L. S. TUNG, S. Y. WONG, P. C. BOYCE, 2010 201 Studies on Homalomeneae (Araceae) of Borneo VI: Homalomena giamensis, a New Species from Sarawak, Malaysian Borneo, with Observations on its Pollination

Tung Lay Soon and Wong Sin Yeng Department of Plant Science and Environmental Ecology Faculty of Resource Science and Technology Universiti Malaysia Sarawak 94300 Samarahan Sarawak, Malaysia Peter C. Boyce Visiting Scientist Pusat Pengajian Sains Kajihayat [School of Biological Sciences] Universiti Sains Malaysia1l800 USM Pulau Pinang, Malaysia

ABSTRACT KEYWORDS A new species, Homalomena giamensis Araceae, Homalomena giamensis, polli­ L.S.Tung, S.Y.Wong & P.C.Boyce, is de­ nation, Coleoptera, Scarabaeidae, Ruteli­ scribed and illustrated from karst limestone nae, Parastasia bimaculata, karst lime­ formations at Kampung Giam, Kuching stone, Sarawak, Borneo. Division, SW Sarawak, to where it is locally endemic. As part of an ongoing project to INTRODUCTION investigate triggers and barriers in specia­ Homalomena is a predominantly Asian tion processes in Homalomena, observa­ genus of terrestrial or lithophytic solitary, tions of the pollination of Homalomena clumping, rarely scandent, very rarely giamensis were undertaken to record climbing, mesophytic, rarely helophytic, inflorescence mechanics, interactions with usually aromatic herbs occurring mainly insect visitors, and determine the pollina­ in shady perhumid or everwet forest. The tor. Five insect genera visited during genus has several centers of diversity in SE anthesis-Diptera: Drosophilidae (Coloca­ Asia, notably Sumatera and New Guinea, siomyia); Coeleoptera: Chrysomelidae: with the greatest representation on Borneo (Chaloenus & Dercetina sp.); Nitidulidae where there are perhaps in excess of 300 (genus undetermined), and Scarabaeidae: species, virtually all endemic, and with Rutelinae (Parastasia bimaculata Guerin). fewer than 20 species formally described The pollinator was determined to be the (Wong & Boyce, in press). ruteline scarab. Parastasia bimaculata is a Homalomena in Sarawak is presently rather widespread species in tropical Asia, the subject of study by a taxonomic and and has previously recorded as a pollinat­ systematic consortium coordinated from ing agent for one Bornean Homalomena. A UNIMAS (for outputs to date see, e.g., review of pollination studies of tropical Boyce & Wong, 2008, 2009; Boyce & Wong, Asian aroids to date is given. in press; Boyce & Wong, in prep.; Boyce et al., 2010; Wong & Boyce, in press; Wong et al., in press; Ng et al., in prep.). This paper Corresponding author: sywong@frst. is the first in a series investigating incidence unimas.my of pollinator guild niche partitioning as one 202 A ROJDEANA, Vol. 33

Plate 1. Homalomena giamensis L.S.Tung, S.Y.Wong & P.C.Boyce. A. Plant in habitat. ote the glossy bright green leaves. B. Detail of the petiolar sheaths and prophyll. C. Detail of sinus. D . Peti ole showing the pul vinus. E. Inflorescence at late pistillate anthesis. Note the spadix has extended beyond the spathe limb, the Colocasiomyia fli es o n the spathe limb, and the Pm'astasia beetl e visible inside the inflorescence. Note too that the length of the lower spathe exceeds that of the spathe limb. L. S. TUNG, S. Y. WONG, P. C. BOYCE, 2010 203 of the mechanisms to attempt to explain the thence tubular-apiculate for ca. 2-3 mm; extremely high rates of local and micro­ posterior lobes short, straight, ovate to endemic sympatric and parapatric often ovato-triangular, sinus rather narrow, lobes synchronously flowering species that occur 6-9 cm long; midrib raised abaxially green in Homalomena. when fresh, adaxially sunken slightly into lamina, ca. 3 mm wide, with ca. 9-11 TAXONOMY primary lateral veins