Plant Syst Evol DOI 10.1007/s00606-012-0725-2

ORIGINAL ARTICLE

A new of (Ruscaceae s.l., ) from southern , field observations on its flowering and possible by flies (Phoridae)

Nikolay A. Vislobokov • Andrey N. Kuznetsov • Dmitry D. Sokoloff

Received: 23 June 2012 / Accepted: 16 October 2012 Ó Springer-Verlag Wien 2012

Abstract The genus Aspidistra with almost 100 species Introduction is the most diverse in southern and northern Viet- nam. We describe a new species Aspidistra phanluongii The genus Aspidistra, variously placed in Convallariaceae, N.Vislobokov from southern Vietnam. Species of Aspi- Ruscaceae or s.l. (Conran and Tamura 1998; distra are herbs characteristic of Southeast Asian forests. Rudall et al. 2000; Chase et al. 2009; Takhtajan 2009; Kim Flowering of these can be considered cryptic. There et al. 2010), is remarkable in the rapid discovery of new are no recorded field observations on phenology, biology of species during recent decades. Conran and Tamura (1998) flowering or pollination of Aspidistra. Previously proposed reported ca. 11 species of Aspidistra from Himalayas, pollinators of Aspidistra include collembolans, amphipods, Indochina, southern China and south-western Japan. Ten fungus gnats (Diptera: Mycetophilidae, Sciaridae) and even years later, Tillich (2008) recognised 93 species of Aspi- slugs. We present observation data on the flowering of distra, and following his review several new species have A. phanluongii in the wild. Flowers were visited by flies of been described, mainly from southern China (e.g. Lin et al. (Phoridae) and ants. Megaselia flies, well- 2009; Xu et al. 2010; Gao and Liu 2011; He et al. 2011; known pollinators of numerous tropical species, are Lin et al. 2011). Vietnam is the second hotspot of new shown to be likely pollinators of Aspidistra phanluongii. species discovery in Aspidistra after China (Liang and The present study does not provide evidence of pollination Tamura 2000; Bogner and Arnautov 2004; Bra¨uchler and by fungus gnats. Ngoc 2005; Tillich 2005; Tillich et al. 2007; Tillich and Averyanov 2008; Averyanov and Tillich 2012; Averyanov Keywords Aspidistra Flies Flower et al. 2012). Southern China and an adjacent part of biology Pollination Phoridae Southern Vietnam northern Vietnam is the centre of total species diversity of Aspidistra. Some species inhabit the southern part of Electronic supplementary material The online version of this Indochina. Aspidistra larutensis De Wilde and Vogel article (doi:10.1007/s00606-012-0725-2) contains supplementary (2006), the southernmost species of Aspidistra, was material, which is available to authorized users. described from Peninsular Malaysia, and placed in synon- ymy under A. longifolia Hook.f., which is also found in N. A. Vislobokov (&) D. D. Sokoloff Department of Higher Plants, Faculty of Biology, Thailand and (Phonsena and De Wilde 2010). We M.V. Lomonosov Moscow State University, describe a new species of Aspidistra from Dong Nai 1, 12, Leninskie Gory, 119234 Moscow, Russia Reserve in southern Vietnam. This is the southernmost e-mail: [email protected] known locality of Aspidistra in Vietnam, and one of N. A. Vislobokov A. N. Kuznetsov D. D. Sokoloff southernmost for the genus. The new species was collected Joint Russian-Vietnamese Tropical Scientific and Technological in one of the few surviving lowland rainforests of southern Centre, Cau Giay, Hanoi, Vietnam Vietnam (Kuznetsov 2003) at elevation of 30–90 m, whereas many other members of the genus have been A. N. Kuznetsov A.N. Severtsov Institute of Ecology and Evolution of Russian collected in montane forests between 300 and 800 m Academy of Sciences, Leninsky 33, 119071 Moscow, Russia elevation. 123 N. A. Vislobokov et al.

