104 AROIDEANA, Vol. 28 Flowering Phenology and Beede Pollination in infaustum N.E.Br. ()

M. Sivadasan and Raghavan Kavalan* DepanrnnentofBotany University of CaJicut CaJicut University P.O. Kerala----673 635 India (e-mail: [email protected]) .. Present address: Jawahar Navodaya Vidyalaya, Almati Dam Site, Bijapur Dist., Karnataka-5S6 201, India

ABSTRACT deals with temperate genera (Gibernau, 2003). The only tropical aroid that Tberiophonum injaustum N.E.Br. (Ara­ is fairly well-known in this regard is Amor­ ceae), an endemic species of SW India, phophallus, mainly from the work of Pijl consists of small tuberous . The spe­ (937). The works of Hubbard (895), cies was found to have protogynous inflo­ Cleghorn (913), Knoll (926), Pijl (1937), rescences and to be visited by several Meeuse & Hatch (960), Dormer (960), kinds of beetles. Beetles belonging to the species Anotylus rubidus Cameron (Cole­ Monteith (973), Williams & Dressler optera, Staphylinidae) are found to be the (976), Golubev & Volokitin (983), Shaw main potential pollinators. Other species & Cantrell (983), Valerio (984), Gotts­ observed were only casual visitors. A de­ berger & Amaral (984), Pellmyr (985), tailed account on the process of canthar­ Young (986), Whitehill (993), Patt et al. ophily in this species is given with explan­ (995), Gibernau et al. (999), & Gibernau atory illustrations. et al. (2003) are some of the significant works on members of Araceae. In India KEYWORDS not much work has been done on this as­ pect especially in Araceae, and only scat­ Araceae, cantharophily, India, Tberio­ tered and fragmentary information is avail­ phonum. able. The work of Barnes (934) who de­ scribed the mode of pollination and pol­ INTRODUCTION linators in Arisaema occurring in the The family Araceae is mainly tropical in Nilgiri Hills, and that of Banerji (947) on distribution, with a few temperate genera, trilobatum are worth mention­ constituted of herbs and vines comprising ing. A detailed account of cantharophily in about 105 genera and about 3,300 species Amorphophallus hohenacheri (Schott) (Mayo et al., 1997). It is characterised by Eng1. & Gehrm., with a brief review on the a spadix which is subtended by a spathe. general pollination biology with special Tberiophonum B1. of the subfamily Ar­ reference to Araceae was provided by Si­ oideae is a small genus of five species en­ vadasan & Sabu (989). Beath (996) pro­ demic to South and Central India with one vided the details of pollination in Amor­ species extending to Sri Lanka (Sivadasan phophallus johnsonii. Gibernau (2003) & Nicolson, 1982). Tberiophonum injaus­ provided an up-dated list of aroid polli­ tum N.E.Br. is restricted to SW India. nators which is the latest covering all the Information on pollination biology of available data on this aspect. the members of Araceae is meagre as most According to the most far-reaching gen­ of the information available on this aspect eralizations, as far as is known the Araceae M. SIVADASAN, R. KAVALAN, 2005 105 are considered to be exclusively ento­ cinity by the commencement of SW mon­ mophilous, and in most cases the polli­ soon (June-September). nators are beetles (Coleoptera) and flies It is a small, tuberous herb reaching a (Diptera) (termed cantharophily and my­ height of about 12 to rarely 30 