on each side, diverg­ ing at 30°-80° from the midrib, adaxially Homalomena giamensis L.S.Tung, impressed abaxially slightly raised, distal­ S.Y.Wong & P.C.Boyce sp. nov. most veins curved slightly towards the apex when near the margin, interprimary veins Haec species notabilis ab combinatio barely distinguished from the primary folHs cordatis claroviride nitens, subtus lateral veins, alternating irregularly with non-glauca, venis primariis impressis primaries; secondary venation obscure, spathe inferior cum lamina spathae long­ running parallel to the interprimary veins, iore et inflorescentia mascula per anthesin striate; tertiary venation not visible; all quam resin producto. Typus: MALAYSIA, veins running into a slightly thickened Sarawak, Kuching Division, Siburan, Kam­ intermarginal vein. Inflorescences up to pung Giam, 01°19'16.1"; 110°16'16.7", 20 5 together, erect and smelling powerfully of June 2009, P.CBoyce & Wong Sin Yeng AR- anethol (anise) at anthesis, thence decli­ 2558 (holotypus SAR). nate, subtending prophyll, short, usually Paratype: Malaysia, Sarawak, Kuching less than 5 cm long; peduncle to ca. 25 cm Division, Siburan, Kampung Giam, Air long X ca. 5 mm diam. at the spathe Terjun Giam, 01°19'11.2"; 110°16'11.4", 7 insertion, glossy pale green. Spathe ca. 9- Feb. 2006, P.G.Boyce, leland ak Kisai & 11 em long, tightly furled prior to anthesis, Wong Sin YengAR-1691 (SAR). Plates 1 & 2. lower spathe inflating at pistillate anthesis Medium, somewhat robust, strongly ar­ and the margins gaping to partially reveal omatic (camphor) evergreen glabrous me­ spadix, margins also recurving, spathe limb sophytic herbs to ca. 1 m tall. Stem loosening at pistillate anthesis, thence pleionanthic, erect to ascending, ca. 3 cm inflating and then opening wide, spathe thick, medium green, internodes to ca. 1 cm interior white at anthesis, exterior pale long. Leaves up to ca. 15 together, ca. 6 glossy green with numerous faint longitu­ per module; modules starting with a dinal striae; spathe fully closing post conspicuous 2-keeled short-duration pro­ anthesis, and then turning pale green and phyllto 30 cm long, medium-green, glossy; eventually dark green; lower spathe petiole erect, 40-45 cm long, sub-terete ovoid-ellipsoid, twice as long at spathe except for the distal-most portion where limb, ca. 6-7 cm long, moderately con­ very shallowly dorsally channeled with stricted at the junction of the spathe limb, rounded margins, obscurely pulvinate ca. the constriction coinciding with junction of 2/3 along length, pulvinus ca. 4 cm long, the staminate and pistillate flower zones; petiole bases clasping, petiole bright green spathe limb ovato-triangular, ca. 3-4 cm with obscure longitudinal darker green long X ca. 3.5 cm wide (at staminate striae; petiolar sheath ca. 6-10 cm long, anthesis), limb margins very slightly recur­ ca. 1/3 of petiole length, equal, persistent, ving at staminate anthesis, apex stiffly decurrent at apex, margin erect, pale glossy mucronate to ca. 4 mm long. SPadix green; lamina rather broadly ovato-sagit­ exceeding the spathe by rapid spadix tate, 18-30 cm long X 14-25 cm wide, extension at anthesis onset, ca. 10.5 cm rubbery-leathery and somewhat quilted long, briefly stipitate; spadix contracting between the primary and interprimary post anthesis and entirely enclosed in veins, glossy bright green adaxially, paler persistent spathe throughout the fruiting green abaxially, base cordate to sagittato­ period; stipe ca. 3 mm long X 2 mm diam., cordate, tip acute, acuminate for ca. 2 cm weakly gibbose-cylindrical, and inserted 204 AROIDEANA, Vol. 33 slightly obliquely on peduncle, glossy Notes white; pistiUateflower zone ca. 3.75 cm Homalomena giamensis belongs to the long X