest and low (“bons ai”) forest on clay d erived from u ltrama fic rock. We as sign an IUCN R ed List prel imina ry statu s as Data Def icient.
aBsTr ak
Inilalarawan sa ulat na ito ang isang bagong species ng halaman mula sa Pilipinas, ang Vaccinium hamiguitanense. Kahawig ng bagong
specie s ang V. gitinge nse Hook f., ng unit mas maliit ang mga dahon, bawat isa ay may 2 hanggang 4 na kapansin-pansin at halos pantay-
pantay na mga um bok sa parehong g ilid (mga glandu la), mas k akaunti ang b ulaklak k ada kumpol, b awat isa ay mas m aikli ang t angkay na
pinalilibut an ng maliliit ngun it magagaspang na buhok, at ang floral di sk ay mak inis. Ang bagong specie s ay matatagpuan lamang sa Mt.
Hami guitan Ra nge Wildlif e Sanctuar y sa lalaw igan ng Davao Or iental, isl a ng Mindan ao, partik ular sa mat ataas na ba hagi ng kag ubatan at
sa kag ubatang “bon sai,” kung saa n ang lupa ay luwad at h ango sa batong u ltramaf ic. Binibig yan namin n g paunang st atus na Data D eficient
ang bagon g species sa I UCN Red List.
key words: Endemic, IUCN R ed List, Min danao, new spe cies, Philip pines, Vaccinium
inT roducTion
The genus Vaccin iu m L. (Ericaceae: Vaccinoideae: Vaccinieae) comprises ca. 450–500 species distributed
among all continents except Australia and Antarctica, with its center of diversity in Malesia (Sleumer 1966–
1967; Fang & Stevens 2005; Vander Kloet & Dickinson 2009; Argent 2014). The taxonomic limits of the genus
are still uncertain, with one treatment provisionally placing most of the Malesian species of Vac cinium in an
expanded concept of Agapetes, rendering Vaccin iu m as comprising ca. 140 species (Stevens et al. 2004), and
another segregating the mainly Bornean genus Rigiolepis (Hook. f.) Sleumer of ca. 23 species from Vaccinium
(Argent 2019). DNA sequence data strongly suggest that the genus is highly para- or polyphyletic (Kron et al.
2002), but much more data will be needed to resolve the classific ation of Vacc inium and the Vaccinieae (Stevens
et al. 2004). For the purpose of the present study, we follow the circumscription of Va cc inium as presented in
Sleumer (1966–1967) and Vander Kloet & Dickinson (2009).
The two most comprehensive taxonomic treatments of Vac ciniu m for the Philippines are those of
Copeland (1930) and Sleumer (1966–1967). Copeland’s work focused specifically on the Philippines, whereas
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This article has been licensed as Open Access by the author(s) and publisher.
This version supersedes any other version with conflicting usage rights.
282 Journal of the Botanical Research Institute of Texas 14(2)
Sleumer’s more up-to-date work was part of the treatment of the genus for all of Malesia. As based on these
publications and more recent species discoveries (Co et al. 2002; Salares et al. 2018), there are 34 species of
Vac ciniu m currently recognized in the Philippines, 31 of which are endemic (Argent 2008; Pelser et al. 2011
onwa rds).
During field work in June of 2015 and 2019 in Mt. Hamiguitan Range Wildlife Sanctuary in Davao
Oriental, Mindanao Island, Philippines, the authors encountered a species of Vacc inium that did not key out in
the species keys of Copeland (1930) and Sleumer (1966–1967) and matched none of the descriptions in these
works. On further morphological comparison of our collections with type material of Malesian Va cci nium at
JSTOR Global Plants (https://plants.jstor.org), herbarium specimens at BRIT, CAS, and CMUH, and images of
in situ plants online, we concluded that the species is new to science. The new species status is also supported
by a photograph of a plant on PhytoImages (Nickrent et al. 2006 onwards) taken at Mt. Hamiguit an by L. Co in
2011 labeled “Vac cinium plant7” but which is clearly this species. It is described and illustrated below.
TaxonoMic Tr eaT MenT
Vaccinium hamiguitanense P.W. Fritsch, sp. nov. (Figs. 1–2). Type: PHILIPPINES. Mindanao island. Davao Oriental:
Municipality of San Isidro, Barangay La Union, Mt. Hamiguitan R ange Wildlife Sanctuary, between second Lantawan and pygmy
forest, at t he peak, 1641 m, 6.73998°N, 126.1821°E, 23 Jun 2015 (fl), P.W. F rit sch 20 27 (holoType: PNH!; isoTypes: BRIT554024!,
CAS490410!, CMUH!).