Species of Aspidistra are commonly present but incon- flowers are held horizontally (and possibly somewhat spicuous herbs in forests of Southeast Asia. Flowers and downwards) lying on the substrate and are likely to be fruits are situated close to ground level and often covered visited by small ground insects. No floral scent was by leaf litter. This is why flowering of Aspidistra can be recognised in A. renatae. According to Bra¨uchler and Ngoc considered cryptic (Phonsena and De Wilde 2010). The (2005), the bright is in high contrast to the dark flowers are nevertheless extraordinarily diverse, and mor- surrounding tissue and could serve as a first attraction point phological characters of the flower are the most important favouring pollination by insects. A ring of non-pigmented in species delimitation (Liang and Tamura 2000; Tillich tissue around the base of the gynoecium may serve as a 2005, 2008). Flower groundplan variation in Aspidistra is light trap, guiding flower-visiting insects towards the remarkable among Asparagales (Remizowa et al. 2010). It proximate part of the flower, forcing them to pass the is expected that the flower structure diversity in Aspidistra anthers. Some floral morphology characters suggest polli- reflects pollination biology. The cryptic nature and pro- nation of Aspidistra species by fungus gnats (Diptera: nounced seasonality of flowering make studies of repro- Mycetophilidae, Sciaridae; Vogel 1978; Tillich 2005). As ductive biology problematic. Many recently described pointed out by Tillich (2005), the pistil in many species species of Aspidistra were discovered in the wild as sterile mimics a fungal fruiting body and the flowers are often dull plants, and only subsequent cultivation in greenhouses in colour, mostly scentless, and develop close to the soil enabled the study of flowers and taxonomic placement (e.g. surface. Bogner and Arnautov (2004) noted that the unu- Bogner and Arnautov 2004; Tillich 2005; De Wilde and sual morphology of A. locii Arnautov & Bogner flower Vogel 2006; Averyanov and Tillich 2012). Since the plants with only a small opening suggests pollination by small appear to be self-incompatible, they are propagated vege- insects, perhaps flies. They also noted high superficial tatively and do not set fruit in greenhouse cultivation (e.g. similarity between the flowers of A. locii and the special- Bogner and Arnautov 2004; Bogner 2005; Tillich 2005). ized inflorescences (syconia) of Ficus, suggesting that Fruit and seed morphology is so far unknown for the flower structure of A. locii must be significant in its pol- majority of the species described from Vietnam (Bogner lination biology. Despite the high diversity of hypotheses and Arnautov 2004; Bra¨uchler and Ngoc 2005; Tillich on pollination biology of Aspidistra, observations of floral 2005; Tillich et al. 2007; Averyanov and Tillich 2012). visitors in the wild are not known. We present work Previous field studies in Vietnam recording flowering of towards filling this gap by field investigation of floral Aspidistra species in the wild did not report any pollinator biology of the here newly described Aspidistra phanluongii visits (Bra¨uchler and Ngoc 2005; Tillich et al. 2007; from southern Vietnam. Tillich and Averyanov 2008). Published data suggest that pollination biology of Aspidistra could be highly diverse. The unusual flowers Materials and methods suggest biotic pollination because pollen is hidden under the stigma and a flower looks like an insect trap. Wilson Plants were observed and collected by the first author in (1889) suggested pollination of the Japanese species southern Vietnam (Dong Nai Reserve) during November– A. elatior Blume by slugs, a hypothesis subsequently December 2011. Sporadic observations and collections adopted in various accounts as a rare example of angio- made in the same locality in other seasons (March, May sperm flowers pollinated by mollusks (e.g. Richards 1986; and October 2011) are also considered. In the description Bell and Hemsley 2000). Kato (1995) performed laboratory of the new species the mean value is given in square experiments (based on flowers collected in the field) brackets after the range of variation of quantitative char- showing the pollen being eaten by the terrestrial acters. These data are based on 50–60 measurements of Platorchestia japonica Tattersall, 1922 (), a leaf characters, ca. 25 measurements of perianth lobe potential pollinator of A. elatior. This hypothesis once characters and 4–8 measurements for other floral charac- again placed Aspidistra among angiosperms with the most ters. Observations on floral visitors were performed unusual pollination ecology. Conran and Bradbury (2007), between 09:00 (occasionally 08:30) and 17:00 during two however, documented successful fruit set in A. elatior periods: 16–22 November 2011 and 9–22 December 2011. cultivated in Australia in conditions where amphipod spe- For observations of flowering stages, flowers and flower cies mentioned by Kato (1995) were absent. They con- buds were labelled individually. Sixty-two flowers were cluded that pollinator substitution is possible in Aspidistra monitored in total. Of these, 24 flowers were observed and discussed soil invertebrates including collembolans from flowering to fruiting. Visual observations as well as and others as possible pollinators. Bra¨uchler and Ngoc video recording and photography were used. Insects were (2005) performed field observations on the flowering of captured by air stream (exhauster), killed by ethyl acetate A. renatae Bra¨uchler in central Vietnam and noted that the fumes and fixed in 70 % ethanol. The number and time of 123 A new species of Aspidistra (Ruscaceae s.l., Asparagales) from southern Vietnam