ca. 1 cm wide, slightly less than ca. Cyrtocladon supergroup (sensu Boyce & 1/3 length of spadix, weakly fusiform; Wong, 2008), and is one of a medium­ pistils ca. 2 mm X 1 mm, rather densely sized species-complex, all currently unde­ arranged, stoutly globose-cylindrical, scribed, with glossy bright green, rather somewhat trapezoidal-compressed where rubbery-leathery cordate leaf laminae with pressed against adjacent pistils, very pale impressed primary venation, powerfully yellow-white, each pistil associated with a aromatic (camphor) vegetative tissues, ventrally located large staminode; style large, soon-degrading prophylls, the length barely differentiated; stigma globose-cap­ of the lower spathe exceeding that of the itate, smaller than ovary diam., pale gray; spathe limb, conspicuous interpistillar sta­ interpistiUar staminodes stoutly abrupt­ minodes, and the staminate portion of the ly globose-clavate on a slender stipe, spadix producing amber-colored resin clavate portion ca. 0.5 mm diam., slightly droplets during anthesis. overtopping the associated pistil, waxy white; suprapistiUar interstice zone Etymology absent; staminate flower zone to ca. 6 cm long, ca. 2/3 length of spadix, lowest From Kampung Giam, the type and only portion composed of a few sterile staminate known locality. flowers, fertile portion producing large numerous amber-colored resin droplets POLLINATION during late pistillate and at onset of staminate anthesis, these particularly to­ History wards the spadix tip, from where they Until recently pollination studies of flow down the spadix, mixing with the tropical Asian Araceae have almost exclu­ pollen strands; staminate flowers well sively been undertaken by Japanese ento­ defined from one another, ca. 2.5 mm X mological researchers, a few non-Japanese 1 mm trapezoid, comprising 3-5 truncate collaborators, and students undertaking stamens, each overtopped by a large, degrees in Japan. Notable among the flat synconnective, anthers marginal Japanese are Masako Honda-Yafuso with pollen extruded in strings and exiting (Honda-Yafuso, 1983), Tamiji Inoue (see via a notch in the synconnective margin, Kato et aI., 2000), Takao Itioka (see Kato et waxy white. Infructescence declinate to al., 2000), Makoto Kato (Kato et al., 2000), pendent, spathe entirely persistent, ca. 14 X Yuko Kumano (later Yuko Kumano-No­ 2 cm, the lower part weakly fusiform, deep mura) (Kumano & Yamaoka, 2006; Ku­ glossy green, peduncle with darker green, mano-Nomura & Yamaoka, 2009), Takashi short striae. Fruits & seeds not observed. Miyake (Miyake, Yafuso, 2003), Kuniyasu Momose (see Kato et al., 2000), Yafuso Distribution Mori (Mori & Okada, 2001), Hiroshi Okada Malaysian Borneo: Sarawak, Kuching (Mori & Okada, 2001), Toyohi Okada Division, Siburan, endemic to karst lime­ (Okada 1975, 1980, 1986, 1987, 1990; stone in the vicinity of Kampung Giam. Okada & Carson, 1980; Okada & Yafuso, 1989), Shoko Sakai (see Kato et al., 2000), Ecology Kohei Takenaka (later Kohei (Takenaka) Takano) (Takenaka et al., 2006), Masanori Terrestrial in deep leaf litter over lime­ J. Toda (Toda et aI., in press, see also stone-derived loams on ridges and moder­ Sultana et aI., 2002; Takenaka et al., 2006; ate slopes beneath lowland perhumid Yafuso et al., 2000), Seiki Yamane (see evergreen broadleaf forest on and adjacent Kato et al., 2000), Masako Yafuso (see to karst limestone. 35-50 mas!. Miyake & Yafuso, 2003), and Ryohei L. S. TUNG, S. Y. WONG, P. C. BOYCE, 2010 205

