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September 2005

A new species of Pharaxonotha (Coleoptera: ), probable pollinator of the endangered Cuban cycad, Microcycas calocoma (Zamiaceae)

Ramiro Chavez Museo Nacional de Historia Natural, Cuba

Julio A. Genaro York University, Department of Biology, Toronto ON, M3J 1P3, Canada

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Chavez, Ramiro and Genaro, Julio A., "A new species of Pharaxonotha (Coleoptera: Erotylidae), probable pollinator of the endangered Cuban cycad, Microcycas calocoma (Zamiaceae)" (2005). Insecta Mundi. 70. https://digitalcommons.unl.edu/insectamundi/70

This Article is brought to you for free and open access by the Center for Systematic Entomology, Gainesville, Florida at DigitalCommons@University of Nebraska - Lincoln. It has been accepted for inclusion in Insecta Mundi by an authorized administrator of DigitalCommons@University of Nebraska - Lincoln. INSECTA MUNDI, Vol. 19, No. 3, September, 2005 143

A new species of Pharaxonotha (Coleoptera: Erotylidae), probable pollinator of the endangered Cuban cycad, Microcycas calocoma (Zamiaceae)

Ramiro Chaves Museo Nacional de Historia Natural Obispo no. 61, Habana Vieja 10100, Cuba [email protected]

Julio A. Genaro York University, Department of Biology 4700 Keele St., Toronto ON, M3J 1P3, Canada [email protected]

Abstract: The new species Pharaxonotha esperanzae (Coleoptera: Cucujoidea: Erotylidae: Pharaxonothinae) is described. It feeds on the pollen of the endangered Cuban cycad Microcycas calocoma (Miq.) A. DC. and breeds in its male cones. Its potential role as a pollinator of Microcycas is discussed.

Resumen: Se describe la especie nueva: Pharaxonotha esperanzae (Coleoptera: Cucujoidea: Erotylidae: Pharaxonothinae). Ésta se alimenta del polen del cicadal cubano amenazado Microcycas calocoma (Miq.) A.DC y se reproduce en su cono masculino. Se discute su papel potencial como agente polinizador.

Key words: Pharaxonotha, Erotylidae, Languriidae, new species, Cycadales, pollination.

Palabras claves: Pharaxonotha, Erotylidae, Languriidae, especie nueva, Cycadales, polinización.