Haec species Vaccinio gitingensi Elmer simillima, sed ab eo foliis parvioribus ad margine crenulatis, floribus inflorescentiae paucioribus,
pedic ellis brev ioribus puber ulis mur iculatis, d isco glabro d iffert.
Shrubs, terrestrial, evergreen to 1.5 m tall, densely branched. Branchlets with white, erect, straight trichomes
to 0.14 mm long, mature branchlets dark maroon, slightly compressed and often ridged, 0.5–0.9 mm t hick, not
lenticellate, outer surface grayish, peeling on older br anchlets; perennating buds compres sed-ovoid, to 0.8 mm
long, with several obscurely overlapping scales. Leaves persistent on older branchlets, densely crowded, spi-
rally arranged; petiole in vivo green or often red, 1.2–1.8 mm × 0.5– 0.6 mm, 1.8–2.6 times a s long as wide, with
white erect straight trichomes to 0.14 mm long, glabrescent on leaves of old branchlets, in cross-section abaxi-
ally rounded, adaxially nearly flat; leaf blade elliptic or slightly obovate, larger leaves on each branchlet 8.8–
16.0 × 4.0–6.5 mm, cor iaceous, both s urfaces bri ght pink when young, glabrou s except occasion ally puberule nt
at base, smooth, abaxial surface without punctae, light green in vivo, light brown in sicco, adaxial surface green
in vivo, dark brown in sicco, base cuneate, margin crenulate by 2 to 4 impressed more or less evenly distributed
crenations (glands) per side, thinly recurved, apex obtuse to rounded, often emarginate, the very tip with a
gland and often a few cilia, impressed marginal glands 2 to 4 per side, scattered along length of margin, ca. 0.14
mm diam., midvein slightly raised abaxially, planar or sulcate adaxially, secondary veins 1 to 4 on each side of
midvein, the first one or two pairs arising from the base, the remainder along the midvein, arc-ascending,
slightly raised or obscure abaxially, obscure adaxially, tertiary veins faintly evident or obscure. Inflorescences
axillary or usually pseudoterminal, racemose and developing beyond confines of perennating bud, 1 per axil,
ca. 1.6 cm long at anthesis, (2-) 4- to 6-flowered; rachis white-puberulent and occasionally scattered orange-
muriculate with glandular trichomes up to 0.06 mm long; bracts subtending pedicels ± foliaceous, persistent,
greenish and often flushed red in vivo, brown in sicco, narrowly elliptic to suborbicular, planar or occasionally
cucullate, 5.9–8.3 × 3.5–4.0 mm, coriaceous, margin entire or crenulate by 1 or 2 impressed crenations
(glands) per side, apex obtuse to subrounded or slightly emarginate, often with a few cilia at very tip. Flowers
articulated at junction with pedicel, 6 –7 mm long. Pedicel nodding, 2.5–3.5 × 0.5–0.6 mm at anthesis, white-
puberulent and orange-muriculate; bracteoles caducous, 2, borne at base of pedicel, narrowly deltoid to
oblong, planar or nearly so, 0.14–0.54 mm long, puberulent, margin entire, apex sharply acute. Hypanthium
green or flushed pale red and shining in vivo, cupuliform, 1.1–1.2 × 1.9–2.4 mm, glabrous or scattered white-
puberulent; caly x limb 1.1–1.3 mm long, glabrous; ca lyx lobes broadly deltoid, 0.5– 0.6 mm long, glabrous both
sides, margin ciliate on upper half, eglandular, apex acute, the very tip eglandular or with a small gland.
