Phytotaxa 433 (1): 055–066 https://www.mapress.com/j/pt/ Copyright © 2020 Magnolia Press

A revision of Vaccinium bullatum (Ericaceae): floral morphology, distribution and typification

MAXIM S. NURALIEV1,2,*, PETER W. FRITSCH3, ANTON S. BEER2, YIHUA TONG4, LEONID V. AVERYANOV5,

ANDREY N. KUZNETSOV1,6 & SVETLANA P. KUZNETSOVA1

1Joint Russian-Vietnamese Tropical Scientific and Technological Center, Cau Giay, Hanoi, Vietnam. e-mail: max.nuraliev@gmail.com 2Department of Higher Plants, Biological Faculty, M.V. Lomonosov Moscow State University, 1, 12, Leninskie Gory, 119234 Moscow, Russia.
3Botanical Research Institute of Texas, 1700 University Drive, Fort Worth, Texas 76107, U.S.A.

4Key Laboratory of Plant Resources Conservation and Sustainable Utilization & Guangdong Provincial Key Laboratory of Digital Botanical Garden, South China Botanical Garden, Chinese Academy of Sciences, Guangzhou 510650, P. R. China.
5Komarov Botanical Institute of the Russian Academy of Sciences, 2, Prof. Popov str., 197376 St. Petersburg, Russia.
6A.N. Severtsov Institute of Ecology and Evolution of the Russian Academy of Sciences, 33, Leninsky av., 119071 Moscow, Russia. *Author for correspondence.

Abstract

A comprehensive morphological and taxonomic revision of Vaccinium bullatum, a species of Ericaceae from China and Vietnam, is presented. Typification of this species is performed and a specimen at P (P04022860) is designated as a second- step lectotype. The floral morphology of V. bullatum is described for the first time, based on the study of living plants in their natural habitat and dry collections. Sectional placement of the species is discussed in the light of new morphological data; its attribution to V. sect. Galeopetalum is partly corroborated. A full list of all available specimens examined, a distribution map, a drawing and colour photographs are included for the species.

Keywords: biodiversity, China, flora, Guangxi, lectotypification, Northern Vietnam, sections, taxonomy, Vaccinium, Xuan Son National Park

Introduction

Vaccinium bullatum (Dop 1930: 700) Sleumer (1941: 446) is an insufficiently known species. It is reported to occupy a narrow area of distribution in western Guangxi Province, China and in several northern provinces of Vietnam (Ho 2000, Hiep 2003, Fang et al. 2005). The available information on the morphology of this species was considerably incomplete up to now, as the accounts contained only descriptions of leafy shoots and infructescences and the flowers were stated to be unknown (e.g., Fang et al. 2005). Other features, such as the morphology of the plant base and the number of ovary locules, were also uncertain. Consequently, the sectional placement of V. bullatum within Vaccinium Linnaeus (1753: 349) remained highly questionable (e.g., Vander Kloet & Dickinson 2005 vs. Vander Kloet & Dickinson 2009).

Here, we fill a number of gaps in the knowledge of V. bullatum. We provide the first description and illustrations of its flowers based on a recent collection from Vietnam with ample inflorescences made by the first author and supplemented by several other collections. We also update the description of the entire plant, based on all the available material. We discuss the sectional placement of V. bullatum in light of the newly obtained morphological data. Critical study of herbarium collections allowed amending the identification of the specimens earlier assigned to V. bullatum, which is reflected in the identification list provided here. Then, we review the known geographical distribution of this species and summarize our results on a map. Finally, we provide a second-step lectotypification of V. bullatum according to our finding that it is currently typified by two specimens (herbarium sheets).

Material & methods

The specimen Nuraliev M.S. 1071 identified as Vaccinium bullatum was collected during an expedition of the Russian- Vietnamese Tropical Centre in Xuan Son National Park, Phu Tho Province, Vietnam, in June 2014. Its duplicates are deposited at BRIT and MW (Seregin 2018). In addition, the herbarium collections of V. bullatum and several similar species of Vaccinium kept in A, ACAD, GXMG, GXMI, HITBC, HN, IBK, IBSC, L, LBG, LE, MO, P and PE were examined. Detailed information on herbarium collections of V. bullatum and two similar species examined for this study are provided in the Appendix. The photographs of living plants were taken in Xuan Son National Park with a Pentax Optio W80 digital camera. The description and drawings presented here are based on herbarium collections and photographs of living plants. Measurements of the specimens of V. bullatum were performed on dried plants with a ruler, and also under a Carl Zeiss Stemi 2000 dissecting microscope equipped with a digital camera calibrated with an object micrometer to document small plant parts. The description of the flower is mostly based on the specimens Nuraliev M.S. 1071 and Averyanov L.V., Hiep N.T. VH 4961, partly rehydrated in heated water.

