70 AROIDEANA, Vol. 34 and Visitors of Aroid : an addendum

Marc Gibemau Laboratoire d'Ecologie des Forets de Guyane (Ecofog - UMR CNRS 8172), BP 709 97387 Kourou, France [email protected]

ABSTRACf according to Tricia, I was the only one following the rule and she used thus to Data on aroid pollinators or inflores­ receive regularly a letter from France ... cence visitors were reviewed lately by I only met Julius once in September 2008 Gibernau (2003), documenting the pollina­ also during the annual lAS sale & show. As tors of 49 genera and about 125 . many people can say, he was the kind of This addendum adds information on 35 person who struck in your mind. Before genera, of which 9 are newly documented, meeting, we used to exchange e-mails, and about 60 species. In summary, we have particularly on flowering or pollination some information on pollinators or inflo­ questions. His last e-mail before my depar­ rescence visitors on 58 genera and about ture to Miami was in summary "I have 165 species. Such numbers are very low in prepared tens of questions for you"; and comparison of the family diversity (more my reply has been "I will do my best to than 110 genera and about 4,000 species). answer all of them." Effectively, we dis­ The pollination of entire groups of cussed on different occasions and particu­ is still unknown. The knowledge on the larly one evening during which we emptied pollination of each is discussed. many beers. Julius'S thirst of knowledge was immense and of course I was not able KEYWORDS to answer all his questions!!!!

Araceae, Coleoptera, Diptera, Hymenop­ INTRODUCflON tera, Pollination. Inflorescences of Araceae are mainly DEDICATION insect-pollinated and a review on Aroid inflorescences visitors and pollinators has I would like to dedicate this paper to been published (Gibernau, 2003). This Patricia Frank and Julius Boos. review presented 176 references of studies I met (Pa)Tricia on a few occasions in and observations on insects visiting inflo­ Sarawak during the International Aroid rescences of 49 genera and 125 species of Congress and in Miami during the 2008 Araceae. Studies at the family level have annual lAS sale & show at the Fairchild shown that Aroid floral traits are adapted to Tropical Botanical Garden in Miami. I was the different types of pollinating insects the invited speaker and she organized (Chouteau et at., 2008; Gibernau et at., perfectly my stay in Miami. She was very 2010b). But data on Aroid pollinators are kind to me and always "in good humor" as still scarce in comparison with the family we say in French. I still remember her diversity with more than 115 genera and guiding me through her marvelous garden. about 4,000 species (Boyce & Croat, pers. When I was a member of the lAS Board of comm.). Here is an addendum to the 2003 Directors, we had to vote on several review on Aroid inflorescences visitors and decisions via internet but the vote was to pollinators. In this paper, I summarize the be printed, signed and mailed. Apparently, data on Araceae visitors and pollinators by M. GIBERNAU, 2011 71 completing the references since 2003 and cases only one or a small number of species citing several older references omitted in of a given has been documented. Gibernau (2003). The purpose of this work Thus a generalization about the pollination is not to give an exhaustive review of this of the whole genus from the reports now subject but rather an up-to-date statement available becomes hazardous and may in of the subject, and to develop new remarks fact hide the existence of a greater diversity on aroid pollination. Another important in insect-aroid interactions. consideration is that the duckweeds (pre­ viously Lemnaceae family) are now includ­ DISCUSSION ed within the Araceae (Cabrera et al., 2008) Once again, in spite of their "unique" are not detailed in the present study. Just worth mentioning, no clear data are avail­ and very constant design, aroids have developed pollination relation­ able on duckweeds pollination except that their inflorescences/ are protogy­ ships with a great diversity of insects: bees, nous as are all known aroids (Gibernau et beetles and . If we combine the information of the previous review (Giber­ at., 2004); animal pollination is vaguely nau, 2003) and the present paper, pollina­ mentioned (may be water mites?) as well as tors or inflorescence visitors have been water, wind or self pollination (Landolt et studied or mentioned for a total of 58 at., 1998; Tsatsenko & Malyuga, 2002). genera and about 165 species. Such num­ bers are still very low in comparison to RESULTS jPe family diversity (more than 115 genera The pollinators and!or inflorescence and about 4,000 species). The pollination visitors of 35 genera and about 60 species of entire groups