Yamaoka (see Kumano & Yamaoka, 2006; Insect identifications were undertaken Kumano-Nomura & Yamaoka, 2009). with reference to various web resources, Collaborators and students include notably www.beetle-diversity.com, www. Abang Abdul Hamid (see Kato et ai., coleoptera.org, and www.coleop-terra. 2000), Hampton L. Carson (see Okada & com. Insect vouchers are deposited in the Carson, 1980), Maklarin B. Lakim (see Entomological Museum, Faculty of Re­ Sultana et ai., 2000), Maryati Bte Mohamed source Science and Technology, University (see Toda et ai., in press), Nguyen Manh Malaysia, Sarawak. Cuong (see Sultana et ai., 2000), Farhat Sultana (Sultana et ai., 2000; see also Study Site Yafuso et ai., 2000), Yin Jian-Tao (see Takenaka etai., 2006, and Wen Shuo-Yang The study was carried out at Kam­ (see Takenaka et ai., 2006). pung Giam, Daerah Siburan, Kuching The overwhelming majority of aroid Division, Sarawak, Malaysia (01°19'16.1"N; pollination research in Asia has focussed 110 0 16'16.7''E, 50 m asl); this is the type on Drosophilidae, a family of flies (Diptera) location of Homaiomena giamensis, and commonly called fruit flies, and in partic­ the species is as far as known, endemic to ular has been directed at investigations of this locality. The population occurs as synhospitality between pollinator and in­ scattered individuals and small clumps florescence. While without doubt many along and down a narrow ridge below aroids are Drosophilidae pollinated, in­ perhumid broadleaf forest adjacent to cluding Furtadoa M.Hotta (Mori & Okada exposed karst limestone outcrops, and a 2001), the sister taxon to Homaiomena, large limestone bluff reaching to 300 m. there is considerable circumstantial evi­ The study was undertaken throughout dence, and recently hard data, that while December 2009, a period marking the early Homaiomena is visited by various Droso­ . part of the northeast monsoon, with daily philidae, they may be in the main beetle temperatures ranging from 23°-31 °C (73 0 - pollinated, it has been properly document­ 86°F), and daily afternoon and nighttime ed for one species (Kumano & Yamaoka, heavy prolonged precipitation, often deliv­ 2006; Kumano-Nomura & Yamaoka, 2009) ering > 3 cm of rain in an hour. One striking morphological feature of many Homaiomena are the conspicuous Homalomena Species Present at Giam interpistillar staminodes. The role of stami­ nodes as beetle rewards is well-document­ Homaiomena giamensis occurs sympat­ ed in other angiosperms, notably basal rically with H. borneensis Ridl., H. insignis dicots (see Armstrong & Irvine, 1990; End­ N.R.Br., H. griffithii (Jack.) Hook.f., and an ress, 1984a,b) but understanding their role as yet unidentified species. Based on in aroids is of rather recent realization. observations for the past 3 yr H. giamensis Documented in Neotropical Dieifenbachia flowers synchronously with H. borneensis. and Gearum (Spathicarpeae) which are Although differing markedly in vegetative also beetle pollinated (Young, 1986; Gon­ morphology, both have similarities in s;alves & Maia, 2006). inflorescence gross morphology, including It is perhaps notable that Homaiomena the lower spathe exceeding the length of inflorescences with interpistillar stami­ the spathe limb, conspicuous staminodes, nodes undergo distinctive changes in color and staminate flowers with a well-devel­ when preserved in ethanol, with the pistils oped synconnective. Both produce a floral staining dark brown while the staminodes odor reminiscent of anethol (anise). One remain white. Although yet to be con­ striking difference in their floral morphol­ firmed, we suspect that the ovaries are ogy is that H. giamensis produces conspic­ provided with distasteful chemicals, per­ uous amber-colored resin droplets from the haps tannins, to discourage herbivory, staminate portion of the spadix during the explaining the different reaction in ethanol. floral transition from pistillate to staminate 206 ARO IDEANA, Vol. 33