Terminology of stamen structure generally follows Hermann & Palser (2000) and Stevens et al. (2004). Because the anthers of most Ericaceae invert in the course of stamen development, their inner side (i.e., that facing the flower center) is morphologically abaxial, and the outer side is adaxial. Hermann & Palser (2000) used these terms in an opposite way (i.e., as if the anthers are not inverted). To avoid confusion, we prefer to use the terms “ventral” (inner) and “dorsal” (outer) for the sides of the mature anthers (see also Bukhari et al. 2017). Similarly, we disregard anther inversion, and describe the distal (apical) part of the anther as seen in the mature condition. Then, we follow the interpretation of the calyx tube and the inferior ovary that is accepted for angiosperms in general (Leins & Erbar 2010, Ronse De Craene 2010). According to this interpretation, the calyx is arranged at the top of the inferior ovary and consists of a tube and/or free lobes. We found the alternative way used in some treatments of the genus Vaccinium and other Ericaceae (e.g., Fang et al. 2005, Watthana 2015), i.e., description of the calyx tube as being adnate to the ovary and the calyx limb as a part of the calyx free from the ovary, confusing and morphologically incorrect. Finally, the swollen transitional part between the base of the tap root and stem that is often present in Vaccinium and related genera, especially in their epiphytic representatives, has been described in various ways, including “tuber”, “lignotuber”, “tuberous root” and “root swelling” (Sleumer 1967, Vander Kloet 2002, Stevens et al. 2004, Fang et al. 2005, Watthana 2015). Because the morphological nature of this structure remains unclear, we prefer to follow Stevens (1969) and use the noncommittal term “swollen stem base or root”.

Taxonomic treatment

Vaccinium bullatum (Dop) Sleumer (1941: 446) (Fig. 1–3).

Literature:—Fang (1991: 101, Fig. 26 (2, 3)), Hiep & Ho (1996: 92, as “Vaccinium bulbatum”), Ho (2000: 617), Hiep (2003: 447), Fang et al. (2005: 486, Fig. 672 (2, 3)), Tang (2011: 651).

Basionym:—Agapetes bullata Dop (1930: 700).

TYPE:—VIETNAM. Tonkin [Hoa Binh]: [Da Bac district, Doan Ket, Phu Canh Mt.] massif de Nui bien, près Chobo, 1198 m, 1 September 1926, Poilane E. 13090 (lectotype: P, first-step lectotype designated by Sleumer 1941: 446, second-step lectotype designated here: P: P04022860!; isolectotypes: A: 00015986!; L: L0007945!; P: P04022861!). Images of lectotype and isolectotypes available at:

http://mediaphoto.mnhn.fr/media/151574415835835NFsQ4JfZZ2ve7v, http://mediaphoto.mnhn.fr/media/1515744161197PKkPh96Cq0sg0Fs7, https://s3.amazonaws.com/huhwebimages/C10081C41588474/type/full/15986.jpg, http://medialib.naturalis.nl/file/id/L0007945_MLN/format/master

Description

Shrubs, epiphytic or epilithic, evergreen, to 2 m tall, sparsely branched, with fusiform swollen stem bases or roots. Branchlets, petioles, leaf blades, inflorescence rachises, bracts, pedicels, bracteoles, calyx, corolla, ovary, and fruit with white erect or ascending straight or more or less curved trichomes to 0.3(to 0.5) mm long mixed with more scattered erect to appressed ferrugineous (darkening to black on old branchlets and leaf blades) stalked glandular trichomes to 0.10(–0.15) mm long, often glabrescent. Mature branchlets reddish brown, more or less terete but irregularly ridged, 3–4 mm thick, densely lenticellate (Fig. 2a–c); epidermis grayish, peeling around lenticels; perennating buds with multiple overlapping scales. Leaves alternate, scattered, distichously arranged (Fig. 2a–c, 3c); petiole 1–3 mm long, 2.3–2.8 mm in diam.; leaf blade ovate or narrowly ovate, or rarely narrowly oblong-ovate or lanceolate, larger leaves on each branch 9–16 × 3.9–8.1 cm, 2.1–3.3 times as long as wide, coriaceous, green in vivo and dark green or brown in sicco adaxially, pale green in vivo and brown in sicco abaxially, both sides slightly shiny, adaxial surface bullate, i.e., with convex sections flanked by sulcate major veins; adaxial surface glabrous except puberulous at base along major veins; abaxial surface with white trichomes mainly on major veins and glandular trichomes mainly on surface proper, with both types of trichomes when caducous leaving a faintly punctate scar; base subrounded, rounded, subtruncate, or subcordate; margin entire, slightly revolute, with one sessile dark basal gland 1.0–1.2 mm in diam. per side that often forms a shallow lobe at margin (Fig. 2b); apex acute to caudate-acuminate, or rarely obtuse; midvein sulcate adaxially but with a strong central ridge, strongly raised abaxially; secondary veins pinnately arranged, sulcate adaxially, raised abaxially, 7 to 12 on each side of midvein or to 20 when including smaller veins that do not reach the margin, more or less arc-ascending, often meeting in a closed loop before reaching margin, often some veins straighter, smaller, and not reaching margin situated between larger arching veins; tertiary veins slightly sulcate adaxially, raised abaxially.