of Araceae is still un­ are listed in Table 1. Nine aroid genera are known. newly documented namely: , Following the classification proposed by , , , Pip­ Mayo et at. (1997), I comment below in tospatha, Pothos, , Scin­ summary what is known on pollinators or dapsus and (Table 1). Since inflorescence visitors in the several aroid 2003, 42 new studies have been published subfamilies and tribes. on aroid pollination indicating a growing interest of this topic. SubfamUy Gymnostachydoideae It is important to note that the list given in No mention according to my knowledge Table 1 does not distinguish between has been published on the pollination of pollinators and visitors. Although aroid Gymnostachys even if some other aspects inflorescences can be visited by several of its biology have been studied (Shaw, types of insects, only a few - and sometimes 1997; Shaw et al., 1997). only one - are the real pollinators for each species (Blackith & Blackith, 1993; Sivada­ SubfamUy Orontioideae san & Kavalan, 2005; Garcia-Robledo et at., 2005b; Sultana et at., 2006; Takenaka et at., A few studies have dealt with Lysichiton 2006; Barriault et at., 2009, 2010; Maia et at., and Symplocarpus species in both North 2010; Punekar & Kumaran, 2010; Quilichini America and East Asia. It appears that et al., 2010; Tung et al., 2010). Second, except perhaps for L. americanus, these when there are several species of insects, genera have a mixed-pollination system efficiency in achieving pollination can implying several taxa of insect, and maybe differ between pollinators, some species wind-pollination (see review in Gibernau, carrying few or no grains and acting 2003). Such a generalist pollination system as "simple" visitors (Albre et at., 2003; has been indirectly suggested by a multi­ Garcia-Robledo et al., 2005a; Takenaka et variate analysis of the floral traits associated al., 2006; Barriault et at., 2009, 2010; with the pollination (Chouteau et at., 2008; Quilichini et al., 2010). Third, in several Gibernau et al., 2010b). Almost nothing is Table 1. Aroid pollinators/visitors cited in the literature between 2003 and 2010, plus a few older references omitted in the previous review (Gibernau, 2003). Subfamily/tribe Genera Bees Beetles Flies Others References Orontioideae Symplocarpus Honey bees Kevan, 1989 Orontioideae Lysichiton Tanaka, 2004 / Anthurium Curculionidae Franz, 2003, 2007 Anthurieae Euglossine Hentrich et al., 2007,2010 Pothoideae/ Pothos hymenoptera Yadav, 1998 Potheae / Spathiphyllum Euglossine Hentrich et al., Spathiphylleae 2010 Monsteroideae/ Monstera Nitidulidae Chouteau et al., Monstereae 2007 Monsteroideae/ Scindapsus Drosophilidae Sultana et al., 2006 Monstereae Monsteroideae/ Rhaphidophora hymenoptera Yadav, 1998 Monstereae beetles flies Zhu & Croat, 2004 Lasioideae Dracontioides flies Boos, 1997 Gon,.:alves, 2005 Lasioideae Anaphyllopsis Nocturnal Gibernau et al., insects? 2010a Cockroaches? / Gearum Dynastinae Bogner & Spathicarpeae Gon,.:alves, 1999 Gon,.:alves & Maia, 2006 Aroideae/ Dynastinae Maia et al., unpub!. Spathicarpeae Table 1. Continued. Subfamily/tribe Genera Bees Beetles Flies Others References Aroideae/ Drosophilidae Okada, 1986, 1987 Homalomeneae Sultana et al., 2006 Scarabaeidae Kumano & Yamaoka, 2006 Chrysomelidae Kumano-Nomura & Yamaoka, 2009 Scarabaeidae . Tung et al., 2010

Aroideael Drosophilidae Sultana et al., 2006 Homalomeneae Aroideael Scarabaeidae Maia et al., 2010 Philodendreae AroideaelCaladieae Dynastinae Garcia-Robledo et al., 2004, 2005a, 2005b Garcia-Robledo, 2010 AroideaelCaladieae Dynastinae Maia& Schlindwein, 2006 Aroideael Necrophagous flies Austen, 1896 Thomsonieae Beetles Yadav, 1998 Trigona (Apidae) Nitidulidae Drosophilidae Punekar & Kumaran, 2010 Hybosoridae Muscidae Hetterscheid, Staphylinidae unpubl. Ph.D. Scarabaeidae Rutelinae Table 1. Continued. Subfamily/tribe Genera Bees Beetles Flies Others References Aroideae/Schisma - Schismatoglottis Drosophilidae Sultana et al., 2006 toglottideae Aroideae/Schisma - Drosophilidae Sultana et at., 2006 toglottideae Aroideae/ Diptera Yadav, 1998 Ctyptocotyneae Aroideae/ Lagenandra Diptera Yadav, 1998 Ctyptocotyneae Aroideae/ Neurochaetidae McAlpine, 1978, Colocasieae 1988 Drosophilidae Okada, 1986, 1987 Ivancic et at., 2005 Sultana et at., 2006 Aroideae/ Drosophilidae Toda & Okada, Colocasieae 1983 Okada, 1986, 1987, 1990 Ivancic et al., 2004, 2008 Miyake & Yafuso, 2005 Sultana et al., 2006 Aroideae/ Steudnera Drosophilidae Takenaka et al., Colocasieae 2006 Sultana et at., 2006 Aroideae/ Diptera Yadav, 1998 Arisamateae Barriault et al., 2009, 2010 Table 1. Continued. Subfamily/tribe Genera Bees Beetles Flies Others References Aroideael Not found Quilichini et at., Ambrosineae 2005 AroideaelAreae Diptera Yadav, 1998 AroideaelAreae