cm (Fig. 1). ophily respectively). Bees (Hymenoptera) The corm is somewhat subglobose to cy­ are rarely reported as pollinators as in the lindric in shape, ca. 0.5-4.00 cm thick and case of Spathiphyllum (Williams & Dress­ 0.7-1.8 cm diam. The leaves have distinct ler, 1976) and Anthurium (A Beath, T. petioles which arise from the top of the Croat, pers. comm.), especially those with tuber. The lamina is glabrous, reticulate­ sweet-scented flowers such as the mem­ veined, and is highly variable in size and bers of Anthurium sect. Calomystrium. shape, narrowly to broadly hastate-sagit­ Beetles have the oldest fossil record of any tate, rarely broadly ovate-cordate, 4.6-12 of the four anthophilous orders of insects cm long, lateral lobes rounded to ovate or viz., Coleoptera, Lepidoptera, Diptera and with acute tip. The inflorescence is borne Hymenoptera. Thus, it is widely presumed on an erect peduncle, which arises in leaf that beetles were the pollinators of the ear­ axils, and during its early developmental liest entomophilous angiosperms (Gray­ stage is completely encased by the vagi­ urn, 1984). nate leaf-base. This species, like many oth­ Floral morphology is the most important ers in the genus, also reproduces by veg­ element in determining the system of pol­ etative means. lination of a particular type of blossom. Structural features have long been recog­ STUDY SITE AND METHODS nised as a diagnostic specialization for par­ The study was carried out at the Calicut ticular broad categories of insect-pollina­ University Campus and its surroundings tion. Faegri & Pijl (1979) gave an account (latitude: 11°35'-45'N, longitude: 75°45'- of the syndrome of cantharophily. It in­ 50'E, at an altitude of 40-60 m; average cluded the features such as a pollination annual rainfall of ca. 300 cm; and temper­ unit with few visual attractions, no nectar ature ranging from 17-35°C). Flowering guide, generally large, flat, cylindric or period is from June-September and begins shallow, bowl-shaped-sometimes closed, one month after the first rains. easy of access, dull colour, frequently Flowering individuals in the study area greenish or white, and strong odour­ were located and tagged. A total of 50 in­ fruity or aminoid. The beetle-pollinated florescences were observed and data col­ flowers are protogynous. In addition to the lected during the study period. Observa­ protogyny, cantharophily is generally as­ tions were made on the morphology and sociated with phenomena like thermogen­ phenology of the inflorescences along esis, musty odour, crowded pistillate and with the details of the pollinators and their staminate flowers enclosed by bracts dur­ activities. Insect visitors to the inflores­ ing a brief nocturnal anthesis. cences were collected, and preserved in Many beetles of different families are re­ 70% alcohol for identification. Insects ported to be involved in the pollination of were also collected from other popula­ Araceae, with most of the beetles from the tions along with inflorescences to verify family Scarabeidae (Grayum, 1984). But whether the pollinators are common to all Staphylinidae are also common in Araceae populations. Voucher specimens of all outside the New World (Gibernau, 2003). beetles are deposited at Commonwealth Institute of Entomology, London.