Introduction et al. 1995, Tang et al. 1999, Wilson 2002, Hall et al. 2004, Tang 2004). Cycadales (cycads) are the oldest group of in- Within Erotylidae (which now includes taxa sect-pollinated plants (Norstog and Nicholls 1997). previously in Languriidae) is the genus Pharaxono- Many have developed mutualistic interac- tha Reitter 1875 (= Planismus Casey 1890) belong- tions that can be host-specific (Donaldson 2003). ing to the subfamily Pharaxonothinae, which is one There are different hypotheses on the antiquity of the most primitive in the family (Leschen 2003). and original association (whether with gymno- Of all genera known to pollinate cycads, sperms or angiosperms) of the insects known to Pharaxonotha is the most important based on the pollinate cycads. Some of these interactions appar- fact it has been associated with three of eleven ently have recent origins (Crowson 1991, Oberpri- cycad genera: Zamia, , and Ceratozamia (Tang eler 1995a, 1995b). Nevertheless, the fossil record, 1987, Vovides 1991). This genus of little was phylogenetic studies and biogeography support the taxonomically redefined by Sen Gupta and Crow- existence of Coleoptera feeding on gymnosperms son (1971). At present, there are 11 described since the Jurassic, and those associated with cones species for the world (Leschen 2003). Pakaluk (1988) are considered precursors of important families of described two new species from Costa Rica and phytophagous beetles (Crowson 1991, Farrell 1998). presented a key including the four New World Primitive beetles of the family Erotylidae are among species. These are: P. kirschii Reitter (= Thalisella the oldest extant pollinators of cycads. Their bioge- conradti Gorham), P. floridana (Casey) [= P. zami- ography supports the hypothesis that they pollinat- ae Blake], P. clarkorum Pakaluk, and P. confusa ed cycads before the Cretaceous diversification of Pakaluk. angiosperms (Hall et al. 2004, Tang 2004). Erotyl- Pharaxonotha kirschii, the type species, is na- ids are known to pollinate species in six of the tive to Mexico and Central America, but as a stored eleven cycad genera, but only eight of these genera product pest, it now has a much wider distribution had been surveyed up to now (Tang 1987, Vovides that includes North America, South America and 1991, Ornduff 1991, Forster et al. 1994, Donaldson Europe (Pakaluk 1988). Pharaxonotha floridana, 144 Volume 19, No. 3, September, 2005, INSECTA MUNDI from the southeastern United States (Leschen and Materials and Methods Skelley 2002), is hosted by Zamia integrifolia [= Z. floridana (Jones 1993)] and was the first erotylid Both adult beetles and larvae were collected cited as a mutualistic pollinator of a cycad (Tang from male cones of M. calocoma, in an area of 1987). Pharaxonotha clarkorum and P. confusa, Parque Nacional Viñales, Viñales Township, Pinar from Costa Rica, feed on pollen and breed in male del Río Province, Cuba. This is a well-preserved cones as P. floridana, but in Zamia skinneri and Z. area where pollination and seed production occurs fairchildeana, respectively (Pakaluk 1988). annually. There are Pharaxonotha species yet to be de- Measurements were made with a stereo micro- scribed. All of which are considered mutualistic scope from 25 adult Pharaxonotha esperanzae sp. pollinators and are apparently host specific with nov., mounted or taken from alcohol: total body other New World Zamiaceae of the genera Zamia, length from anterior head edge to elytral apex; Dioon and Ceratozamia (Vovides 1991, Norstog et maximum body width; pronotum, length along its al. 1992, Cruz 1999, Pérez-Farrera et al. 2000, midline from anterior to posterior edge, and at its González 2004, P. Skelley and M. Thomas, pers. widest point; elytral length from base adjacent to comm. 2004). The genera Zamia and Dioon also scutellum to its apical edge; head width; dorsal have mutualistic pollinators among curculionoids interocular distance; and ventral interocular dis- in the family Belidae (Tang 1987, Vovides 1991, tance (following Pakaluk 1988). Other measure- Norstog et al. 1992). The New World Zamiaceae ments taken were antennal length and width of genera: Chigua (endemic from Colombia) and Mi- antennomeres VIII and IX. crocycas (monotypical endemic from Cuba) have no Genitalia were studied from five specimens. previously reported insect pollinator (Tang 2004). Genitalia were dissected, cleared 1 hour in 10% Prior to this study, there were no reported mem- KOH and preserved in 70% alcohol or slide mount- bers of Pharaxonotha from the West Indies. ed with euparol. Two specimens were dissected to The endemic Cuban cycad, Microcycas caloco- study mouthparts and wing venation. Drawings ma (Miq.) A.DC., is a threatened plant listed as were made from photographs taken by digital cam- Critically Endangered (CR), according to IUCN era through a microscope, and corrected or detailed criteria (Peña et al. 1998, Stevenson et al. 2003). A after additional observations were made. very important factor contributing to this status is its scarce reproduction, fundamentally due to lim- ited pollination (Cendrero 1940, Foster and Ro- dríguez 1942, Peña et al. 1997a, 1997b, Vovides et Order Coleoptera: Superfamily Cucujoidea al. 1997, Peña et al. 1998). This has been attributed Family Erotylidae: Subfamily Pharaxonothinae to the supposed extirpation or population decline of the yet unknown pollinator in most of its localities Pharaxonotha esperanzae, n. sp. (Norstog et al. 1986, Peña et al. 1997a and b; Vovides et al. 1997). Nevertheless, some preserved Description. There were no observable external localities showed a habitual seed production (Risco sexual dimorphism. Body oblong, oval (Fig. 1B), et al. 1984, Peña et al. 1988, Peña et al. 1997a), length 3.49-4.21mm (x =3.91mm), 2.50-3.06 times which proved the existence of a pollinator. It was maximum width (Table 1), and approximately 4.0 believed the unknown pollinator could be a curcu- times maximum body height. Body finely pubes- lionoid of the family Belidae. The discovery and cent, cuticle between punctures on frons and prono- identification of this pollinator was an urgent pri- tum minutely granulate, dull; general colour red- ority for the development of a conservation strategy dish brown, elytra generally uniform in colour, (Vovides et al. 1997). some structures with dark stains of variable inten- The objective of this work is to present the first sity and form, generally without fixed pattern. record for the genus Pharaxonotha in Cuba, de- Head: Eyes prominent, coarsely faceted (9 facets in scribe Pharaxonontha esperanzae and discuss the longitudinal axis), wide ventrally in relation to potential function of this species as the pollinator of dorsal width (Figs. 1B, 1C, and Table 1), with short, Microcycas. inconspicuous setae of variable density. Antennae relatively short, length less than pronotal width and slightly more than head width (Table 1); anten- nomere relative lengths approximately INSECTA MUNDI, Vol. 19, No. 3, September, 2005 145