Table 1 Floral visitors to Aspidistra phanluongii Observation Total of observation Number of observed/ Mean observed flies Number of observed ants Mean ants per min time interval (min) for all days captured flies per min of observations of observations

9:00–11:00 346 10/6 0.029 4 0.012 11:00–13:00 312 9/2 0.029 17 0.054 13:00–15:00 174 19/9 0.11 9 0.052 15:00–17:00 194 19/7 0.098 2 0.01 insect visits were noted in the field journal; these data are Morphological description summarized in Table 1. For floatability testing, 22 fruits were used. For scanning electron microscopy (SEM), the Herbaceous perennial rhizomatous evergreen plant. Rhi- material was dissected in 70 % ethanol and subsequently zome creeping, hypogeous, Ø 4.5–8 [6.4] mm, cataphylls transferred to 100 % acetone via 80 and 96 % ethanol and pale green up to 10 cm long, roots numerous, Ø up to 4 mm. a mixture of ethanol (96 %) and acetone (100 %) in pro- Leaves solitary, distinctly divided into and lamina. portion of 1:1. The material was then critical-point dried in Petiole 10–40 [20] cm long, Ø 2–4 [3] mm. Lamina fair a HITACHI HCP-2 critical point drier using liquid carbon green, lanceolate to elliptic, basally cuneate and distally dioxide. Dried samples were mounted onto stubs using acuminate, 17–31 [23.5] cm long, 3–9 [6.5] cm wide, with double-sided sticky tape or nail polish, coated with plati- 3–4 secondary veins at both sides of the midrib. Peduncle num and palladium using an Eiko IB-3 ion coater and hypogeous, white or pale green, 1–5 [2.3] cm long, Ø 1,8–3 observed using a Cam Scan S-2 scanning electron micro- [2.3] mm, with 5–8 scale leaves, two upper scales largest, scope at Moscow University. adjoining to perianth, widely ovate, acuminate, up to 11 mm long (Fig. 1a, b). Flowers crowded, epigeous, perianth 10.5–16.5 [13.6] mm high, Ø 10–16 [13] mm, tube widely Results bowl-shaped, dark-violet inside and outside, mouth of tube hole Ø 4–9 [6] mm; lobes 6 (very rarely 4, 5, 7, 8) (Fig. 1a, d, e), Aspidistra phanluongii N.Vislobokov sp. nov. reflexed, oblong, dark-violet or dull-green outside, white inside (pale green inside in young flowers), 5.0–7.5 [6.1] Holotype: Vietnam, Prov. Dong Nai, Vinh Cuu District, mm long, 3–4.5 [3.9] mm wide, lobes basally with two 50 km NNE from Bien Hoa, Dong Nai Reserve, Ma Da prominent keels 0.5 mm high. Stamens usually 6, at the Forest District, 11°22043.700N, 107°03041.100E, 18 Nov base of the perianth tube, filament 0.5–0.8 [0.6] mm long, 2011, N.A. Vislobokov M0111 (MW), including all parts in anthers 1.5–2 [1.6] mm long, anthers latrorse (Fig. 1c). liquid collection at Moscow University. Pistil table-shaped (Fig. 1b), ovary inconspicuous Paratypes (all in the same locality): 19 Nov 2011, N.A. (Fig. 2a), superior, tricarpellate, in each carpel 2 seed buds Vislobokov M0211 (MW), including all parts in liquid (Fig. 2b), style short, 2–2.5 [2.3] mm, stigma circular, disc- collection at Moscow University; 4 Mar 2011, A.N. Kuz- shaped, three-lobed, each lobe distally bifurcate, upper sur- netsov, M.S. Nuraliev, D.D. Sokoloff 396 and 397 (MW), face white in the centre, distally violet, sometime violet including vegetative parts and fruits in liquid collection at mottled, Ø 10–13 [12] mm. Pollen grains globular, inaper- Moscow University; 7 June 2011, A.B. Vassilyeva s.n., turate, Ø 30–40 lm (Fig. 2g), exine irregularly reticulate, the vegetative parts and fruits in liquid collection at Moscow cross-points of the net strongly thickened (Fig. 2h). Fruits University. usually 3-seeded (rarely 4–6-seeded) (Fig. 1f), dark-violet to Affinity: Aspidistra phanluongii is similar and probably black, spherical, slightly oblate in vertical direction, 9–15 closely related to A. opaca H.-J. Tillich, because both have [12] mm long, Ø 10–22 [14] mm, somewhat three-lobed, a widely urceolate six-lobed perianth and a table-shaped with rough protuberances. The shape of the seeds is like a pistil with short style and circular stigma. The new species segment of a sphere, surface somewhat wrinkled (Fig. 2e). differs from A. opaca by lobes of perianth oblong (not deltoid), with upper surface white (not dull purple), anthers Etymology 1.5–2 mm (not 2–2.5 mm) long, stigma with white and violet patches (not white), perianth 10–16 mm (not ca. The species is named in honour of Dr. Phan Luong 20–25 mm) in diam. Aspidistra opaca was described from (1948–2008), a graduate of the Kharkov State University. Vietnam, Prov. Khanh Hoa, Khan Nam (Tillich 2005), Since 1990 Dr. Phan Luong has headed the terrestrial which is approximately 200 km NE from Ma Da. ecology investigation in the Joint Russian–Vietnamese

