Pollinators and Visitors of Aroid Inflorescences

Marc Gibernau Laboratoire d’E´volution & Diversite´Biologique Universite´de Toulouse III 118 Route de Narbonne, Baˆt. IV R 3—B 2 31062 Toulouse Cedex 4 France e-mail: [email protected]

ABSTRACT view of this subject, as Thomas Croat (2000) did in his history and current status Data on aroid was first sum- of systematic research with , but to marized by Grayum (1984) who docu- give, in the first place, a statement of the mented 35 genera and about 90 . A subject and to develop some remarks on second summary was published in 1997 in aroid pollination. The Genera of Araceae (Mayo et al., 1997) with 38 genera and less than 100 species listed including data from Grayum (1986, RESULTS 1990). This paper brings the reference list Summarizing data from Grayum (1984, up to date since 1997, documenting the 1990) and Mayo et al. (1997), and includ- pollinators of 49 genera and about 125 ing omitted and new publications, the pol- species. These numbers are still very low in comparison with the diversity of the Ar- linators of 49 genera and about 125 spe- aceae family which contains 105 genera cies are documented in Table 1. These and about 3,300 species. Some questions numbers are still very low in comparison on aroid pollination are developed in the with the diversity of the Araceae family discussion. which contains 105 genera and about 3,300 species (Mayo et al., 1997). Thus, KEY WORDS pollinators are cited for only 47% of the genera. Araceae, Coleoptera, Diptera, floral The references not previously published morphology, Hymenoptera, pollination. document pollinators from 25 known gen- era and from 8 new genera namely: Agla- INTRODUCTION onema, , , , Pi- Data on aroid pollinators was first sum- nellia, Rhaphidophora, and marised by Grayum (1984) who docu- . mented 35 genera and about 90 species. A It is important to note that the list given second summary was published in 1997 in in Table 1 does not distinguish between The Genera of Araceae (Mayo et al., 1997) pollinators and visitors. Although aroid in- with 38 genera and less than 100 species florescences can be visited by several listed including additional data from Gra- types of insects, only a few—and some- yum (1986, 1990). In 13 years, the polli- times only one—are the real pollinators nators (and sometimes the pollination bi- for each species (Bogner, 1981; Shaw & ology) of only three new genera and Cantrell, 1983; Valerio, 1984; Pellmyr & about 10 species were documented. In this Patt, 1986; Young, 1986; Patt et al., 1995). paper, I summarise the data on Araceae Second, when there are several species of pollinators by completing the references insects, efficiency in achieving pollination since 1997 and citing several omitted ref- can differ between pollinators, some spe- erences earlier than 1997. The purpose of cies carrying few or no pollen grains and this work is not to give an exhaustive re- acting as ‘‘simple’’ visitors (Young, 1988a). Table 1. Adapted from Mayo et al. (1997), references cited after Grayum (1984, 1986, 1990) with new and update references on aroid pollinators/visitors. Subfamily/ Genera Bees Beetles Flies Others References Orontioideae Lysichiton Staphylinidae Anthomyiidae Hesse, 1980 Lauxaniidae Pellmyr & Patt, 1986 Various flies Willson & Hennon, 1997 Tanaka, 1998 Orontioideae Symplocarpus ‘‘hive Staphylinidae