Tberiophonum Staphylinidae Sivadasan & Kavalan, 2005 AroideaelAreae Beetles Yadav, 1998 AroideaelAreae Necrophagous flies Austen, 1896 Blowflies- Angioy et at., 2004 Calliphoridae AroideaelAreae Necrophagous flies Austen, 1896 Calliphoridae Blackith & Blackith, 1993 Muscidae Fanniidae Piophilidae AroideaelAreae review Gibernau et at., 2004 Halictidae Diaz & Kite, 2006 Staphylinidae Sphaeroceridae Linz et at., 2010 Drosophilidae Urru et at., 2010 Stokl et al., 2010 Quilichini et at., 2010 AroideaelAreae Staphylinidae Diptera midges Boyce, 2008 76 AROIDEANA, VoL 34 known about the pollination of the third dospatha, Stenospermation), even if genus, Orontium. It is probably pollinated beetles have been mentioned for Rho­ by insects, such as flies and beetles with no dospatha. Drosophilid flies have been more indications as to specific species observed to oviposit on the inflores­ involved (Klotz, 1992). cences of Scindapsus and may be pollinators (Sultana et at., 2006). For Subfamily Pothoideae Monstera no clear information is avail­ able, bees and beetles have been • Tribe Potheae. No study of pollinators mentioned visiting their inflorescences for the three genera of this tribe (Pothos, (see review in Gibernau, 2003). The Pothoidium, Pedicellarum), except one only published study has recently indi­ mention of Hymenoptera in Pothos cated one species, M. obliqua, to be (Yadav, 1998). pollinated by nitidulid beetles in French • Tribe Anthurieae. Only one genus is Guiana (Chouteau et at., 2007). The included in this tribe, Anthurium, but it pollination ecology of Rhaphidophora is the most diverse of the family (with africana by a scarab beetle has been more than 1,200 species, Croat, pers. studied in West Africa, but not pub­ comm.). A great diversity of pollination lished (Beath, 1993), whereas Hyme­ systems exists in this genus: bees noptera are mentioned for Asian species collecting resins and/or oily fragrances (Yadav, 1998). Interestingly, a bee­ (Hentrich et at., 2007, 2010), curculionid pollination system has been indirectly pollination looking for a feeding, and suggested for Monstereae based on a probably a reproductive site (Franz, multivariate analysis of some floral traits 2003, 2007), and humming-bird pollina­ associated with pollination (Chouteau et tion, the unique mention of vertebrate at., 2008; Gibernau et at., 201Ob). pollination in Araceae (Kraemer & Schmitt, 1999). It is not unlikely that further studies will reveal new pollina­ Subfamily Lasioideae tion systems in this genus. The pollinators from the five Asian genera are not known (, Cyr­ Subfamily Monsteroideae tosperma, , Podotasia, ). In South America, no clear data are • Tribe Spathiphylleae. No mention of available on the pollinators of the four pollinators for the genus Hotochtamys. genera (Anaphyllopsts, Dracontium, Dra­ In contrast, the pollination of the genus contioides, ). A recent study over Spathiphyllum appears to be variable. several years was unable to determine the On one hand, bees have been observed pollinators of Anaphyllopsis suggesting a to collect pollen, wax or oily fragrance very rare event or maybe night-pollination (Hentrich et at., 2010), on the other (Gibemau et at., 2010a). The genera Dra­ hand flies looking for nectar (and/or a contium and Dracontioides are suggested reproductive site?) (see review in Gi­ to be -pollinated (Boos, 1997; Zhu & bernau, 2003). Croat, 2004; Gon~alves, 2005). Nitidulid • Tribe Anadendreae. No mention of beetles have been observed in spathes of pollinators for the genus Anadendrum, Urospatha (T. Boos, pers. observ.). Interest­ the unique genus of this tribe. ingly, the only African species, • Tribe Heteropsidae. No mention of (cited as ) senegaiense, is ap­ pollinators for the genus Heteropsis, parently pollinated by nitidulid beetles in the unique genus of this tribe. West Africa (Knecht, 1983). • Tribe Monstereae. The pollinators from In multivariate analyses on some floral five genera are not known (Allosche­ traits associated with pollination, the three mone, Amydrium, , Rho- studied species of Lasioid are grouped with M. GIBERNAU, 2011 77 the two Orontioid species Lysichiton genera have shown that Gearum (Gon­ camtschatcense and Symplocarpus renifo­ \;alves & Maia, 2006) and Taccarum lius. This original pollination group inter­ (Maia et al., unpub!.) are pollinated by mediate between fly- and bee-pollination scarab beetles (Dynastidae). systems was hypothesized to represent a • Tribe Philodendreae. The only genus of generalist (undefined?) pollination system this tribe, Philodendron, has been the (Chouteau et aI., 2008; Gibernau et al., model of many pollination works. All 2010b). the actual results indicate the same pollination system implying scarab bee­ Subfamily Calloideae tles (Dynastidae) even