Plate 2. Homalomena giamensis L.S.Tung, S.Y .Wong & P.C.Boyce. A . Inflorescence at pistillate anthesis. Note the spathe margins are gaping to reveal the spadix, and that th e margins have also recurved. Note, too, the conspicuous resin-like droplets on the staminate portion of the spadix, and that th e spadix has extended beyond the spathe. L. S. TUNG, S. Y. WONG, P. C. BOYCE, 2010 207 anthesis whereas H. borneensis does not. mas) started at ca. 1,600 with the lower Interestingly, species related to H. hanneae spathe inflating within an hour. By ca. P.C.Boyce, S.Y.Wong & Fasihuddin B. 2,100, the lower spathe loosened to the Ahmad also produce amber-colored resin, extent that a longitudinal ( <2 mm) gap was but are otherwise florally and vegetatively created down the convolute margin. The distinct from H. giamensis, differing among inflorescence remained in this phase for ca. other characters by the spathe limb ex­ 3 hr. During this period, no insect visitors ceeding the lower spathe, producing a were present. floral fragrance of 2-butanol (sweet apricot) at anthesis, infructescence spathe turning Day 2 deep red during fruiting, petiolar sheathes By ca. 0200 the lower spathe had red with marcescent margins, and leaf loosened and inflated to widen the mar­ laminae with conspicuous extrafloral nec­ ginal gap to ca. 0.5 em (widest), while the taries. The species studied by Yuko Ku­ spathe limb remained closed, and showed mano and Ryohei Yamaoka and reported no signs of inflation. At ca. 0300, the upper as H. propinqua (Kumano & Yamaoka, spathe started to open, initially with the 2006; Kumano-Nomura & Yamaoka, 2009) small gap (ca. 0.5 em wide) and at the same is an undescribed species of the Hanneae time the spadix extended upwards through complex (H. propinqua Schott = H. ros­ the gap, lengthening by 2 em in less than an trata Griff., a colonial helophyte; see hour; this spadix extension marked the Wong, Boyce & Fasihuddin, in press). onset of detectable odor production (ane­ thole) which persisted until completion of Pollination Observation entire anthesis cycle. By ca. 0430, the upper Observations were carried out over the spathe was almost fully open, and it was whole anthesis period, enabling onset possible to observe the spadix. By ca. 0500 times, duration and inter-anthesis activity the upper spathe was almost fully open, to be monitored and recorded. All obser­ (ca. 3 em wide) and by ca. 0630 opened to vations of inflorescence movement were its fullest extent (ca. 3.5 em wide) and correlated to odor production, and arrival lower spathe was fully inflated (ca. 3.0 em and departure of insects to the inflores­ diam.). At approximately 0715 insects cence. began to appear, landing on the exterior and interior of the spathe limb. Colocasio­ Dayl myia flies mainly on exterior of the spathe, and some of the flies entering the lower At ca. 13.00 the lower spathe loosened spathe chamber but not remaining. Small slightly, but showed no signs of inflation, dark orange Nitidulidae beetles (genus the lower spathe reclosed tightly at 1,600, undetermined) were observed to visit the the cause of which is not yet understood. exterior of the spathe, and also enter the Pistillate anthesis (indicated by wet stig- lower spathe but not remain for any great