Corolla in bud closed and strongly 5-ribbed in line with petal midveins, at anthesis gamopetalous for ca. 5⁄6 of
total length, white, broadly urceolate, 4.6–5.0 × ca. 2.6 mm, outside glabrous except on lobes, inside glabrous;
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Fritsch et al., A new species of Vaccinium from the Philippines 283
Fig. 1. Vaccinium hamiguitanense. A. Flowering branchlet. B. Leaf, abaxial view. C. Pedicel and flower. D. Distal portion of pedicel, hypanthium, calyx,
and style. E. Ovary in cross section showing axile placentation and five locules, each locule with incomplete false partition emanating from ovary wall
and two locules shown in each of four of the locules. F. Distal portion of filament, and anther, oblique-lateral view. G. Stamen, ventral view. H. Infruc-
tescence showing bracts, bracteoles, and fruit. A foliaceous bract subtends each pedicel, and the bracteoles can be seen as minute narrowly deltoid
structures at the bases of the two proximal pedicels. A–G based on P.W. Fritsch 2027 (BRIT, CAS) and images of the living plants; H based on P.W. Fritsch
1984 (BRIT, CAS) and images of the living plants. Illustration by Samantha Peters.
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284 Journal of the Botanical Research Institute of Texas 14(2)
Fig. 2. Images of living Vaccinium hamiguitanense and its habitat. A. Habitat at peak of Mt. Hamiguitan of low (“bonsai”) forest. B. Habit with leaves
predominantly in abaxial view. C. Branchlet with inflorescence. D. Inflorescence with flowers in bud and (in apical view) at anthesis. E. Immature fruit
with persistent bracts. F. Submature fruit, apical view. Note small glands at apex of appressed persistent calyx lobes. B, C, P.W. Fritsch 2027; photographs
by P.W. Fritsch. D–F, Plants and Lichens of the Southern Philippines Survey 725; photographs by J.G. Opiso.
corolla lobe s 5, ovate, ca. 1.0 × 0.5 mm, white- setose abaxially on midvein, apex obtuse. Sta mens 10, monomor-
phic, distinct from each other, 2.8–3.5 mm long; filaments linear, 1.9–2.0 mm, white-villous on all sides, also
spars ely echinulate in l ines, with tr ichomes to 0.34 mm long; anthers 1.2–1.9 mm long, anther cel ls echinulate,
0.7–1.0 mm long, tubules parallel, broadly cylindric, 0.6–0.8 mm long, about as wide as cells, opening by
oblique ventrally oriented apical pores, pore apex rounded, spurs slightly upcurved, 0.2–0.4 mm long. Ovary
5-locular but appearing somewhat pseudo-10-locular with false partitions extending ca. 0.16–0.20 mm from
inner wall; ovules in two columns per locule, each column separated by false partition; disk glabrous; style not
exserted from corolla, 3.5–4.0 mm long, glabrous. Infructescence with persistent bracts (persisting even after
mature fruit is gone), bracts planar or slightly cucullate, often incurved and touching fruit; fruit pseudo-10-
locular, in vivo turning burgundy and then dark purple to black at maturity, subglobose, 3.8–8.0 × 4.0–7
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Fritsch et al., A new species of Vaccinium from the Philippines 285
mm, glabrous or sparsely puberulent, persistent calyx lobes incurved and appressed to disk, lobe apex with a
small gland; disk ca. 2 mm in diameter, glabrous.
Etymolog y.—The specific epithet refers to Mt. Hamiguitan, to which the species is endemic.
Distribution, Habitat, and Phenolog y.—The new species is endemic to Mt. Hamiguitan Range Wildlife
Sanctuary in Davao Oriental Province of Mindanao Island. It is known from Tropical Upper Montane Rain
Forest and “bonsai” forest (i.e., predominant vegetation of densely crowned sclerophyllous trees and shr ubs of
low stature) on ridges and at the peak of Mt. Hamiguitan at 1081–1641 m elevation. It is common, growing at
least in part in clay derived from ultramafic bedrock. The species is known to flower and fruit in June.
Additional specimens examined (Paratypes).—PHILIPPINES. Mindanao island. Davao Oriental: Municipality of San Isidro, Ba rangay La
Union, Mt. Hamiguitan R ange Wildlife Sanctuary, between second Lantawan and pygmy forest, 1201 m, 6.73069°N, 126.1795°E, 22 Jun
2015 (fr), P.W. Fr it sc h 19 84 (BRIT55402 6!, CAS490 408!, CMUH!, PNH!); ibid., along “ Superfer ry Trail,” 1081 m, 6.72857°N, 126.17730°E , 17
Jun 2019 (fl, fr), Plants a nd Lichens of the So uthern Philipp ines Survey 725 (B RIT!, CM UH!, PNH!).