if other insects can be found within the inflorescences The pollination ecology of is not (review in Gibernau, 2003; Maia et al., known. Old references (review in Giber­ 2010). nau, 2003) mentioned visiting flies. A • Tribe Homalomeneae. Drosophilid flies modern study is particularly necessary to have been identified as pollinators of evaluate the potential attractive role of the Furtadoa (Sultana et al., 2006). Pollina­ white 'flag' spathe. Such work would be tors of Homalomena are not clearly interesting in regard to the recent phyloge­ identified. On one hand, drosophilid netic results that place Calla among the flies have been observed within their Aroideae taxa and not in an intermediate inflorescences (Okada, 1986, 1987; Sul­ position between the basal taxa with tana et at., 2006). On the other hand, bisexual flowers and the most derived ones scarab beetles were also observed (Le. Aroideae) with unisexual flowers (Kumano & Yamaoka, 2006; Kumano­ (Cabrera et at., 2008). Nomura & Yamaoka, 2009; Tung et at., 2010). Thus the pollination situation in Subfamily Zamiocu1cadoidae Homalomena needs to be clarified. In multivariate analyses on some floral • Tribe Zamioculcadeae. No mention of traits associated with pollination, the pollinators for the two genera of this genus Homalomena is distributed both tribe (Zamiocutcas, Gonatopus). within fly- and beetle-pollinated species • Tribe Stylochaetoneae. No mention of (Chouteau et al., 2008; Gibernau et at., pollinators for the genus Stytochaeton, 2010b). Several hypotheses can be the unique genus of this tribe. proposed. First, Homatomena species have a mixed pollination system impli­ cating both flies and scarab beetles. Subfamily Aroideae Second, some Homalomena species are fly-pollinated and others are beetle­ • Tribe Dieffenbachieae. No pollinators pollinated. Third, Homalomena species are known for the genus , on are pollinated by only one type of the contrary several studies have illus­ insects, flies or beetles, the second type trated the pollination of being inflorescence visitors. species (see review in Gibernau, 2003). • Tribe Anubiadeae. Only one reference Scarab beetles (Dynastidae) are the is available on one species from West pollinators of different Dieffenbachia Africa, gigantea, indicating species even if other insects have been nitidulid beetles as visitors and/or pol­ observed and/or collected in the inflo­ linators (Knecht, 1983). rescences. • Tribe Schismatoglottideae. The pollina­ • Tribe Spathicarpeae. No mention of tors of four genera are unknown pollinators for six genera of this tribe (Hottarum, , Phyma­ (, , Gorgoni­ tarum, Heteroaridarum). The two stud­ dium, , , ied genera, Piptospatha and Schismato­ ). Recent studies on two glottis, are pollinated by drosophilid 78 AROIDEANA, Vol. 34

flies (Sultana et al., 2006; Toda et al., Ancbomanes, Nepbtbytis and Pseudoby­ 2006). The same is probable for Ar­ drosme are visited by beetles (Nitiduli­ idarum nicolsonii in Sarawak even if dae) and flies. Trigona bees observed beetles (nitidulidae and staphylinidae) on inflorescences of Ancbomanes might have been observed to eat the male probably be only visitors (review in flowers (fertile and sterile) (Gibemau & Gibemau, 2003). In multivariate analy­ Boyce, unpub!.; Bogner, pers. observ. ses on some floral traits associated with in Knecht, 1983). Interestingly, the pollination, some species of Ancbo­ genus Scbismatoglottis, with a disjunct manes and Nepbtbytis have typical fly distribution (Asia - America), may pollinated floral characters while other present different pollination systems species have more mixed floral charac­ on the two continents. In multivariate ters (Chouteau et al., 2008; Gibemau et analyses on some floral traits associated aI., 2010b). Further studies are needed with pollination, the genus Scbismato­ to assess if pollination is achieved in glottis is distributed within both fly- and these genera by a mixed-pollination beetle-pollinated species (Chouteau et system or by only one type of insect. al., 2008; Gibemau et al., 2010b). • Tribe Aglaonemateae. No • Tribe Cryptocoryneae. The two genera mention for the genus Ag/aodorum. of this tribe, Cryptocoryne and Lagen­ On the other hand is andra have very similar floral morphol­ certainly pollinated by drosophilid flies ogy and ecology. No proper study on (review in Gibemau, 2003). But the their pollination ecology has been results from multivariate analyses on published. But many observations and some floral traits associated with polli­ insect collections are available, suggest­ nation suggest that Aglaonema species ing fly pollination by a trap mechanism possess floral characters typical of in aquatic habit (Yadav, 1998; see beetle-pollination (Chouteau et a/., review in Gibemau, 