Drosophilidae Wind (self- Trelease, 1879 bees’’ Psychodidae pollina- Moodie, 1976 Chloropidae tion) Grimaldi & Jaenike, 1983 Ceratopogoni- Various in- Camazine & Niklas, 1984 dae vertebrates Uemera et al., 1993 Chironomidae Simuliidae Cecidomyiidae Muscidae Pothoideae/ Anthurium Euglossine Curculionidae Cecidomyiidae Self-pollina- William & Dressler, 1976 Anthurieae Staphylinidae Drosophilidae tion Madison, 1979 (wind?) Croat, 1980 Humming- Valerio & Villalobos, 1980 bird Beath, 1998 Kraemer & Schmitt, 1999 Schwerdtfeger et al., 2002 Monsteroideae/ Monstera Trigone Scarabaeidae Madison, 1977 Monstereae Ramirez & Gomez, 1978 Ramirez, 1980 Grayum, unpubl. data Monsteroidae/ Rhaphidophora Scarabaeidae Beath, 1993 Monstereae Monsteroideae/ Rhodospatha ‘‘beetles’’ Schatz & Grayum, unpubl. data Monstereae Table 1. Continued. Subfamily/tribe Genera Bees Beetles Flies Others References Monsteroideae/ Spathiphyllum Euglossine Chrysomelidae Drosophilidae William & Dressler, 1976 Spathiphylleae Trigone Tephritidae Montalvo & Ackerman, 1986 Halictidae Lewis et al., 1988 Vogel, 1990 Chuah et al., 1996 Beath, 1998 Lasioideae Cyrtosperma Nitidulidae Knecht, 1983 Lasioideae Dracontium ‘‘myiophilous’’ Grayum, pers. observ. Lasioideae Urospatha Nitidulidae J. Boos, pers. observ. Calloideae ‘‘small flies’’ Krause, 1908 Knuth, 1909 / Scarabaeidae Madison, 1979 Dieffenbachieae Bawa & Beach, 1981 Valerio, 1984 Young, 1986, 1988a, 1988 b, 1990 Seres & Ramirez, 1995 Beath, 1998, 1999 Grayum, pers. observ. Aroideae/ Scarabaeidae Hubbard, 1895 Philodendreae Schrottky, 1910 Madison, 1979 Gottsberger & Amaral, 1984 Ramirez, 1989 Gottsberger & Silberbauer- Gottsberger, 1991 Seres & Ramirez, 1995 Grayum, 1996 Croat, 1997 Table 1. Continued. Subfamily/tribe Genera Bees Beetles Flies Others References Beath, 1998 Gibernau et al., 1999, 2000 Gibernau & Barabe´, 2002 Grayum, pers. observ. Aroideae/ Furtadoa Drosophilidae Mori & Okada, 2001 Homalomeneae Aroideae/ Scarabaeidae Yafuso & Okada, 1990 Homalomeneae (neotropics) Drosophilidae Momose et al., 1998 Chrysomelidae (Asia) Sultana et al., 2002 In Malaya Grayum, pers. observ. Schatz, pers. observ. (Malaya) Aroideae/ ‘‘bees’’ Scarabaeidae Madison, 1979 Montrichardieae Ramirez & Brito, 1992 Gibernau et al., 2003 Grayum & Schatz, unpubl. data Aroideae/ Nitidulidae Knecht, 1983 Anubieae Scarabaeidae Aroideae/ Nitidulidae J. Bogner, pers. observ. Schismatoglot- tideae Aroideae/ Staphylinidae J. Bogner, pers. observ. Schismatoglot- tideae Aroideae/ Drosophilidae J. Bogner, pers. observ. Schismatoglot- tideae Aroideae/ Scarabaeidae Madison, 1981 Caladieae Beach, 1982 Pellmyr, 1985 Pellmyr, pers. comm. Table 1. Continued. Subfamily/tribe Genera Bees Beetles Flies Others References Aroideae/ Staphylinidae Madison, 1981 Caladieae Aroideae/ Scarabaeidae Madison, 1979 Caladieae Croat, 1981 Beath, 1998 Grayum, pers. observ. Aroideae/ Euglossine Scarabaeidae Drosophilidae Schrottky, 1910 Caladieae (Nitidulidae) Madison, 1979 Valerio, 1988 Tsacas & Chassagnard, 1992 Seres & Ramirez, 1995 Moron, 1997 Beath, 1998 Goldwasser, 2000 Garcia-Robledo et al., 2003, unpubl. data Grayum, pers. observ. Aroideae/ Amorphophal- Trigone Asilidae Knuth, 1909 Thomsonieae lus Cetoniidae Platystomatidae Backer, 1913 Nitidulidae van der Pijl, 1937 Scarabaeidae Bogner, 1976 Silphidae Mayo et al., 1982 Staphylinidae Sivadasan & Sabu, 1989 