123 N. A. Vislobokov et al.

Fig. 1 Diversity of flowers and fruits of Aspidistra phanluongii. a Common type of flower structure with six lobes of perianth. b Longitudinal section of typical flower. c Cross- section of typical flower (stigma and lobes removed). d Uncommon type of flower structure with five lobes of perianth. e Uncommon type of flower structure with eight lobes of perianth. f Cross section of fruit with three seeds and three abortive ovules. lb lobes of perianth, p peduncle, pr perianth, s seed, sg stigma, st stamen, white arrow abortive ovule, black arrow stamen in substigmal area

Tropical Research and Technological Centre (JRVTRTC); (Meliaceae) and Lagerstroemia calyculata Kurz (Lythra- he has been the organizer of and a participant in all ceae). The forest is situated on yellow ferrallitic soil, drying expeditions of the Tropical Centre since 1995. out between the wet seasons. Despite an extensive search, only one compact population of Aspidistra phanluongii,about Distribution and ecology 600 9 600 m2, has been found. Within the population, plants are abundant. No clear habitat specificity has been observed. The species is known only from the type locality, a lowland closed-canopy rainforest. The upper part of the canopy is Flower and fruit biology dominated by Dipterocarpus dyeri Pierre, Shorea roxburghii G. Don. (Dipterocarpaceae), Irvingia malayana Oliv. ex Flowering was recorded from October to December. We Benn. (Irvingiaceae), Sandoricum koetjape (Bum. f.) Merr. made no field observations in the period from June to

123 A new species of Aspidistra (Ruscaceae s.l., Asparagales) from southern Vietnam

Fig. 2 Reproduction biology of Aspidistra phanluongii. a Longitudinal section of flower bud (scale bar 1 mm). b Tricarpellate ovary with six ovules (ovary wall partly removed) (scale bar 300 lm). c Megaselia fly in side view (scale bar 1 mm). d Head of Megaselia fly in front view (scale bar 100 lm). e Surface of seed (scale bar 20 lm). f Germinated pollen grain on the papillate upper surface of stigma (scale bar 30 lm). g Pollen grain (scale bar 10 lm). h Surface of pollen grain (scale bar 1 lm). ov ovary wall. p peduncle, pr perianth, pt pollen tube, sb seed bud, sg stigma, st stamen, white arrow ovary