THERIOPHONUM INFAUSTUM OBSERVATIONS Theriophonum infaustum, an endemic Floral Morphology species of SW India grows abundantly on The inflorescence is solitary, completely the Calicut University Campus and its vi- enclosed by the petiolar sheath in the ear- 106 AROIDEANA, Vol. 28

.~. :'" -", ~::~.-...... <:: .. /.~,,,...... :::'_._ .. -...... ,...... 2:::..-.....

E u C") E u

D

Fig. 1. Tberiopbonum inJaustum N.E.Br.: A. Habit; B. Inflorescence; C. Inflorescence with spathe cut open to show spadix; D. Spadix. (f: female flowers; i: interstice; m: male flowers. n: neuter flowers; sat: stalk of spadix-appendix; sa: spadix-appendix).

ly stage. Peduncle ca. 5.5 cm long and 4 with a zone of neuter flowers ca. 3 rom rom diam., greenish to purplish. Spathe long, followed by naked slender interstice erect, to about 6 cm long, differentiated, ca. 6 rom long, followed by male portion into a basal completely convolute tubular ca. 5 rom long and terminating in a creamy portion, and an upper initially convolute, purplish sterile appendix ca. 2.5 cm long but later with a spreading yellowish-green with a distinct stipe ca. 3.5 rom long. Pis­ limb portion separated by a slight constric­ tillate flowers 4-8, in a single whorl; ovary tion between the two; the basal convolute ovoid, yellowish green, unilocular with 4- tubular portion is 1.0-1.6 cm long, 0.6-1.2 5 basal and apical sub-orthotropous cm diam., upper limb portion oblong, 3-5 ovules; stigma sessile, discoid. Neuter cm long and 1-1.5 cm broad. The overlap­ flowers subulate to subclavate, verrucu­ ping of the margins of the spathe is either lose, creamy purplish, ca. 3.5 mm long. to the right or to the left as observed ear~ Staminate flowers 23-48, closely arranged, lier by Barnes (1938). The spadix is sessile, purplish, sessile and truncate, each with 2 almost equalling or slightly shorter' than distinct anther lobes, dehiscing by apical the spathe, to ca. 4.5 cm long, with basal pores. Fruit a berry, 6-8 per infructes­ female portion ca. 2 mm long, contiguous cence, each to ca. 1 cm long and 5 rom M. SIVADASAN, R. KAVALAN, 2005 107 diam., ovoid, green, 2-3-seeded. Seeds to escape from the tube, crawling up, but broadly ovoid with basal white tissue. on every attempt they fell down because of the smoothness of the inner surface of Phenology and Pollination the basal tube of the spathe. The beetles Mechanism carrying pollen moved across and overthe pistillate flowers, Simultaneously effecting The spadices were observed to be pro­ pollination. The release of odoudasted for togynous. The exact time of receptivity of a period of 3.5 to 6 hours, but with a rel­ pistillate flowers is difficult to determine atively short period of maximum intensity. because there is no clear indication to this When the odour started fading, the spathe­ effect. Initially the spathe is completely limb began to roll and close forming a tu­ rolled tightly around the spadix (Fig.2A). bular structure, leaving only a small open­ At maturity, the spathe unrolled and ing. at the top. At this stage, the spathe at opened during the afternoon of the first the constriction between the limb and the day. A detailed study of the spathe open­ basal tube pressed the spadix (Fig. 2E) ing revealed that the margin of the limb -thereby completely closing the basal tube began to unfold or unroll from the top to­ with the insects trapped dnside the tube wards the base to its basal constriction _and unable to escape. Figs. 2B,2C). Generally the unfolding took ·Theanthesis of-the male flowers com­ place between 1.30 P.M. and 4 P.M. and menced during the afternoon of the sec­ the spathe attained its maximum opening ond day between 2,30P:M. and 5.30 P.M. with maximum width of spathe limb (Fig. The anthers dehisced by the apical pores 2C), between 4.00 P.M. and 6.30 P.M. The releaSing pollen grains in masses. They spathe limb stands erect with the tip'slight­ were deposited at the base of the tube ly incurved and .in some cases it touched formed by the rolled limb just above the the tip of the spadix-appendix. At this po­ closed basal tube. At this stage the spathe sition, the sterile appendix of the spadix and the neuter flowers -became more or and the staminate flowers are completely less flaccid and the constriction became exposed. The basal tubular portion ofthe loose (Fig. 2F), allowing insects .trapped spathe enclosing the pistillate and neuter for a day inside the tube to crawl up. flowers is not tightly closed at this stage so While they crawled up, their bodies were that there is space between the spathe­ covered with pollen fallen from above and tube and the spadix with pistillate and also that accumulated at the base of the neuter flowers (Fig. 2D). This facilitates .limb. No special organ or organs of the easy access of small insects into the tube. insects were noticed as depositories and Simultaneous with the opening of the carrier organs of pollen: -Pollen grains spathe, a foetid and pungent -odour was were found all over the body of the in­ emitted by the spadix-appendix. The in­ sects. The insects carry thepoHen -and if tensity of the odour reached at its maxi­ they visit other iriflorescences, get en­ mum when the spathe has fully opened trapped as mentioned earlier, and when (between 4.00 P.M. and 6.30 P.M.). The they escape by crawling over.the pistillate beetles carrying pollen grains from other flowers, effect cross-pollination. inflorescences that they have previously The spathe showed signs of degenera­ visited were probably attracted by this tionon the third day onwards. No insects odour and alight on either the smooth spa­ were found to remain in the -inflorescence dix-appendix or on the inner surface of on the third day, except a few small dead the spathe-limb. The beetles alighting on ones. The first symptom of. degeneration them slipped off and fell down into the of the spathe was the inwar.dcurling of the tube of the spathe and are trapped there. spathe-margin (Fig. 2G) while the spathe Since the flowers are protogynous, the and spadix still stand erect. Gradually the stigma is receptive during this period. A colour of the spathe started fading, the large number of such trapped insects tried spadix-appendix shrunk, lost its colour 108 AROIDEANA, Vol. 28