B. Detail of resin-like droplets on spadix staminate zone. C. Detail of pistillate zone ca. halfway through pistillate anthesis. Note that many of the interpistillar staminodes have already been eaten. D. Detail of pistillate zone towards end of pistillate anthesis. No staminodes remain. E. Inflorescence at end of staminate anthesis, the spathe is reclosing and the spadix has contracted to be fully enclosed within the spathe. The debris visible on the spadix is pollen paste. Just visible are two species of Chrysomelidae (a Chaloenus and perhaps a Dercetina) plundering pollen waste. F. Developing infructescence. Note that the spadix is one again fully enclosed within the persistent spathe. 208 AROIDEANA, Vol. 33 time. Metallic blue chrysomelid (Cbaloe­ period for this particular inflorescence. At nus) beetles, and also the ruteline Para­ this time activity by Chrysomelidae (Cba­ stasia bimaculata were observed to enter loenus sp. and a previously unobserved the upper spathe and lower spathe respec­ Dercetina sp.) was strongly suggestive of tively. Both crawled on the spadix, with pollen plundering. only the Parastasia descending the spadix to the pistillate zone. CONCLUSIONS At ca. 1,200, a small chewed hole was observed on the staminate zone, this • The anthesis period per inflorescence is caused by one of the Parastasia. At the ca. 43 hr, comprising a pistillate phase same time, droplets of an amber-colored lasting 24 hr, an inter-anthesis phase resin-like substance appeared on the sta­ lasting 15 hr, and a staminate phase minate flower zone, exuded from between lasting a maximum of 4 hr. the individual flowers. The exuding of the • The primary attractant appears to be resin-like substance marked the end of the odor. Although an inflorescence is pistillate phase, the stigma droplets were readily visible during the first 11 hr of already dry. This inter-anthesis phase lasted the pistillate phase, it is not until floral for 15 hr. During this long phase, numerous odor production starts that insects start individuals of the ruteline scarab Parasta­ to arrive. It is speculated that odor sia bimaculata were observed to remain production, together with suitable food inside the lower spathe chamber, were they rewards, and an appropriately sized fed on the interpistillar staminodes, and lower spathe chamber, is linked to were also mating. No other beetles nor flies 'filtering' the optimal pollinator from remained in the lower spathe during this within the guild of early morning-active phase, although periodically Chrysomeli­ insects. dae spent time on the spadix, and small • The pollinator of Homalomena giamen­ clouds of Colocasiomyia were seen 'cir­ sis is Parastasia bimaculata Guerin. cling' the inflorescence. Both presumably This is the only insect to visit the attracted by the continuing floral odor inflorescence during the pistillate production. phase, remain in the lower spathe throughout the inter-anthetic period, Day 3 and the only insect witnessed to collect At ca. 0300 onset of staminate anthesis pollen on leaving the inflorescence was indicated by the production of the during the staminate phase. This same white string-like pollen strands, these species has been shown to pollinate a mixed with the aforementioned resin drop­ Homalomena species in Borneo (Ku­ lets to form a 'paste'. At this time initially the mano & Yamaoka, 2006; Kumano-No­ Parastasia remained on pistillate zone mura & Yamaoka, 2009). within the lower spathe. At ca. 0500, the • While it is not clear what, if any, role entire staminate zone was covered by the Colocasiomyia and the unidentified pollen and at this time the beetles were Nitidulidae play in the pollination, it is observed to crawl up the spadix, crossing highly likely that the two Chrysomeli­ the staminate zone, in so doing their legs dae (Cbaloenus sp. and Dercetina sp.) were coated with both pollen strings and are pollen plunderers. pollen 'paste'. Most flew away at this time, • The interpistillar staminodes are 'sacri­ although a few remained, together with ficial' structures functioning as food two species of Chrysomelidae which ap­ rewards for the pollinator: Parastasia peared to be feeding on the pollen paste. At bimaculata. Given that the pollinator is ca. 0630, the upper spathe was observed to trapped in the lower spathe for 15 or close, and was closed tightly at ca. 0715. more hours, and in that time is active, This marked the end of the flowering and often mating, a food resource is L. S. TUNG, S. Y. WONG, P. C. BOYCE, 2010 209