September, but the absence of fruits and old flowers in become violet (Fig. 3a). The violet bud (Ø 10–15.5 October strongly suggests that anthesis did not take place [12] mm) widens during the subsequent 8–9 days and in September. Flowering is asynchronous in the population. ultimately opens. Green margins of perianth lobes become While first opened flowers were recorded in the beginning visible 1–2 days before flowering. A flower opens gradually of October, new floral buds were recorded as late as 19 during 1 day and possibly also during the night (Fig. 3b). December. By the next morning the flower is fully open. The upper Young green flower buds (Ø 4.5–8 [6] mm) become surface of perianth lobes is pale green in recently open distinguishable among forest litter at the ground level. Buds flowers (Fig. 3c). After 2–3 days of anthesis the perianth are covered with green scales. During the next 7–9 days, lobes become pure white (Fig. 3d). The presence of pollen bud diameter increases, peduncle and scales grow, and thus grains on the stigma and perianth even in incompletely open the bud appears above the forest litter. Gradually, the buds flowers suggests pollinator visits. Germinated pollen grains

123 N. A. Vislobokov et al.

Fig. 3 Flowering stages of Aspidistra phanluongii. a Violet (left) and green (right) flower buds, b opening flower bud, c young flowers, d mature flower, e old flower, f different stages of flowering of the same plant

were detected on the upper surface of the stigma (Fig. 2f). and remain recognisable until at least February. Different According to our observations, insects continue to visit the stages of flowering can be observed in different flowers of flowers during the first 5–7 days of anthesis. No flower the same plant (Fig. 3f). scent or nectar secretion was recorded. Peduncles with mature fruits are generally longer than Evidence of flower senescence appears at the 10th day those at the flowering stage (2–6 [3.9] cm long). Out of 24 of flowering. At first the stigma becomes darkish, and later flowers observed from flowering to post-anthetic stages, 10 the perianth lobes fade (Fig. 3e). During the subsequent (ca. 40 %) produced fruits. By late December, mature fruits 5–6 days the process of senescence spreads to all parts of were recorded in many plants, and new flower buds did not the flower except the fertilized ovary. The peduncle does continue to appear in the population. During February and not elongate significantly during the first days after flow- May 2011, plants with mature and undamaged fruits ering. Peduncles of unfertilized flowers are not abscised attached to peduncles were common, but flowering plants