2003; Jacobsen, 2008; Gibemau et al., 2010b). unpub!. data) • Tribe Culcasieae. For the two genera of • Tribe Zomicarpeae. No mention of the tribe (Cutcasia, ), beetles pollinators for the four genera of this (Nitidulidae) and flies (Drosophilidae) tribe (Zomicarpa, Zomicarpella, Ulea­ have been collected within their inflo­ rum, ). rescences. In multivariate analyses on • Tribe Caladieae. No pollinator mention some floral traits associated with polli­ for three of the genera of this tribe nation, these two genera appear to (Scapbispatba,jasarum, ). For distribute in separate positions. Cu/ca­ the three other genera (Caladium, sia clusters within the fly-pollinated Cblorospatba and ), many species whereas Cercestis appears with­ studies and observations indicate a in the beetle-pollinated species (Chou­ scarab beetle pollination (review in teau et at., 2008; Gibemau et at., 2010b). Gibemau, 2003; Maia & Schlindwein, Further studies are needed to assess if 2006). In Xantbosoma, many different pollination is achieved in these genera insects (euglossine bees, drosophilid by a mixed-pollination system or by flies, scarab beetles) have been ob­ only one type of insect. served in the inflorescences (review in • Tribe Montrichardieae. This tribe con­ Gibemau, 2003), but recent studies tains only the genus Montricbardia. indicate that certainly only scarab bee­ Observations of bees visiting inflores­ tles are pollinators, other insects have cences may provide erroneous informa­ been probable visitors (Garcia-Robledo tion about its pollination that is et al., 2004, 2005a, 2005b; Garcia­ achieved by scarab beetles (review in Robledo 2010). Gibemau, 2003; Gibemau et at., 2003). • Tribe Nephthytideae. The inflorescenc­ • Tribe Zantedeschieae. The only genus es of the three genera of this tribe of this tribe, Zantedescbia, may be M. GIBERNAU, 2011 79

pollinated in its spontaneous distribu­ able to detect any pollinators, rare non­ tion area by beetles, but accurate field flying small insects or arthropods were studies are needed (see review in observed visiting the inflorescences Gibemau, 2003). (Quilichini et al., 2005). • Tribe Callopsideae. No mention of • Tribe . Only one genus of this pollinators for the genus , the tribe has not been studied from a unique genus of this tribe. pollination point of view, namely La­ • Tribe Thomsonieae. Observations have zarum. The pollination of all the other shown that there is a great diversity of eight genera has been studied (Arum, visiting insects (bees, beetles, flies) in Biarum, Dracunculus, , Heli­ Amorpbopballus, but until recently only codiceros, Sauromatum, 7beriopbonum one study was published on its pollina­ and Typbonium), and it's achieved by tion indicating beetle pollination for the necrophilous or saprophilous flies and/ African species A. jobnsonii (Beath, or beetles (review in Gibemau, 2003; 1993). Recent observations suggest a Blackith & Blackith, 1993; Yadav, 1998; great diversity of pollination systems in Angioy et al., 2004; Gibemau et aI., this genus implying bees, beetles and/ 2004; Sivadasan & Kavalan, 2005; or flies by deception or not (Punekar & Boyce, 2008; Espindola et al., 2010; Kumaran, 2010; Hetterscheid, unpub!. Linz et al., 2010; St6kl et al., 2010; Ph.D). , a close re­ Quilichini et al., 2010; Urru et al., 2010) lated genus, presents floral odors and a except for one known exception (Diaz pollen morphology very similar to & Kite, 2006; Urru et al., 2010). Amorpbopballus species pollinated by • Tribe Arisaemateae. The two genera of carrion flies and beetles, such hypoth­ this tribe and Arisaema are esis must be confirmed by a field study pollinated by various mycophilous and (Kite and Hetterscheid, 1997; Kite et al., saprophilous flies (and a few beetles) 1998; van der Ham et al., 2005; Hetter­ and thunder bugs (Thysanoptera) (re­ scheid, unpub!. Ph.D). view in Gibemau, 2003; Barriault et al., • Tribe Arophyteae. No mention of polli­ 2009, 2010). nators for the three genera of this • Tribe Colocasieae. No pollinator men­ Malagasy tribe (Aropbyton, Carlepby­ tion is available for three out of the six ton, Colletogyne). genera of the tribe (, , • Tribe Peltandreae. No mention of pol­ ). The pollinators of Coloca­ linators for the genus Typbonodorum, sia and Alocasia species are drosophilid while pollination of is highly flies ovipositing on the inflorescences. specialized by flies (Chloropidae) ovi­ Interestingly, each aroid inflorescence is positing on the inflorescences (review visited by two fly species, one ovipos­ in Gibemau, 2003). iting on the staminate zone, the other • Tribe Arisareae. The only genus of this on the pistillate one but not damaging tribe, , seems to be pollinated the (Toda & Okada, 1983; by diverse mycophilous or saprophilous Okada, 1986, 1987, 1990; Ivancic et al., flies (review in Gibemau, 2003). 