Fig. 2. A-H: Inflorescence of Theriophonum injaustum N.E.Br.-Different stages during and after pollination: A. Inflorescence with completely convolute spathe; B-C. Different stages of unfolding of spathe from top to the base of the limb; D. A portion of the spathe removed showing the space between the spadix and the spathe; E. A portion of the spathe removed showing the closely rolled spathe with the spadix; F. A portion of the spathe removed showing the gap between the spadix and the spathe after anthesis; M. SIVADASAN, R. KAVALAN, 2005 109 and the whole inflorescence along with Table 1. Insect visitors of Therio­ the peduncle bent down (Fig. 2H) and phonum infaustum. touched the ground. Within 7-10 days all parts of the spathe and spadix degenerat­ COLEOPTERA ed except the fertilized pistillate flowers, LEIODIDAE which eventually became the fruits. Ptomophaginus sp. STAPHYLINIDAE Insect Visitors Oxytelus incisus Motschulsky Several kinds of insects were collected Anotylus latiusculus (Kraatz) from the inflorescences. Out of the six Anotylus rubidus (Cameron) species of beetles (Coleoptera) collected Anotylus sp. nr. A. rubidus from the inflorescences, only two were (Cameron) observed regularly in and around the in­ Ph ilonth us flavocinctus Motschul­ florescences. A list of insect visitors of sky Tberiophonum injaustum is presented in Atheta sp. Table 1. The taxa of Anthocoridae of Het­ DIPTERA eroptera could not be identified. PSYCHODIDAE The beetles belonging to the species PSYCHODINAE Anotylus rubidus Cameron (Figs. 21, 2J) of Genus et sp. indet. Staphylinidae (Coleoptera) and another unidentified species of this genus, and HEPTEROPTERA seemingly related to this species, were ANTHOCORIDAE found to be frequent visitors and consid­ Genus et sp. indet. ered to be the potential pollinators. The NABIDAE other species of beetles appeared to be ca­ Phorticus? cingalensis sual visitors; perhaps they might also sometimes facilitate cross-pollination, but are not considered as regular pollinators. dix-appendix or on the inner side of the The same species were also collected from spathe-limb in search of the source of the different populations of Tberiophonum in­ odour, but lost their grip and fell down jaustum occurring in parts of Kerala other into the basal tube. The beetles remained than the present study site. in the spathe-tube for about 24 hours. The members of the species Philonthus Then they began to leave the inflores­ flavocinctus Motschulsky were captured cence on the evening of the second day, only once during the study and members most leaving between 3 P.M. and 6 P.M. of Heteroptera (Table 1) were found to be The Anotylus species captured immediate­ only occasional visitors. ly after leaving the inflorescence were No insect-visits to the inflorescences found to carry large amounts of pollen on were observed during the forenoon of the their legs and body. first day. The first visits were by species of Anotylus which arrived by the evening of DISCUSSION the first day when the odour emitted by the spadix-appendix was strong and the Tberiophonum injaustum was observed spathe started to close. They easily located to be pollinated mainly by the beetles of the inflorescence and alighted on the spa- the species Anotylus rubidus Cameron of

G. Inflorescence with the margins of the spathe rolled; H. The inflorescence in drooping position after pollination: 1. Anotylus rubidus (Cameron)-the pollinator beetle; J. Single hind wing-enlarged. 110 AROIDEANA, Vol. 28