Fig. 1. Pharaxonotha esperanzae, n. sp. A) Head, dorsal view; B) Habitus, dorsal view; C) Habitus, ventral view, without left legs; D) Coarse and fine punctations on elytra; E) Posterior wing; F) Posterior wing with anal cell; G) Left protibia with tarsus, posterior view; H) Aedeagus, lateral view; I) Aedeagus, ventral view; J) Ovipositor, ventral view; K) Larva dorsal view. 146 Volume 19, No. 3, September, 2005, INSECTA MUNDI

Table 1. Body sizes and size relations, from 25 adult specimens of Pharaxonotha esperanzae, n. sp. x= medial value, SD=Standard deviation. Sizes (mm) x S.D. Rank Total length 3.91 0.22 3.49-4.21 Elytral length 2.61 0.12 2.38-2.82 Pronotal length 0.79 0.05 0.64-0.87 Maximum width 1.400.061.27-1.51 Pronotal width 1.12 0.05 1.03-1.23 Head width 0.830.030.77-0.87 Size relations Maximum length / width 2.79 0.12 2.50-3.06 Head width / dorsal interocular distance 1.53 0.04 1.46-1.64 Head width / ventral interocular distance 2.17 0.08 2.05-2.33 Dorsal / ventral interocular distances 1.42 0.05 1.25-1.50 Antennal length/ head width 1.11 0.05 1.02-1.19 Antennal length / pronotal width 0.83 0.04 0.77-0.92 IX/VIII antennomere widths 1.28 0.09 1.12-1.67 Pronotal length / width 0.70 0.03 0.59-0.76 Elytral length / pronotal length 3.33 0.17 3.00-3.81

18:13:13:8:9:9:9:9:16:15:17, antennal club slightly occiput-gular sutures; occiput-gular sutures end wider than preceding segments, club three seg- anteriorly in elongated tentorial pits. Thorax. mented and oval in cross section. Mandibles large, Pronotum transverse (Table 1), convex and nearly at rest prominent in front of and laterally to labrum oval; anterior angles not developed, curved ven- (Fig. 1A), right mandible with two apical teeth, left trally, not visible dorasally; posterior angles round- with two or three teeth. Terminal maxillary pal- ed; punctures similar to larger punctures on frons pomere visible in dorsal view projecting from under (Fig. 2); base with pit on each side of midline, each retracted mandible by length or more of antennal with a short groove extending onto disc. Elytra scape (Fig. 1A). Apex of labial palp project slightly length 3.00 - 3.81 times pronotal length (Table 1); forward of mandible, and visible from above. Men- lacking marginal line at base; with 10 weakly tum trapezoid, almost triangular, fine and densely impressed striae, represented by imperfectly aligned punctured, without lateral pockets; submentum no rows of punctures slightly coarser than on frons defined. Supraocular striae and transverse line of and separated 1 to 3 diameters; scutellary striole vertex present (Figs. 1A and 2); punctation on frons with 8 – 12 punctures; interval punctures fine coarse and separated about 1 diameter near trans- (diameters approximately 1/3 of strial punctures), verse line on vertex, becoming fine and sparse punctures disorganized in disposition, frequently toward clypeus; clypeus punctures become finer with two at the same level (Fig. 1D); all elytral and sparser toward apical margin where they dis- punctures with a short seta. Hind wings as in Fig. appear; frontoclypeal suture obsolete at middle; 1E, anal cell absent or, if present, small (Fig. 1F, labrum lacking punctures. Occiput with two strid- observed in only one wing of one specimen). Pros- ulatory files, dorsal punctures nearly as coarse and ternum finely punctured, punctures denser on pro- dense as in neighbouring frons, punctures extend cess between coxae. Procoxal cavities slightly open ventrally as wedges. Eyes edged with ridge ventral- externally. Mesosternum coarsely and densely punc- ly, more prominent on posterior side. Pregula dense- tured. Mesepimeron turns inward slightly between ly punctured, somewhat limited by ventral-lateral mesosternum and metasternum. Punctures on ocular ridges. Transverse gular ridge or groove metasternum finer and sparser than on proster- absent, but with transverse unpunctured band that num. Protibia (Figs. 1G and 2) slightly expanded defines posterior limits of pregula and postocular and angled to the external apex, angle obtuse. ridges, band narrows laterally and is limited poste- Tarsomeres I-IV slightly elongated, conical shape, ro-laterally by punctured wedges of occiput; poste- I longer than II, II length almost equal to III, IV rior gula smooth, shiny, with very fine transverse half length of II, V slightly longer than I+II togeth- striations, narrower anteriorly, laterally limited by er. Abdomen. Punctures on ventrites fine and INSECTA MUNDI, Vol. 19, No. 3, September, 2005 147