2004, 2005, 2008; Miyake & Yafuso, • Tribe Ambrosineae. The only genus of 2005; Sultana et al., 2006). Other insects this tribe, Ambrosina, has been recently have been mentioned visiting the inflo­ studied. Its complex inflorescence struc­ rescences but apparently they are not ture suggests a complicated pollination pollinators. Recently the pollination mechanism with a small animal being ecology of Steudnera has been studied able to move from a lower (dorsal) indicating a similar fly pollination sys­ "male" chamber from one inflorescence tem except that only one fly species is to an upper (ventral) "female" chamber present (Takenaka et al., 2006). of another inflorescence (Barabe et aI., • Tribe Pistieae. Vague mention of insects 2004). But field observations were not observed in the minute inflorescence of 80 AROIDEANA, Vol. 34

Pistia, but no published study on its Ambrosina and Arisarum (Araceae). pollination is available. Can . .f. Bot. 82(2): 282-289. Barriault, I., M. Gibernau & D. Barabe. In conclusion, data on the pollination 2009. Flowering period, thermogene­ ecology and biology on aroids has greatly sis, and pattern of visiting insects in increased in the last 10 years with about 70 (Araceae) in papers published, compared to 145 be­ Quebec. Botany 87: 324-329. tween 1879 and 2000. But many tribes and ---, D. Barabe, L. Cloutier & M. genera are still non-documented. In several Gibernau. 2010. Pollination ecology other cases, the data are not conclusive and reproductive success in ]ack-in­ between a mixed pollination system imply­ the Pulpit (Arisaema triphyllum) in ing at least two types of pollinators or Quebec (Canada). Bioi. 12(1): pollination by only one type of insects 161-171. associated with other visiting insects. In Beath, D. N. 1993. Biology afforest Araceae these cases further studies are needed to in Ghana. unpub\. Ph.D. Thesis, assess which is the real pollination system. University of Aberdeen, U.K. 274 p. An indirect approach has been recently Blackith, R. & R. Blackith. 1993. Diptera developed through multivariate analyses (Insecta) attracted to the Dragon's based on some floral traits associated with mouth lily (Dracunculus vulgaris pollination (Chouteau et al., 2008; Gibernau Schott) in Ireland. Bull Irish Biogeo. et at., 2010b). These analyses were able to Soc. 16: 2-7. separate four combinations of floral traits: Bogner, ]. & E. G. Gonr,:alves. 1999. The generalist, bee-, fly-, beetle-pollinated taxa. genus Gearum N. E. Br. (Araceae: Such results are exploratory. Even if these Tribe Spathicarpeae). Aroideana 22: analyses can predict from a set of floral 20-29. character the probable pollination type of Boos, ]. O. 1997. Observations on New an unknown species, they must be consid­ World Araceae - Lasieae. Aroideana ered as hypotheses to be further tested in 20: 13-26. the field or experimental conditions. Boyce, P. C. 2008. A taxonomic revision of Biarum. Curtis's Bot. Mag. 25(1): 2- ACKNOWLEDGMENTS 119. I wish to thank Derek Burch for con­ Cabrera, L. I., G. A. Salazar, M. W. Chase, structive comments and corrections on the S.]. Mayo,]. Bogner & P. Davila. 2008. manuscript. Phylogenetic relationships of aroids and duckweeds (Araceae) inferred LITERATURE CITED from coding and noncoding plastid DNA. Amer. .f. Bot. 95(9): 1153-1165. Albre, ]., A. Quilichini & M. Gibernau. Chouteau, M., M. McClure & M. Gibernau. 2003. Pollination ecology of Arum 2007. Pollination ecology of Monstera italicum (Araceae). Bot. .f. Linn. Soc. obliqua (Araceae) in .]. 141: 205-214. Trap. Beol. 23(5): 607-610. Angioy, A.-M., M. C. Stensmyr, I. Urru, M. ---, M. Gibernau & D. Barabe. 2008. Puliafito, I. Collu & B. S. Hansson. Relationships between floral charac­ 2004. Function of the heater: the dead ters, pollination mechanisms, life horse arum revisited. Proc. Roy. Soc. forms and habitats in Araceae. Bot. .f. London, ser. B 271: S13-S15. Linn. Soc. 156: 29-42. Austen, E. E. 1896. Necrophagous diptera Diaz, A. & G. C. Kite. 2006. Why be a attracted by the odour of flowers. An. rewarding trap? The evolution of floral Mag. Nat. Hist. Zool. Bot. Geo!. 18: rewards in Arum (Araceae), a genus 237-240. characterized by saprophilous polli­ Barabe, D., C. Lacroix & M. Gibernau. nation systems. Biol . .f. Linn. Soc. 88: 2004. Aspects of floral morphology in 257-268. M. GIBERNAU, 2011 81