Conservation Assessment.—Because Vaccinium hamiguitanense has been collected from only a single locality,
we categorize it as Data Deficient (DD) in accordance with the IUCN Red List Criteria (IUCN Standards and
Petitions Subcommittee 2019). However, the collections made of the species have all been from within the
boundaries of Mt. Hamiguitan Range Wildlife Range Sanctuary and this t axon has been afforded protection. It
is a common species along the trails traversed by the authors at the elevations within which it grows, with at
least 100 individuals seen.
Discussion.—Vaccinium hamiguitanense can be placed in V. sect. Bracteata Nakai sensu Sleumer (1966–
1967) by its multi-flowered racemes, c alyx lobes shorter than t he hypanthium, corolla consisting of one homo-
geneous layer without a membranaceous wing at the sinuses, and anther tubules opening by short introrse
pores. The sectional classification of Va ccinium has been revised by Vander Kloet and Dickinson (2009), with
V. sect. Bracteata divided into several smaller sections. Using the key in their treatment, one can place the new
species in V. sect. Euepigynium Schltr. by its perennating buds being monomorphic, one per leaf axil, and cov-
ered by > 2 scales; leaves persisting > 18 months and having palmate venation; inflorescence rachis being
robust and bracteate; hypanthium being fused to the ovary; and ovary being pseudo-10-locular, the disk being
scarcely if at all visible when in ripe fruit.
In the key to the Malesian species of Vacc inium sect. Bracteata in Sleumer (1966–1967), the new species
would fall within lead 1 of couplet 1: subtending bract of each flower conspicuous, ± foliaceous, persisting for
some time during anthesis, mostly so to the fruiting stage, and lead 1 of couplet 2: leaves manifestly cren(ul)ate
or cren(ul)ate-serrate all along t he margin. Under lead 1 of couplet 2, the spe cies keys best to V. gitingense Elmer
by its calyx lobes not being callose-thickened at the apex and lacking any apical or marginal glands (although
some of the lobe s can have a faint gl and). However, the new species is di stinguishable from V. gitingense by hav-
ing smaller leaf blades (8.8–16 × 4.0–6.5 mm versus 15–35 × 8–15 mm), crenulate leaf blade margins (versus
crenulate-serrulate), inflorescences 3- to 5-flowered (versus 5- to 10-flowered), pedicels 2.5–3.5 mm long at
anthesis (versus 7–15 mm), pedicels puberulent and muriculate (versus glabrous), and a glabrous disk (versus
pubescent). Under lead 2 of couplet 2, the species keys best to lead 1 of couplet 29 by having glabrous leaves
0.8–1.5 × 0.4–1 cm, calyx lobes not callose-thickened at the apex and lacking glands (although some of the
lobes can have a faint gland), anther tubules as long as and as wide as the cells, and anthers with 2 spurs.
However, it fits neither of the taxa under this lead (V. scortechinii King & Gamble and V. mi q u e lii Boerl. var.
miquelii) because they both have leaf blades with entire margins except for one pair of marginal glands within
the lower third. The new species can be further distinguished from V. scortechinii by leaves abaxially without
punctae (versus with punct ae), hypanthium glabrous (versus pubescent), and corolla glabrous (versus densely
pubescent on both sides). It can be further distinguished from V. mi qu elii var. miquelii by its leaves abaxially
without punctae (versus with punctae), inflorescences 3- to 5-flowered (versus 10- to 20-flowered), hypan-
thium glabrous (versus pubescent), corolla glabrous inside (versus pubescent), and disk glabrous (versus
pubescent).
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286 Journal of the Botanical Research Institute of Texas 14(2)
In the key to Bornean species of Vacc inium in Argent (2019), the new species keys to V. coriaceum Hook. f.
by its leaf blades < 20 mm long and 15 mm wide, leaf margins with at least a pair of glands (crenations of
Sleumer 1966–1967), and inflorescence > 3-flowered. However, the new species is easily distinguished from V.
coriaceum (va r. coriaceum sen su Argent 2019) by its inflorescences 3- to 5-flowered (versus > 5-flowered), pedi-
cels muriculate (versus not muriculate), anthers without gland-tipped trichomes (versus with gland-tipped
trichomes), anther tubules about as wide as the cells (versus much narrower than the cells), anther pores
obliquely cut (versus transversely cut), and disk glabrous (versus pubescent).