Fig. 2. Pharaxonotha esperanzae, n. sp. Scanning Electron Microscope photograph showing dorsal view of head, with partial pronotum and protibiae. sparse as on metasternum. Aedeagus as in Figs. 1H male cone of Microcycas calocoma, (deposited in and 1I. Ovipositor as in Fig. 1J. Museo Nacional de Historia Natural de Cuba [MNHNCu]). Paratypes. (24 specimens) same Larva. (Fig. 1K) Maximum observed length 7.5 locality as holotype: 20.viii.2002, coll. R. Chaves, mm. Head pale brown, body segments yellowish, (N=6: 5 in MNHNCu [sexed 1 male] and 1 in except dorsal bands of dark brown granules. Gran- Colección Zoológica del Instituto de Ecología y ules separated by approximately one diameter. Sistemática de Cuba [CZACC]); 14.viii.2003, coll. R. Medial line without granules is weakly defined, Chaves (N=2, in MNHNCu); 16.viii.2003, coll. R. present on all thoracic and abdominal segments Chaves (N=1, in MNHNCu); 19.viii.2003, coll. R. except last one. Chaves (N=5: 4 in MNHNCu [sexed 1 male] and 1 in CZACC); 3.ix.2003, colls. R. Chaves and J. A. Diagnosis. Pharaxonotha esperanzae n .sp. differs Genaro (N=10: 7 in MNHNCu [sexed 6 females and from other New World species of Pharaxonotha by 1 male], 1 in CZACC and 2 in Florida State Collec- lacking the basal margin on the elytra and possess- tion of in Gainesville [FSCA]). ing supraocular striae. All presently described New World Pharaxonotha have a distinct basal margin Etymology: The specific name was chosen to ho- on the elytra and lack supraocular striae. nour Prof. Esperanza Peña García, outstanding Cuban researcher in plant conservation, especially Types. Holotype. Male. CUBA, provincia Pinar in Microcycas calocoma. The use of her Spanish del Río, municipio Viñales, Parque Nacional Viñales, name reminds us that the discovery of this species 3.ix.2003, collectors R. Chaves and J. A. Genaro, in 148 Volume 19, No. 3, September, 2005, INSECTA MUNDI gives hopes of achieving successful conservation of rhynchinae is the responsible for the exit holes in the plant. the microsporophylls that were observed in some old male cones as mentioned by Vovides and collab- Comments: Adult beetles were found feeding on orators (1997). pollen, inside 6 dehiscent male cones of M. caloco- Members of Pharaxonotha are the most fre- ma. There were 1 to 13 adult beetles per cone. Five quently observed pollinators of New World Zami- cones were just beginning to shed pollen. The sixth aceae (Tang 1987, Pakaluk 1988, Vovides 1991, cone had already shed its pollen and its microsporo- Norstog et al. 1992, Cruz 1999, Pérez-Farrera et al. phylls were starting to rot. This cone produced five 2000, González 2004). Given that they are known adults and more than 250 larvae of different sizes. as pollinators in so many New World cycads, it is These were feeding on pollen, and larger instars probable that they are the pollinators for cycads could also be found feeding on the parenchyma where no pollinator is known. To date, P. esperan- inside the microsporophylls. This represents the zae has been the only flying insect found repeatedly first record for Pharaxonotha from Cuba and West on male cones of Microcycas at the optimal moment Indies. for pollination in an area where natural pollination occurs every year. These facts strongly support the Discussion hypothesis that P. esperanzae is the pollinator for Microcycas. Before this work, Coleoptera had been found as Nevertheless, to have complete certainty about pollinators in three New World Zamiaceae (Zamia, its role as a pollinator, P. experanzae has yet to be Dioon and Ceratozamia). These are cucujoids of the found in female cones and demonstrated it carries family Erotylidae (Pharaxonotha, that pollinate all pollen to micropils in megasporangia. This could be three genera) and curculionoids of the family Bel- achieved by careful experiments, as those recently idae (Rhopalotria and Parallocorynus, that polli- done by some researchers of cycad-pollinator inter- nate Zamia and Dioon respectively) [Norstog et al. actions (Donaldson 1997, Mound and Terry 2001, 1986, Tang 1987, Vovides 1991, Norstog et al. 1992, Terry 2001, Wilson 2002, Hall et al. 2004). Further Cruz 1999, Pérez-Farrera et al. 2000]. Considering studies are required to discover if other species are that members of a clade can share morphological involved in cycad pollination in this area or in other characters, ecological and behavioural preferenc- localities with seed production. es, Vovides and his collaborators (Vovides et al. The discovery of P. esperanzae is an important 1997, Sosa and Ogata 1998) extrapolated the polli- step forward in identifying factors influencing the nation type of Microcycas, and suggested that its natural reproduction of Microcycas, and why some pollinators could be insects of the same genera or populations do not reproduce. The interaction be- closely related. This reasoning and typical weevil tween P. esperanzae and Microcycas needs to be traces found in old male cones, drove them to considered in any management plan to conserve believe that Mycrocycas’ pollinator would probably the remaining population of Microcycas. This inter- be one of the belid genera found in Zamia and Dioon action also adds support to the hypothesis that all (Vovides et al. 1997). cycads are insect-pollinated (Wilson 2002, Donald- In male cones that had passed the moment of son 2003). Furthermore, it stresses the major im- dehiscence, with rotting in progress, we found portance of primitive Erotylidae as pollinators of other insects that are still under study. None of cycads, and especially of Pharaxonotha, which is them is a member of the family Belidae. Only one now associated with four of eleven cycad genera. curculionoid species, not yet identified, of the sub- family Cryptorhynchinae (Curculionidae), was re- Acknowledgments peatedly present in those cones. Members of this subfamily have not been recorded as pollinators of We gratefully thank Gregoria, Jesús Serrano cycads, and the morphology of this species does not and their family for the extraordinary hospitality fit to the general appearance of those that do and help given; John Donaldson, John Hall, Rich- pollinate (R. Oberprieler, pers. comm. 2002). Based ard Leschen, Roy Osborne, Esperanza Peña, Miguel on this, the weevil probably does not serve as a A. Pérez-Farrera, Paul Skelley, William Tang, Irene pollinator (C. Lyall, pers. comm. 2002). Besides, Terry, Michael Thomas and Sebastián Vélez, for this weevil is not present in a proper moment to supplying pertinent literature; René Pérez for perform pollination. It is probable that this Crypto- mounting specimens; Antonio López, Esteban INSECTA MUNDI, Vol. 19, No. 3, September, 2005 149