Staphylinidae (Coleoptera), which were (Gottsberger & Silberbauer-Gottsberger, considered as the main potential pollina­ 1991). tors. Other species observed were only ca­ Adaptation for effective pollination and sual visitors. Apart from Araceae, canthar­ subsequent trapping of beetles inside the ophily is prominent in many families of the spathe tm the pollen shedding of the host Magnoliales, Annonales (Faegri & Pijl, 1979; were reported earlier in Typhonium (Ba­ Thien, 1974, 1980; Armstrong & Drum­ nerF, 1947; Monteith, 1973; Armstrong, mond, 1986), Nymphaeaceae(Prance, 1979). Typhonium and 7beriophonum 1980), and Palmae (Henderson, 1986). share many of these characteristics. Both In beetles, the sense of smell is much genera are in the subtribe Arinae of the more highly developed than the visual subfamily . The inflorescence of sense, and'beetle-flowers' are often white Typhonium generally opened in the morn­ or dull in colour and frequently have ing .and remained odourless until late in strong odours. These odours are usually the afternoon. On the evening of the first fruity, spicy, or like certain fermentation­ day, the spathe stood erect and emitted a odours and distinct from the sweet-odour foetid odour which attracted the dung­ of flowers pollinated by bees, moths and beetles of the genus Onthophagus (Scara­ butterflies (Raven et al., 1976). The com­ baeidae), which are the potential pollina­ binationof less striking flower (spathe) tors, to the blossoms. Dakwale & Bhatna­ colour, (green, purple or pale .green with gar (1997) reported that 7beriophonum purple tinge on thespathe~tube), strong crenatum is pollinated by crepuscular foetid odour, absence of nectar, peculiar Ceratopogonid flies, blood-sucking female midges, and hence has a very different inflorescence structure with the· spathe di­ pollination syndrome from that of7berio­ vided into lower· chamber (tHbe) and an phonum injaustum. Theriophonum is upper limb, and spadix with basal fertile closely related to Typhonium within the portion and an apical sterile barren spa­ tribe and they represent the basal dix-appendix, protogyny,etc. are the syn­ taxa of this tribe (Renner & Zhang, 2004). drome of beetle pollination. Protogyny is The tribe Areae is composed of 9 genera an essential element in thefunction·ofthis (Mayo etal., 1997), and except for the ge­ type of trap~blossoms. Protandry would-be nus Lazarum, pollinators are known for the prevalent form of dichogamy in taxa each of the genera (Gibernau, 2003). Sta­ pollinated by bees or flies whereas pro­ philinidae, associated with other beetles or togyny would prevail in taxa poUinated by flies, have been mentioned in , beetles, wasps or other vectors. , , and some The inflorescences of 7beriophonum in­ (Gibernau, 2003); while ap­ jaustum have a strong unpleasant smell pears to be pollinated by Scarabaeidae emitted from the spadix, which is doubt­ and flies, and by only cal­ less for the purpose of .attracting beetles liphorid flies. It is interesting to note that involved in their cross-pollination. The in­ Typhonium is a genus pollinated only by sects were irresistibly attracted by the beetles with staphylinids among them. strong odours, whether or not the blos­ In contrast to .thecondition in Typhon­ somsoffersome compensatory attractant ium, the spathe opened only in the after­ (pollen, food bodies, etc.) (Faegri & Pijl, noon in 7beriophonum injaustum and the 1979). The importance of odour as a bee­ odour is emitted simultaneously with the tle-attractant has been demonstrated in opening of the spathe, and lasts for a pe­ many of the tropical aroid species of the riod of 3.5 to 6 hours. The spathe never genera like Arisaema (Barnes, 1934), Ty­ bent backwards as in the case of Typhon­ phonium (Cleghorn, 1914; Banerji,1947; tum. Here the spathe remained erect even Monteith, 1973; Pijl, 1953), Arum (Knoll, after anthesis of staminate flowers. This is 1926; Dormer, 1960), Amorphophallus (Si­ an advantage to this species for getting vadasan & Sabu, 1989) and Philodendron pollinated by its own pollen grains and for M. SIVADASAN, R. KAVALAN, 2005 III dispersal of potten by the pollinators. As carrion beetles (Phaeocrous amplus) in the case of Typhonium, the beetles in a Ghanaian rain forest. ]. Trop. trapped are not offered any nutritive re­ Ecol. 12:409---418. wards by the inflorescence. From this it is Cleghorn, M. E. 1913. Notes on the polli­ evident that the insects are attracted by the nation of Colocasis antiquorum. ]. odour produced by the spadix-appendix Asiat Soc. Bengal 9:313. and accidentally trapped inside the ---. 1914. A note on the floral mech­ spathe-tube. anism of Typhonium trilobatum.]. & Proc. ASiat. Soc. Bengal 10:421---424. ACKNOWLEDGMENTS Dakwale, S. R. & S. Bhatnagar. 1997. Midge-trapping behaviour and floral The authors wish to thank the Depart­ biology of Theriophonum crenatum ment of Science, Technology and Environ­ Blume (Araceae). Curro Sci. 797:606- ment, Government of Kerala for providing 608. funds for a Research Project on Biosystem­ Dormer, K.]. 1960. The truth about polli­ tatic study of the genus Theriophonum Bl., nation in Arum. New Phytol. 59:298- and the present study was undertaken as 301. a part of the project. The invaluable ser­ Faegri, K. & L. van der PijI. 1979. The Prin­ vices rendered by the Director and the ciples of Pollination Ecology, 3rd ed. concerned staff of the C.A.B. International Pergamon Press, New York. Institute of Entomology, London, UK. for Gibernau, M. 2003. Pollinators and visitors identification of the insects are gratefully of Aroid inflorescences. Aroideana acknowledged. The various help by Mr. M. 26:66-83. S. Kiran Raj, Research Associate working Gibernau, M., D. Barabe, P. 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