In the artificial key to the species of Philippine Vaccin iu m (Copeland 1930), the new species would key to
V. myrtoides Miq. by the following characters: leaves < 3 cm long, flowers in racemes, racemes with a foliaceous
bract subtending each pedicel, and pedicels 1 cm long or less, not longer than the flowers. The new species is
easily distinguished from V. myrtoides, however, by its leaf margins crenulate by 2 to 4 impressed more or less
evenly distributed glands (crenations of Sleumer 1966–1967) per side (versus 1 or 2 impressed glands in the
lower third); inflorescences 3- to 5-flowered (versus 4- to 12-flowered); pedicels 2.5–3.5 mm long (versus 0.5–
1.4 mm), muriculate (versus not muriculate); anthers with spurs (versus without); and disk glabrous (versus
pubescent).
Vaccinium hamiguitanense is similar to plants from Dinagat Island, Province of Loreto, Cambinliw-
Redondo Bonsai Forest, 10°21'10.20"N, 125°38'02.08"E, 24 Nov 2014, documented with photographic images
by P.B. Pelser & J.F. Barcelona as displayed on Co’s Digital Flora of the Philippines (Pelser et al. 2011 onwards)
as “Vaccin iu m plant3.” Although the images are lacking flowers, the fruiting branches are similar to the new
species in their densely crowded small elliptic leaves with 2 to 4 marginal impressed crenations (glands) per
side. These plants appear to differ from those from Mt. Hamiguitan at least by their apparently single-flowered
inflorescences (which may place them in V. sect. Oarianthe Schltr.), persistent and much larger bracteoles (up
to ca. 8 mm long), and puberulent hypanthium/fruit. The Dinagat plants may represent another undescribed
species but apparently have yet to be collected in flower.
The discovery of this new species brings the number of currently recognized species of Vacciniu m in the
Philippines to 35. A key to Philippine Vac ciniu m species that updates the keys of Copeland (1930) and Sleumer
(1966–1967) is needed (Argent 2008). Field observation s suggest that there are sti ll new species in t he genus to
be discovered and described (Pelser et al. 2011 onwards; Fritsch et al. 2016), and the species need assessment
as to sectional classification based on morphological and DNA sequence evidence.
The new species is endemic to Mount Hamiguitan Range Wildlife Sanctuary. The only other species of
Vac ciniu m documented from this range are V. cf. barandanum S. Vidal, endemic to the Philippines (24 June
2015, P.W. Fritsch 2048, sterile, BRIT!, CAS!, CMU!, PNH!) and V. gitingense Elmer, endemic to the islands of
Mindanao and Dinagat in the Philippines (24 June 2015, P.W. F r i t s ch 2 0 43, flower, fruit, BRIT!, CAS!, CMU!,
PNH!). The Mount Hamiguitan Range Wildlife Sanctuary covers 6,834 ha located between 6°40'N to 6°47'N
and 126°09'E to 126°13'E. It is known for its unique characteristics and the largest “bonsai” forest in the
Philippines. The substrate in this forest type is derived from ultramafic bedrock, with high concentrations of
iron and magnesium (Amoroso et al. 2009). Perhaps because of this unusual forest type and the general isola-
tion of the range on a narrow peninsula, species endemism is high in the range, with 163 endemic species of
vascular plants documented in 2009 (Amoroso et al. 2009) and with several new endemic species discovered
since (e.g., McPherson 2010; Karger et al. 2012; Naive 2017). With much of the area still unexplored botani-
cally, more plant species new to science are expected to be discovered in the coming years.
acknowled GMe nTs
We thank the office of the Department of Environment and Natural Resources Region XI and Protected Area
Management Board (PAMB) of Mt. Hamiguitan Range Wildlife Sanctuary for the permit, Forester Ruel D.
Colong for logistical support, the assistance of local researchers Alfredo P. Bolante Sr. and Felipe S. Gorme Jr.,
Central Mindanao University for logistical support, the herbarium staff of BRIT, CAS, and CMUH for access to
specimens, Jennifer G. Opiso for photographs, Samantha Peters for the illustration, Jef Mancera for translating
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Fritsch et al., A new species of Vaccinium from the Philippines 287
the Abst ract to Tagalog, and reviewers Pieter Pelser, Val B. Sal ares, and Ken Chambers for helpful comments on
the manuscript. This research was supported by U.S. National Science Foundation grants DEB-1146409, DEB-
1754667, and DEB-1754697, and the Department of Science and Technology (DOST), Philippines.
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