Inflorescences axillary and pseudoterminal, indeterminate (racemose and developing mostly beyond the confines of the perennating buds), 1(to 3) per axil, 1.5–9.0 cm long at anthesis, 8- to 14-flowered (Fig. 2b, 3e); peduncle and rachis green flushed red in vivo; bracts caducous, maroon to black in vivo, narrowly deltoid, cucullate, 1.6–2.5 × 0.6–1.0 mm, chartaceous, adaxially glabrous, abaxially pubescent, margin entire, apex acute. Flowers articulated with pedicel, 5-merous, 5.5–7.5 mm long (Fig. 1a, 2d–g). Pedicels green flushed red in vivo, 4–7(–9 in fruit) mm long and 0.65–0.75(–0.80) mm thick at anthesis, slightly expanded to 1.00(–1.15) mm towards apex; bracteoles 2, borne at base of pedicel (or slightly above), opposite, caducous, maroon to black in vivo, narrowly deltoid, cucullate, 1.4–1.7 × 0.4–0.6 mm, chartaceous, margin entire, apex sharply acute. Calyx (without ovary) green flushed red in vivo, erect, cupuliform, 1.8–2.2 mm long (Fig. 2f,i); connate portion (tube) 0.25–0.30 mm long; lobes 5, slightly incurved (appressed to corolla), narrowly ovate-deltoid to oblong-deltoid, 1.3–2.4 × 0.8–1.1 mm, pubescent on both sides but pubescence less dense than on ovary and tube, margin entire, smooth (e.g., without sessile glands), apex acute, without terminal gland. Corolla in bud closed and strongly 5-ribbed in line with petal midveins (Fig. 2b); gamopetalous for ca. 2/3 length, at anthesis whitish except bright red along petal midveins, broadly campanulate, ca. 5–6 mm long, 4.0–5.5 mm in diam., outside with glandular trichomes mainly within ca. 0.5 mm on each side of petal midvein and more sparsely white-puberulent closer to midvein, inside glabrous; lobes 5, reflexed in vivo, erect in sicco, deltoid or narrowly deltoid, ca. 2.0 × 2.2 mm, sometimes with several long setae adaxially near apex (Fig. 1a, 2d–g). Stamens 10, dimorphic with respect to pores and spurs, free from corolla, distinct from each other but slightly conjoined by lateral sides of thecae, ca. 4.5–5.0 mm long (Fig. 1, 2d,e,g,h); filaments bent inwards, ca. 1.5–2.3 mm long, margins sparsely white-villous in sicco (white-lanate in vivo) with trichomes to 0.35 mm long (Fig. 1); anthers dorsifixed with filament attached at distal part of theca, ca. 3.6–4.5 mm long, introrse, with 2 spurs each (Fig. 2g,h); anther thecae ca. 1 mm long, papillate; anther tubules parallel, ca. 3.5 mm long, opening by strongly oblique ventral apical pores 0.90–1.19 mm long on antesepalous anthers and 1.20–1.35 mm long on antepetalous anthers, pore apex acute (Fig. 1); anther spurs borne dorsally at base of tubules, strongly curved, ca. 0.8–1.1 mm long, those on antesepalous stamens extending laterally outside of antepetalous anthers and strongly overlapping with spurs of next antesepalous stamens, those on antepetalous anthers strongly hooked outward below spurs of antesepalous stamens (Fig. 1). Antesepalous anthers slightly closer to flower center than antepetalous anthers (Fig. 1b). Ovary inferior, dark green in vivo, slightly obconic (Fig. 1a, 2f,g), 0.8–1.7 mm long, 1.7–2.0 mm in diam., terete in vivo, slightly 5-ribbed in bud in sicco, 5- locular with incomplete false septa protruding to not more than 1/2 of each locule (Fig. 3a,b); disk annular, occupying the entire area within the corolla tube, 1.45–1.90 mm in diam., glabrous or pubescent (Fig. 2g,i); style slightly exserted from corolla (Fig. 2d–h), ca. 4.7–5.0 mm long, glabrous or basally pubescent. Infructescences 4.7–13.4 cm long. Fruit (berry) reportedly white, globose, 6.0–7.5 × 6.5–7.5 mm excluding calyx, fleshy, pubescent, pseudo-10-locular, bearing persistent calyx with erect-incurved lobes 1.5–3.5 mm long (Fig. 3d).