Espindola, A, L. Pellissier & N. Alvarez. ceae) in French Guiana.]. Trop. Ecol. 2010. Variation in the proportion of 19: 103-107. visitors of ---, D. Macquart & G. Przetak. 2004. along its distributional range in rela­ Pollination in the genus Arum - a tion with community-based climatic review. Aroideana 27: 148-166. niche analyses. Oikos. doi: 10.1111/ ---, M. Chouteau, K. Lavallee & D. j. 1600-0706.2010. 18937.x Barab~. 201Oa. Notes on the Phenol­ Franz, N. M. 2003. Systematics of Cy­ ogy, Morphometry and Floral Biology clanthura gen. n., a new genus of of Anaphyllopsis americana (Ara­ Derelomini (Coleoptera: Curculioni­ ceae). Aroideana 33: 183-191. dae). Insect Syst. Evol. 34: 153-198. ---, M. Chartier & D. Barabe. 201Ob. ---. 2007. Pollination of Anthurium Recent advances towards an evolu­ (Araceae) by derelomine flower wee­ tionary comprehension of Araceae vils (Coleoptera: Curculionidae). Int. pollination. Diversity, Phylogeny, and ]. Trop. BioI. 55: 269-277. Evolution in the : Pro­ Garcia-Robledo, c., G. Kattan, C. Murcia & ceedings of the Fourth International P. Quintero-Marin. 2004. Beetle polli­ Conference on the Comparative Biol­ nation and predation of Xantho­ ogy of the Monocotyledons. O. Seberg, soma daguense (Araceae) in an An­ G. Petersen, A S. Barford & J. Davis. dean cloud forest in Colombia.]. Trop. Aarhus University Press: 101-114. Beol. 20: 459-469. Gon~alves, E. G. 2005. A revision of the --, --, -- & --. 2005a. genus Dracontioides Engl. (Araceae), Equal and opposite effects of flo­ including a new species from Bahia, ral offer and spatial distribution on . Aroideana 28: 21-31. fruit production and pre-dispersal --- & A C. D. Maia. 2006. New seed predation in Xanthosoma da­ evidence of pollination in Gearum brasiliense (Araceae - Spathicarpeae). guense (Araceae). Biotropica 37(3): Aroideana 29: 148-151. 373-380. Hentrich, H., R. Kaiser & G. Gottsberger. ---, P. Quintero-Marin & F. Mora­ 2007. Floral scent collection at the Kepfer. 2005b. Geographic variation perfume flowers of Anthurium rubri­ and succession of arthropod commu­ nervium (Araceae) by the kleptopar­ nities in inflorescences and infructes­ asitic orchid bee Aglae caerulea (Eu­ cences of Xanthosoma (Araceae). glossini). Beotropica 13: 149-155. Biotropica 37(4): 650--656. --, -- & --. 2010. Floral ---. 2010. Restoration of plant-pollina­ biology and reproductive isolation by tor interactions: pollination neighbor­ floral scent in three sympatric aroid hood and asymmetric pollen flow species in French Guiana. Plant BioI. between restored habitats in a beetle 12(4): 587-596. pollinated aroid. Restor. Beol. 18(Sl): Ivancic, A., V. Lebot, O. Rouspard, J. 94-102. Quero Garcia & T. Okpul. 2004. Gauthier, M.-P., L. D. Barabe & A Bru­ Thermogenic flowering of (Colo­ neau. 2008. Molecular phylogeny of casia esculenta, Araceae). Can.]. Bot. the genus Philodendron (Araceae): 82: 1557-1565. delimitation and infrageneric classifi­ ---, O. Rouspard, J. Quero Garcia, V. cation. Bot.]. Linn. Soc. 156: 13-27. Lebot, V. Pochyla & T. Okpul. 2005. Gibernau, M. 2003. Pollinators and visitors Thermogenic flowering of the giant of Aroid inflorescences. Aroideana 26: taro (, Ara­ 66-83. ceae). Can.]. Bot. 83: 647-655. ---, D. Barabe, D. Labat, P. Cerdan & ---, ---, ---, M. Meiteras, T. A Dejean. 2003. Reproductive biology Molisale, S. Tara & V. Lebot. 2008. of arhorescens (Ara- Thermogenesis and flowering biology 82 AROIDEANA, Vol. 34

of Colocasia gigantea, Araceae. ]. Molecular phylogeny of the genus Plant Res. 121: 73--82. Arum (Araceae) inferred from multi­ Kite, G. C. & W. 1. A. Hetterscheid. 1997. locus sequence data and AFLPs. Tax­ Inflorescence odours of Amorpho­ on 59(2): 405-415. phallus and Pseudodracontium (Ara­ Maia, A. C. D. & C. Schlindwein. 2006. ceae). Phytochem. 46: 71-75. (Araceae) and Cy­ ---, ---, M. J. Lewis, P. c. Boyce, clocephala celata (Coleoptera, Dynas­ J. Ollerton, E. Cocklin, A. Diaz & tinae): a well-established pollination M. S. J. Simmonds. 1998. Inflorescence system in the Northern Atlantic rain­ odours and pollinators of Arum and forest of Pernambuco, Brazil. Plant Amorphophallus (Araceae). Reprod. Bioi. 8: 529-534. Bioi. S. J. Owens & P. J. Rudall. Kew, ---, ---, D. M. A. F. Navarro & M. Royal Botanical Gardens: 295-315. Gibernau. 2010. Pollination of Philo­ Kevan, P. G. 1989. How honey bees forage dendron acutatum (Araceae) in the for pollen at skunk cabbage, Symplo­ Atlantic Forest of northeastern Brazil: carpus foetidus (Araceae). Apidologie a single scarab beetle species guaran­ 20: 485-490. tees high fruit set. Int. ]. Plant Sci. Klotz, 1. H. 1992. On the biology of 171(7): 740-748. 1. (Araceae), McAlpine, D. K. 1978. Description and golden club or floating Arum. Aroi­ biology of a new genus of flies related deana 15: 25-33. to Anthoclusia and representing a Knecht, M. 1983. Aracees de Cote d'Ivoire - new family (Diptera, Schizophora, Contribution a l'etude biosystema­ Neurochaetidae). Ann. Natal Mus. tique des representants d'Aracees de 23(2): 273-295. la Cote d'Ivoire. Phanerogamarum ---. 1988. Studies in upside-down flies Monographiae - Tom. XVII. Cramer, (Diptera: Neurochaetidae). Part II. Vaduz, 290 p. Biology, adaptations, and specific Kraemer, M. & U. Schmitt. 