Hybosoridae Beath, 1993, 1996, 1998 Histeridae Weryszko-Chmielewska & Stpiczynska, 1995 Kite & Hetterscheid, 1997 van der Ham et al., 1998 Kite et al., 1998 Table 1. Continued. Subfamily/tribe Genera Bees Beetles Flies Others References Aroideae/ Drosophilidae Yafuso & Okada, 1990 Aglaonemateae Sultana et al., 2002 Aroideae/ Scarabaeidae Letty, 1973 Zantedeschieae Scydmaenidae Singh et al., 1996 Aroideae/ Trigone Nitidulidae Lonchaeidae Knecht, 1983 Nephthytideae Thompson & Rawlins, 1986 Beath, 1993 Aroideae/ Nitidulidae Drosophilidae Knecht, 1983 Nephthytideae Aroideae/ Pseudohydros- Scaphidiidae Choridae Bogner, 181 Nephthytideae me Staphylinidae Sphaeroceridae Aroideae/ Nitidulidae Drosophilidae Knecht, 1983 Culcasieae Beath, 1993 Aroideae/ Nitidulidae Drosophilidae Knecht, 1983 Culcasieae Beath, 1993, 1998 Aroideae/ Nitidulidae Drosophilidae van der Pijl, 1933 Colocasieae Scarabaeidae Anthomyiidae Carson & Okada, 1982 Staphylinidae Neurochaetidae Honda-Yafuso, 1983 Shaw & Cantrell, 1983 Okada & Yafuso, 1989 Vogel, 1990 Yafuso & Okada, 1990 Yafuso, 1993, 1994, 1999 Miyake & Yafuso, 2003 Aroideae/ Drosophilidae Cleghorn, 1913 Colocasieae Carson & Okada, 1982 Honda-Yafuso, 1983 Yafuso & Okada, 1990 Table 1. Continued. Subfamily/tribe Genera Bees Beetles Flies Others References Aroideae/ Centropogoni- Petch, 1928 Cryptocoryneae dae McCann, 1943 Ephydridae Vogel, 1990 Phoridae Aroideae/ Chloropidae Goldberg, 1941 Peltandreae Syrphidae Patt et al., 1995 Aroideae/ Sciaridae Thrips Barnes, 1934 Arisamateae Mycetophilidae Pijl van der, 1953 Dolichopodidae Vogel, 1978, 1990 Phoridae Rust, 1980 Keroplatidae Bierzychudek, 1981, 1982, 1984 Menzel & Martens, 1995 Vogel & Martens, 2000 Aroideae/ ‘‘fungus gnat’’ Hubbard, 1895 Arisamateae Ceratopogoni- Schmucker, 1925 dae Vogel in Jaeger, 1976 Vogel, 1990 Aroideae/ Mycetophilidae Knuth, 1909 Arisareae Sciaridae Vogel, 1978 Psychodidae Koach & Galil, 1986 Chironomidae Herrera, 1988 Drosophilidae Aroideae/ ‘‘flies’’ Mites? Knuth, 1909 Ambrosineae Killiam, 1929, 1933 Aroideae/ ‘‘Curculio- da Silva, 1981 Pistieae nids’’? Aroideae/ Psychodidae Arcangeli, 1886a, 1886b 1 Centropogoni- Gaye, 1885 dae Church, 1908 Ceratopogonidae Table 1. Continued. Subfamily/tribe Genera Bees Beetles Flies Others References Sphaeroceridae Knuth, 1909 Sciaridae Schmucker, 1925 Simuliidae Knoll, 1926 Knoll, 1930 Tonnoir, 1940 Grensted, 1947 Sowter, 1949 Kullenberg, 1953 Prime, 1954, 1960 Lamb, 1956 Dormer, 1960 Braverman & Koach, 1982 Koach, 1985 Papp & Rohacek, 1987 Withers, 1988 Rohacek et al., 1990 Lack & Diaz, 1991 Drummond & Hammond, 1991, 1993 Me´ndez & Obeso, 1992 Boyce, 1993 Kite et al., 1998 Jezek, 2002 Diaz & Kite, 2002 Albre et al., 2003 Drummond & Boorman, 2003 Koach & Galil, unpubl. data Gibernau et al., unpubl. data Table 1. Continued. Subfamily/tribe Genera Bees Beetles Flies Others References Aroideae/ Biarum Scarabaeidae Flies? Koach, 1985 Areae Staphylinidae Sepsidae Empididae Drosophilidae Aroideae/ Staphylinidae Calliphoridae Arcangeli, 1886b, 1886c 1 Areae Dermestidae Muscidae Delpino, 1890a, 1890b Histeridae Sarcophagidae Knuth, 1909 Sylphidae Schmucker, 1930 Malvesin-Fabre, 1945 Boyce, 1986 Seymour & Schultze-Motel, 1999 Aroideae/ Eminium Scarabaeidae Sphaeroceridae Koach, 1985 Areae Staphylinidae Sepsidae Pap & Rohacek, 1987 Ortidae Muscidae Ulidiidae Heleomyzidae Ephydridae Aroideae/ Calliphoridae