important. Post-anthesis, all interpistillar --- & ---. In prep. Studies on staminodes, and the zone of staminodes Homalomeneae (Araceae) of Borneo between the pistillate and staminate X: the Homalomena of Matang, Kuch­ flower zones, show signs of consider­ ing, Sarawak. Gard. Bull. Singapore. able feeding activity. --, --& Fasihuddin, B. A. 2010. • No damage occurs to the pistils and Studies on Homalomeneae (Araceae) very little damage to the staminate of Borneo II: the Homalomena of flowers. This suggests that these struc­ Nanga Sumpa (Batang Ai) - Novel & tures are distasteful, a supposition pre-existing taxa, and notes on Iban backed up by the color reaction of the Usages. Gard. Bull. Singapore. 69(2): inflorescence when placed in ethanol 29-78. (see above). ---, --- & ---. Studies on Homalomeneae (Araceae) of Borneo VIII: additional novel Homalomena ACKNOWLEDGMENTS from Nanga Sumpa (Batang Ai), Sar­ The collaboration and support of the awak. Gard. Bull. Singapore. (In press Sarawak Forestry Department, notably Dr b). Mohd. Shahbudin Sabki, Dr Azahari Bin Endress, P. 1984a. The flowering process Omar, L.c.]. Julaihi, & Lucy Chong is in the Eupomatiaceae (Magnoliales). gratefully acknowledged. This study is Bot. Jahrb. fuer Systematik Pjlanzen­ funded by MOST! E-Science: 05-01-09- geschichte und Pjlanzengeographie SFoo06. All fieldwork has been undertaken 104: 297-319. with Sarawak Forestry Department Research ---. 1984b. The role of inner starni­ Pennits, with the most recent fieldwork nodes in the floral display of some being under Sarawak Forestry Department relic Magnoliales. P. Syst. Evol. 146: Research Permit No. NPW.907.4.2(I)-101 & 269-282. Park Pennit No. 58/20076. Gons,:alves, E. G. & Maia, A. C. D. 2006. New evidence of pollination in llTERATURE CITED Gearum brasiliense (Araceae - Spathi­ carpeae). Aroideana 29: 148--151. Armstrong, ]. E. & Irvine, A. K. 1990. Honda-Yafuso, M. 1983. Interspecific rela­ Functions of staminodia in the bee­ tionship between synhospitalic Dro­ tle-pollinated flowers of Eupomatia sophilella species (Diptera, Drosophi­ laurina. Biotropica 22: 429-431. lidae) inhabiting Alocasia odora Boyce, P. C. & Wong, S. Y. 2008. Studies on Okinawa Is., Japan. KontyU 51: on Homalomeneae (Araceae) of Bor­ 520-527. neo I: Four new species and specula­ Kato, M., Itioka, T., Sakai, P., Momose, K., tion on informal species groups in Yamane, S., Hamid, A. A. & Inoue, T. Sarawak. Gard. Bull. Singapore 60(1): 2000. Various population fluctuation 1-29. patterns of light-attracted beetles in a --- & ---. 2009. Studies on Homa­ tropical lowland dipterocarp forest in lomeneae (Araceae) of Borneo IV: Sarawak. Popul. Bcol. 42: 97-104. Homalomena specimens in the Her­ Kumano, Y. & Yamaoka, R. 2006. Synchro­ barium Beccarianum-Malesia (FI-B) of nization between temporal variation the Museo di Storia Naturale - Sezione in heat generation, floral scents and Botanica "F. Parlatore" dell'Universita pollinator arrival in the beetle-polli­ di Firenze. Webbia 64(2): 169-173. nated tropical Araceae, Homalomena --- & ---. Studies on Homalome­ propinqua. Plant Spec. Bioi. 21: neae (Araceae) of Borneo VIII: clari­ 173-83. fication of early names in Asian Kumano-Nomura, Y. & Yamaoka, R. 2009. Homalomena. Acta Phytotax. Geobot. Beetle visitations, and associations (In press). with quantitative variation of attrac- 210 AROIDEANA, Vol. 33

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