Nomenclatural notes:—The protologue of Agapetes bullata (Dop 1930) includes a description of the species together with the type locality as “Tonkin: massif de Nui-bien, près Cho-bo” and a collector name as “Poilane”. In publishing the combination under the genus Vaccinium, Sleumer (1941) indicated the collection number of the type specimen as well as the herbarium of its deposition: “Poilane n. 13090, Typus in Herb. Mus. Paris”, which appears to fulfill the conditions of lectotypification (Art. 7.11 of ICN, Turland et al. 2018; see also McNeill 2014). However, we have found that there are two specimens of the collection Poilane E. 13090 deposited in P (P04022860 and P04022861) and thus Sleumer’s choice (1941) was the first-step for lectotypification (Art. 9.17). We have selected the specimen at P with barcode P04022860 as the lectotype in the second-step because it has the original label, unlike the other one. The specimens at A (00015986) and L (L0007945) were distributed from P, which is evident from the notes that they contain. These two specimens as well as P04022861 at P are isolectotypes.

Morphological notes:—1. Although the original label of the lectotype states “fleurs blanchatres”, the flowers are in fact absent from the four type specimens that we have studied. This is consistent with the treatment in the Flora of China (Fang et al. 2005) which states “Flowers unknown”. Among the specimens that we were able to locate, several contain flowers, i.e., Averyanov L.V. et al. HAL 12808, Averyanov L.V. et al. LX-VN2334, Averyanov L.V., Hiep N.T. VH 4961, Chinese Botany United Guangxi Team 1698, Huang Y.S., Nong D.X. Y0684, Nuraliev M.S. 1071, Phuong 2043, Qin H.N. et al. 603033, Xu W.B. et al. 9608, most of them made after the publication of Flora of China (Fang et al. 2005). As far as we are aware, the flowers of this species have not been described nor illustrated to date, and thus this is accomplished in our work for the first time.

2. Uncertainty exists regarding the distinction between 5-locular, 10-locular and pseudo-10-locular ovaries in several taxa of Vaccinium. It is most likely that gynoecia of 10 carpels, i.e., the truly 10-locular ones, are not found in Vaccinium s.str. (excluding Rigiolepis Hooker f. 1873: 54, t. 1160) (Stevens et al. 2004, Fang et al. 2005, Vander Kloet 2005, Argent 2019), and the indications of 10-locular ovaries in Vaccinium (e.g., Vander Kloet 2005, Vander Kloet & Dickinson 2009), including V. bullatum (Vander Kloet & Dickinson 2005), are errors likely from inaccurate observations. The absence or presence of false septa, i.e., a 5-locular vs. pseudo-10-locular ovary, has been widely used in the taxonomy of Vaccinium (see below). However, these conditions appear not to be perfectly discrete, as follows from our investigation. Indeed, in V. bullatum, the anthetic false septa are considerably incomplete, and the ovary can be described as 5-locular with V-shaped locules (Fig. 3a,b). The pseudo-10-locular condition that is usually assigned to this species (Fang et al. 2005, Vander Kloet & Dickinson 2009) apparently becomes more evident in fruits. Therefore, this character depends on the stage of the flower/fruit under study, and we thus argue that examination of a range of phenological stages is needed for its reliable evaluation.

Ecology and phenology:—Vaccinium bullatum grows in mountainous areas and limestone hills, in broad-leaved forests, at 600–2000 m a.s.l. Flowering: (February)May–June; fruiting: (July)August–March.

Distribution:—Vaccinium bullatum is distributed in Guangxi Zhuang Autonomous Region, China and the Vietnam provinces of Lao Cai, Cao Bang, Tuyen Quang, Vinh Phuc, Phu Tho and Hoa Binh as well as in the Tam Dao massif (without precise location).
The distribution of V. bullatum is presented in Fig. 4, based exclusively on the specimens examined in our study (see Appendix).