Schnetzler, 1879 Areae Stensmyr et al., 2002 Seymour et al., 2004 Gibernau et al., unpubl. data Aroideae/ Sauromatum Scarabaeidae Sepsidae Dakwale & Bhatnagar, 1982, 1985 Areae Bruchidae Muscidae Otitidae Sarcophagidae Calliphoridae Table 1. Continued. Subfamily/tribe Genera Bees Beetles Flies Others References Aroideae/ Theriophonum Ceratopogoni- Dakwale & Bhatnagar, 1997 Areae dae Aroideae/ Scarabaeidae Cleghorn, 1914 Areae Staphylinidae Banerji, 1947 Nitidulidae van der Pijl, 1953 Ptiliidae Monteith, 1973 Scydmaenidae 1 Arcangeli’s papers were first published in 1883 but they were reprinted in 1886. ihtehgetnme freferences three of example, genera, For number pollinators. their highest about those the are gen- graph The the with on). on named so actually and era three with with nine five reference, two, one with ref- three (13 point). or erences two one, left are by (59%) documented top genera only documented of the the records of (i.e. Most no are pollinators there the (56), genera oid references. 31 with ed pollinators: their about the num- references highest of on the with ber named those are actually graph the genera ref- The three or erences. pollinators. two one, are by the (59%) documented genera only of documented the records (56) of genera Most no aroid are the of there mentioned most As for genera. polli- before, aroid on the for available nators references of number insect-aroid in exis- diversity the interactions. greater hide a fact of in tence becomes may available and now genus hazardous reports whole the the of from has pollination generalization the genus a about given Thus a documented. of small been a species or of one only number cases several in Third, ?? 1. Fig. aedvlpdpliainrelationship pollination developed aroids have design, inflorescence constant very DISCUSSION references. known genus The Arum visitors/pollinators. their with Alocasia, smnindbfr,frms ftear- the of most for before, mentioned As the represents below histogram The mznl nsieo hi ‘nqe’and ‘‘unique’’ their of spite in Amazingly stems ouetdwt 31 with documented most the is atooa Dieffenbachia Xanthosoma, ahhv 0rfrne dealing references 10 have each Arum spriual document- particularly is and with a great diversity of insects: bees, bee- anthesis (e.g. pollen release) hours or a tles and flies (Bown, 2000). Even verte- few days later. brates, e.g. hummingbirds, have recently been mentioned (Kraemer & Schmitt, Three Types of Interactions 1999). As in all pollination systems, pollinators Inflorescence Evolutionary Trends visit flowers looking for a resource. It can be an alimentary resource (stigmatic fluid, Aroids have a characteristic inflores- pollen, floral parts, . . .) or a ‘‘reproductive cence made up of a spadix and a spathe, resource’’ (mating, laying site). in and all the inflorescence morphologies ob- return have developed adaptations to served in aroids can be seen as variations transform this behaviour (e.g. around this same theme (Bown, 2000). resource seeking) into a ‘‘pollinating act’’, From a pollination point of view, there are and thus ensure their reproduction. One two main inflorescence evolutions: interesting point is that not a single species 1) Evolution of unisexual flowers that is known to offer nectar as a reward has allowed the secondary development (Schwerdtfeger et al., 2002). Nectar, a of sterile flowers and