123 A new species of Aspidistra (Ruscaceae s.l., Asparagales) from southern Vietnam were not recorded (Dr. A.B. Vassilieva, JRVTRTC, Ho Chi Discussion Minh, personal communication). Abscised fruits were not recorded throughout the whole period of our observations. Taxonomic status of Aspidistra from Ma Da The fruits are inconspicuous and situated at soil level close to the plant, and their peduncles are subterranean. Broken Comparative morphology suggests that the Aspidistra peduncles suggesting that fruits were eaten by animals population from Ma Da belongs to a new species, here were not recorded. It seems that fruits form at the begin- described as A. phanluongii. Morphologically it is most ning of the dry season (December) and remain attached to similar to A. opaca, a species recently described from the the plant during several months, possibly until the rainy neighbourhoods of Nha Trang (Tillich 2005). There are season. Because there is no evidence of biotic fruit other species resembling A. phanluongii in some charac- dispersal, we tested a hypothesis of fruit dispersal by ters; For example, A. longifolia has flowers similar to floatation during the rainy season. An experiment was A. phanluongii but strongly differs from it in leaf shape. conducted where 22 fruits were placed in water. Out of the Leaves of A. longifolia are long and narrow, without a 22 fruits examined, 18 did not float and 4 floated. All four clear separation between petiole and lamina. Fruits of floating fruits were damaged by unidentified phytophagous A. longifolia are 5–10-seeded (Phonsena and De Wilde insects. Occurrence of juvenile seed-grown individuals of 2010), while fruits of A. phanluongii are usually 3-seeded. A. phanluongii suggests that reproduction by seed plays an Aspidistra longifolia was described from India (Assam), important role. but Phonsena and De Wilde (2010) recognised it in a broad sense, assigning collections from S. China, Thailand and Pollination biology Peninsular Malaysia to the same species. Analysis of published data on Aspidistra morphology According to our observations, flowers were visited by allows the recognition of a species group with similar flies and ants. Flies were recorded on flowers from 08:39 flower (but not leaf) structure, with the flowers typically to 16:30, and ants from 09:15 to 17:20 (Table 1). All trimerous, the perianth bowl-shaped, the tube usually vio- captured flies were identified as members of the genus let, the stigma wide, the stamens attached to the lower part Megaselia (Phoridae; Fig. 2c, d). There were females and of the perianth (below the stigma), and a southern distri- males of Megaselia among the captured visitors (ca. 50 % bution. We propose that these features of floral morphology females). Ants belonged to genera Pachycodyla and are more conservative and taxonomically important than Leptogenys. the leaf characters, but this is yet to be tested using When a fly visits a flower it lands on a perianth lobe. In molecular phylogenetic data. some cases it lands on the vegetative parts of the plant or The inaperturate pollen grains of A. phanluongii are the forest litter first. Megaselia flies can run quickly and similar to those described for A. locii (Bogner and move into the flower using only their legs. They can also Arnautov 2004) and A. typica Baill. (Rudall and Campbell move from one flower to another by running as well as 1999). It seems that pollen morphology is of limited use in flying. From the perianth lobe surface the fly moves into the taxonomy of the genus. the perianth tube and comes to the stigma surface. It remains there for up to 30 s, moving mainly along the Flower and fruit biology and pollination of Aspidistra perimeter of the stigma. There are gaps 1.2–2 [1.4] mm phanluongii wide between the perianth tube and the stigma. Sometimes a fly lingers on a flower before penetrating into the sub- The present work is the first field study of flower biology stigmal area. The Megaselia’s thorax is ca. 0.6 mm in and floral visitors of Aspidistra in Vietnam. Previous diameter, allowing passage through the gap. Stamens are observations of flowering of Aspidistra in Vietnam were located directly under the gaps between the perianth tube scattered and did not cover substantive time intervals. Our and the stigma, hence the fly becomes covered with pollen data suggest that flowering of A. phanluongii takes place at inevitably and completely. Flies loaded with pollen when the end of the wet season and beginning of the dry season leaving flowers of Aspidistra are recorded. (October to December). The time interval between the The ants visited flowers less frequently than the flies beginning of anthesis and mature fruit set is probably a few (Table 1). An ant attempts to thrust its head through the months. Mature fruits remain attached to the plants at gap, but feels resistance. On rare occasions an ant is seen ground level for a considerable period, most likely until the covered with pollen; it immediately begins to clean its beginning of the new wet season. Neither fruit dehiscence antennae and legs. nor fruit abscission was recorded. Decaying fruits are not Video records of floral visitors are presented in the observed during the dry season, and no obvious vectors of Electronic Supplementary Material. fruit dispersal are recorded. It is most likely that seeds are 123 N. A. Vislobokov et al. somehow dispersed and germinate in situ during the wet A. phanluongii (lowland tropical rainforest) differs from season. Reproduction by seed is successful as juvenile habitats typical to most species of Aspidistra (montane plants were repeatedly observed. Whatever the vector of forest). fruit dispersal, it is unlikely to be effective at range extension, as the observed population occupies only Acknowledgments We are indebted