Vernacular names:—Chinese: 泡泡叶越橘; Vietnamese: Sõn trâm phồng; Việt quất lá bọt.

Sectional placement:—There is some uncertainty as to the sectional placement of Vaccinium bullatum. Sleumer (1941) placed the species in V. sect. Galeopetalum J.J.Sm. in Boerlage (1912: 101), characterized by this author as having large evenly scattered evergreen leaves with multiple secondary veins and entire margins, axillary indeterminate racemose inflorescences that develop mostly beyond the confines of the perennating bud, campanulate corolla divided to ca. the middle, two prominent spurs per anther, long-erect anther tubules, and pseudo-10-locular ovaries. Sleumer’s decision was adopted by Fang (1991). Vander Kloet & Dickinson (2005) separated some of the species in this section into the new section V. sect. Calcicolus Kloet in Vander Kloet & Dickinson (2005: 451) on the observation that the species in the latter section actually have determinate inflorescences and these develop entirely within the confines of an enlarging perennating bud, unlike the species remaining in V. sect. Galeopetalum. This separation is supported by phylogenetic analysis based on DNA sequence data (Kron et al. 2002 and unpubl. data, in Vander Kloet & Dickinson 2005), although species sampling within the two sections was limited. Moreover, although not noted by Vander Kloet & Dickinson (2005, 2009), the species of V. sect. Calcicolus lack swollen stem bases or roots (Y.-H. Tong, pers. obs.), whereas those of V. sect. Galeopetalum possess them (Sleumer 1967; Vander Kloet & Dickinson 2005).

Vander Kloet & Dickinson (2005) excluded Vaccinium bullatum from V. sect. Calcicolus perhaps based on its indeterminate inflorescence, and this exclusion is supported by the presence of swollen stem bases or roots in the species (Y.-H. Tong, pers. obs.). They indicated several additional features of this species distinctive from those of V. sect. Calcicolus, namely, a 10-locular ovary and seeds with thin white testa surrounding a green embryo (drying black). However, our study clearly shows that V. bullatum possesses a pseudo-10-locular ovary rather than a 10-locular ovary; this feature thus appears to be consistent with both V. sect. Calcicolus and V. sect. Galeopetalum s.str. Vander Kloet & Dickinson (2009) subsequently referred V. bullatum to V. sect. Euepigynium Schlechter (1919: 174). The basis for this decision, however, is not clear because the members of this section have palmate leaf venation whereas V. bullatum has pinnate venation. The retention of V. bullatum in V. sect. Galeopetalum (excluding the species of V. sect. Calcicolus) appears to be justified in our investigation on the basis of the above characters as well as its slightly conjoined anthers, which also are possessed by some species of this section (Vander Kloet 2002). However, V. bullatum appears to key to couplet 28 in the key to Vaccinium sections in Vander Kloet & Dickinson (2009) (which unites sections Bracteata Nakai (1927: 234), Eococcus Sleumer (1941: 420) and Euepigynium) and both leads in this couplet appear to be false for V. bullatum because of leaves with entire margin and pinnate venation in the species. Moreover, within the key, V. bullatum does not fit V. sect. Galeopetalum because this species possesses a long inflorescence rachis; notably, the assumed character of a short rachis in V. sect. Galeopetalum is in fact not mentioned in any other source. Thus, similar to Stevens (1969), we assert that the species of this section need more detailed morphological documentation. In this regard, our expanded description of V. bullatum can serve as a model for such documentation to make further progress on the delimitation of these and related sections in Vaccinium.

Acknowledgements

We thank Bui Hong Quang, Weibin Xu and James C. Solomon for their help in our examination of herbarium specimens, including providing images of specimens in HN, IBK and MO; Florian Jabbour for finding and imaging the type specimens in P; Gary Li for translating the Chinese herbarium labels and help with the geography of China; Phan Ke Loc and Do Van Truong for help in determining the modern geographical location of the type specimen; and Eugene S. Popov for providing the map of Southeast Asia. The work of MSN and ASB was supported by the Russian Foundation for Basic Research (project 19-54-54007) and conducted in accordance with Government order for the Lomonosov Moscow State University (projects AAAA-A16-116021660105-3 and АААА-A16-116021660045-2, respectively). The work of YHT was supported by the National Natural Science Foundation of China (grant no. 31870180). The work of LVA was financially supported in part by the Vietnam Academy of Science and Technology, international cooperation mission QTRU01.07/18-19 and the Russian Foundation for Basic Research (project 18-54-54005) and was carried out in the framework of the institutional research project of the Komarov Botanical Institute of the Russian Academy of Sciences (project АААА-А18-118031290070-6).

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