then floral function high-energy supply, is not produced in Ar- specialisation (barrier, odor emission, ther- aceae and the stigmatic exudate (e.g. flu- mogenesis, food-reward, . . .). On the con- id) is supposedly a poor substitute from trary, in species with bisexual flowers, all the energetic point of view. Stigmate ex- the flowers have the same functions and udates may have been the first type of participate to all the ‘‘functions’’ from pol- nectar in flowering and occur in linator attraction to fertilisation many primitive flowering plants (Lloyd & through pollen dispersion. Wells, 1992). 2) Development of an enclosing spathe From the flower-pollinator interaction with secondary appearance of a constric- point of view, three situations exist in Ar- tion allowing the capture of insects in con- aceae. tact with the flowers. Consequently, the 1) ‘‘classical’’ flower—pollinator inter- relationship with the pollinators has action. The inflorescence doesn’t play any evolved from a ‘‘two-phases’’ interaction role in the pollinator reproduction. The to a ‘‘one-phase’’ interaction (Meeuse, pollinator visits the inflorescence looking 1978). In species lacking a floral chamber, for a resource (stigmatic fluid, pollen) pollinators cannot ‘‘stay’’ or be captured achieving thereby the pollination (e.g. near the flowers, they have to come and Symplocarpus, Calla, some Anthurium). depart repeatedly. Thus one attraction 2) The inflorescence becomes part of phase is necessary during the female the reproductive cycle of the pollinator. phase to attract pollen vectors and achieve Insects visit Araceae inflorescences not ovule fertilisation. A second attraction only for food rewards (nectar, pollen or phase is needed later during the male floral tissue) but also to meet sexual con- phase to ensure pollen collection and its geners, achieve copulations and some- dispersion. times lay their eggs (e.g. Philodendron, In aroids with exposed inflorescences Dieffenbachia, Peltandra, Colocasia). and long anthesis, the inflorescence 3) The inflorescence dupes the polli- ‘‘needs’’ to attract (e.g. with odour and nators by mimicking the laying site of the heat production) the pollinators every day pollinator (faeces, mushrooms, dead ani- of anthesis. In species with a floral cham- mal, . . .). Hence, the insects visit the inflo- ber, once pollinators have been attracted rescence in order to complete their repro- during the female stage, they are then kept ductive cycle. Through this deceptive at- within the floral chamber by trap mecha- traction, the insects achieve pollination but nisms or rewards (shelter from light, food, without actually receiving any reward (e.g. sexual partners, . . .) until the end of the Arum, Helicodiceros). One can think of a fourth situation in perimental results, to publish them in or- which the pollinator dupes the flower, tak- der to document a domain still poorly ing the resource but not achieving polli- known for Araceae. nation. A well-known example is nectar thieving by bees or bumblebees and even ACKNOWLEDGMENTS birds. Such interaction exists in about 14 I wish to thank Derek Burch and Thom- families although in some instances as Croat for constructive comments and pollination is after all achieved (reviewed corrections on the manuscript. in Maloof & Inouye, 2000). 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