to Tatiana Galinskaya 600 9 600 m2 and other populations were not found (Department of Entomology, Biological Faculty, Moscow State University) for identification of flies visiting Aspidistra flowers, and despite an extensive search. Gennady Dlussky (Department of Biological Evolution, Biological We conclude that A. phanluongii is pollinated by flies Faculty, Moscow State University) for identification of ants. We are belonging to the genus Megaselia (Phoridae). The evidence grateful to Maxim Nuraliev for extensive contribution to field work in is indirect since flower isolation experiments were not February 2011 and helpful discussion, to Anna Vasilieva for col- lecting additional fixed material and field observations and to Alex- performed. We believe the indirect evidence presented here ander Anichkin for generous help and many important suggestions is significant. Megaselia flies were repeatedly observed during field work in November–December 2011. We are grateful to departing from Aspidistra flowers bearing pollen on the Alexei Polilov, Margarita Remizowa and Vadim Mokievsky for body. The observations are documented by video (see helpful discussion, to Leonid Averyanov for copies of his publica- tions, to two anonymous reviewers for important comments and to Electronic Supplementary Material). It is possible that the Maria Vorontsova for helpful comments and linguistic corrections. flies are attracted by wet exudate on the stigma surface. Field work was supported by the Joint Russian–Vietnamese Tropical Megaselia flies were the only flying insects visiting flowers Research and Technological Centre (JRVTRTC). We are grateful to of Aspidistra during the total of 17.1 h of observation, and Trinh Quoc Khanh (General Director of Vietnamese part of JRVTRTC) and Nguyen Dang Hoi (Deputy Director of Vietnamese their behaviour during visits was as expected from pollin- part of JRVTRTC) for their support of this research and help in ators. The size of the flies is suitable for passing through organization of field work, and to our colleagues from JRVTRTC Vu holes between stigma lobes and the perianth, thus entering Manh and Nguyen Vu for their direct help during field work. We are the chamber where the stamens are located. grateful to the staff of the Department of Electron Microscopy at Moscow University for their kind assistance and providing SEM Ants are less plausible candidates for pollination agents facilities. Funding from the Russian Foundation for Basic Research is of A. phanluongii than Megaselia flies. The ants visited gratefully acknowledged (project 12-04-31073). flowers less frequently than the flies. Field observations of ant visits showed that their behaviour on Aspidistra flowers is less focussed than that of flies. It is also important to consider that ants covered in pollen begin to clean their References antennas and legs immediately, likely not transporting Averyanov LV, Tillich H-J (2012) New taxa of Aspidistra (Aspar- pollen to other flowers. No such cleaning was observed in agaceae) from central Vietnam. Turczaninowia 15(1):5–10 the flies. Similarly to previous studies (Kato 1995; Conran Averyanov LV, Hiep NT, Loc PK (2012) Flora and vegetation of and Bradbury 2007), we observed small soil invertebrates areas allied to Phong Nha—Ke Bang. Lambert Academic, such as collembolans in the flowers of A. phanluongii. Saarbru¨cken Bell PR, Hemsley AR (2000) Green plants, their origin and diversity, Although we cannot exclude their role in pollination, we 2nd edn. Cambridge University Press, Cambridge consider this possibility unlikely. It is also unlikely that Bogner J (2005) Aspidistra locii (Convallariaceae), eine ungewo¨hnli- A. phanluongii is self-pollinated, because less than half of che Art aus Vietnam. Der Palmengarten 69:43–45 the observed flowers set fruit and because other species of Bogner J, Arnautov N (2004) Aspidistra locii (Convallariaceae), an unusual new species from Vietnam. Willdenowia 34:203–208 Vietnamese Aspidistra do not produce fruits in cultivation Borba EL, Semir JO (2001) Pollinator specificity and convergence in despite extensive flowering (Bogner and Arnautov 2004; fly-pollinated Pleurothallis (Orchidaceae) species: a multiple Bogner 2005; Tillich 2005). population approach. Ann Bot 88:75–88 Our conclusions support previous suggestions that Bra¨uchler C, Ngoc LH (2005) Aspidistra renatae (Ruscaceae)—a new species from central Vietnam. Blumea 50:527–529 Aspidistra species are likely miophylous (Vogel 1978; Chase M, Reveal J, Fay M (2009) A subfamilial classification for the Bogner and Arnautov 2004; Tillich 2005). We failed to expanded asparagalean families Amaryllidaceae, Asparagaceae document any occurrence of fungus gnats visiting flowers and Xanthorrhoeaceae. Bot J Linn Soc 161:132–136 of A. phanluongii. Megaselia flies and other members of Conran G, Bradbury JH (2007) , amphipods and Oz: niche opportunism between strangers in a strange land. Plant Species Phoridae are well known as pollinators of various tropical Biol 22:41–48 plant species (Young 1984; Hall and Brown 1993; Borba Conran JG, Tamura MN (1998) Convallariaceae. In: Kubitzki K (ed) and Semir 2001; Sakai 2002a, b; Ollerton et al. 2009). The families and genera of vascular plants. Flowering plants. They were apparently not previously recorded as flower . Lilianae (except Orchidaceae), vol 3. Springer, Berlin, pp 186–198 visitors of Aspidistra. Our observations do not exclude the De Wilde WJJO, Vogel A (2006) A new species of Aspidistra possibility that some other species of Aspidistra are polli- (Convallariaceae) from Perak, Peninsular Malaysia. Folia nated by fungus gnats, because the habitat of Malaysiana 6:125–130

123 A new species of Aspidistra (Ruscaceae s.l., Asparagales) from southern Vietnam

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