1999. Possible mating mechanisms. Proc. Linn. Soc. pollination by hummingbirds in An­ N. S. Wales 110(1): 59--82. thurium sanguineum Engl. (Araceae) Miyake, T. & M. Yafuso. 2005. Pollination Pl. Syst. & Evol. 217: 333-335. of Alocasia cucullata (Araceae) by Kumano, Y. & R. Yamaoka. 2006. Synchro­ two Colocasiomyia flies known to be nization between heat generation, specific pollinators for . temporal variations in floral scents Plant Spec. Bioi. 20: 201-208. and pollinator arrivals in the beetle­ pollinated tropical Araceae, Homalo­ Okada, T. 1986. Estimation of the routes of mena propinqua. Plant Spec. Bioi. synhospitalic distribution of the genus 21(3): 173-183. Drosophilella Duda (Diptera, Droso­ Kumano-Nomura, Y. & R. Yamaoka. 2009. philidae), with descriptions of three Beetle visitations and association with new species from Malaysia and In­ quantitative variations of attractants in donesia. Proc. jap. Soc. Syst. Zool. 33: floral odors of Homalomena propin­ 32-39. qua (Araceae). ]. Plant Res. 122: ---. 1987. Further notes on the genus 183-192. Drosophilella Duda with descriptions Landolt, E., I. ]ager-Ziirn & R. A. A. Schnell. of two new species from Indonesia 1998. Extreme adaptations in Angio­ (Diptera, Drosophilidae). Proc. jap. spermous hydrophytes. Encyclopedia Soc. Syst. Zool. 36: 38-45. of Plant Anatomy, Band XIII, Teil. 4, ---. 1990. Subdivision of the genus Gerbriider Borntraeger Verlagsbuch­ Colocasiomyia de Meijere (Diptera, handlung, Berlin. Drosophilidae) with descriptions of Linz, J., J. Stokl, I. Urru, T. KrUgel, M. C. two new species from Sulawesi and Stensmyr & B. S. Hansson. 2010. note on color adaptation of synhospi- M. GIBERNAU, 2011 83

talic species. Proc.Jap. Soc. Syst. Zool. new species of the genus Colocasio­ 42: 66-72. myia de Meijere (Diptera: Drosophili­ Punekar, S. A. & K. P. N. Kumaran. 2010. dae) and its host plant Steudnera Pollen morphology and pollination colocasiaejolia (Araceae) in Yunnan, ecology of Amorphophallus species China. Entomol. Sci. 9(1): 79-9l. from North Western Ghats and Konkan Tanaka, H. 2004. Reproductive biology of region of . Flora 205: 326-336. Lysichiton camtschatcense (Araceae) Quilichini, A., A. Torre & M. Gibernau. in Japan. Aroideana 27: 167-17l. 2005. Preliminary data on the biology Toda, M. J. & T. Okada. 1983. Ecological and reproduction of Ambrosina bassii studies of floricolous Drosophilella in 1. (Araceae) in Corsica. Aroideana 28: Burma with descriptions of three new 37~l. species from Burma and the Philip­ ---, D. Macquart, D. Barabe,J. Albre & pines (Diptera, Drosophilidae). Kon­ M. Gibernau. 2010. Reproduction of tyCt 51(2): 169-184. the West Mediterranean endemic Ar­ Tsatsenko, 1. V. & N. G. Malyuga. 2002. um pictum (Araceae) on Corsica. Lemnaceae Flowering and pollina­ Plant Syst. & Evol. 287(~): 179-187. tion. http://www.mobot.org/jwcrossl Shaw, D. E. 1997. Gymnostachys anceps R. duckweediRusse/biology-e.htm Br.: fruit, germination, and a discus­ Tung, 1. S., S. Y. Wong & P. C. Boyce. sion of the possible means of disper­ 2010. Studies on Homalomeneae (Ara­ sal. Aroideana 20: 71-78. ceae) of Borneo VI: Homalomena ---, 1. H. Bird & 1. 1. Forsberg. 1997. giamensis, a new species from Sara­ Gymnostachys anceps R. Br.: Austra­ wak, Malaysian Borneo, with obser­ lian range and habitat in Southeastern vations on its pollination. Aroideana Queensland. Aroideana 20: 64-70. 33: 201-21l. Sivadasan, M. & R. Kavalan. 2005. Flower­ Urru, 1., J. StcSkl, J. Linz, T. KrOgel, M. C. ing phenology and beetle pollination Stensmyr & B. S. Hansson. 2010. in injaustum N. E. Br. Pollination strategy in Cretan Arum (Araceae). Aroideana 28: 104-112. lilies. Bioi. ]. Linn. Soc. 101: 991- StcSkl, ]., A. Strutz, A. Dafni, A. Svatos, J. 100l. Doubsky, M. Knaden, S. Sachse, B. S. van der Ham, R. W. J. M., G. B. Grob, W. 1. Hansson & M. C. Stensmyr. 2010. A A. Hetterscheid, W. Star & V. H. B. J. deceptive pollination system targeting 2005. Notes on the genus Amorpho­ drosophilids through olfactory mimic­ phallus (Araceae) - 13. Evolution of ry of yeast. Curro Bioi. 20: 1-7. pollen ornamentation and ultrastuc­ Sultana, F., Y.-G. Hu, M. J. Toda, K. ture in Amorphophallus and Pseudo­ Takenaka & M. Yafuso. 2006. Phylog­ dracontium. Grana 44: 252-265. eny and classification of Colocasio­ Yadav, S. R. 1998. Adaptive Significance of myia (Diptera, Drosophilidae), and its phenology and spadix architecture in evolution of pollination mutualism Araceae of Western Ghats of India. with aroid . Syst. Entomol. 31(4): Acta Bot. Yunnan. Suppl. X: 31~0. 684-702. Zhu, G. & T. B. Croat. 2004. Revision of Takenaka, K., J.-T. Yin, S.-Y. Wen & M. J. Dracontium (Araceae). Ann. Miss. Toda. 2006. Reproductive ecology of a Bot. Gard. 91: 59~67.