Gutiérrez and Martín Luís for their guidance and Cycad Society of South Africa, Stellenbosch, help; and the Parque Nacional Viñales and the South Africa. Museo Nacional de Historia Natural for their insti- Farrell, B.D. 1998. “Inordinate fondness” ex- tutional support. We especially thank Michael Th- plained: Why are there so many beetles? Sci- omas and Paul Skelley who supplied specimens and ence 281: 555-559. images of other species for comparisons, and pol- Forster, P., P. Machin, L. Mound, and G. Wil- ished the English translation of this work; Paul son. 1994. Insects associated with reproduc- Skelley and Esperanza Peña for their guidance and tive structures of cycads in Queensland and critical comments on early manuscripts of this northeast New South Wales, Australia. Biotro- work; and Paul Skelley for supplying the Scanning pica 26: 217-222. Electron Microscope photograph. This research was Foster, A. S., and M. Rodríguez San Pedro. supported with partial financing by Juan R. Chaves, 1942. Field studies on Microcycas calocoma. WWF-Canadá, G. Lindsay Field Research Fund of Memorias de la Sociedad Cubana de Historia the California Academy of Sciences (granted to the Natural ‘Felipe Poey’ 16: 105-121. second author), and Museo Nacional de Historia González, F. 2004. Herbivoría en una gimnosper- Natural de Cuba. ma endémica de Colombia, Zamia encephalar- toides (Zamiacaeae) por parte de Eumaeus (Lep- References cited idoptera: Lycanidae). Revista de la Academia Colombiana de Ciencias Exactas, Físicas y Nat- Casey, T.L. 1890. Coleopterological notices II. An- urales 28: 233-243. nals of the New York Academy of Sciences 5: Hall, J. A., G. H. Walter, D. M. Bergstrom, and 307-504. P. Machin. 2004. Pollination ecology of the Cendrero, O. 1940. El Microcycas calocoma, palma Australian cycad Lepidozamia peroffskyana corcho, palma de corcho o corcho. Memorias de (Zamiaceae). Australian Journal of Botany 52: la Sociedad Cubana de Historia Natural ‘Felipe 333-343. Poey’ 8: 175-182. Jones, D. L. 1993. Cycads of the World. Reed, Crowson, R.A. 1991. The relations of Coleoptera Chatsworth, NSW, Australia. 312 pp. to Cycadales. pp. 13-28. In Zunino, M., X. Belles Leschen, R. A. B. 2003. Erotylidae (Insecta: Co- and M. Blas [eds.]. Advances in Coleopterology. leoptera: Cucujoidea): phylogeny and review Asociación Europea de Coleopterología, Barce- (Part 1). Fauna of New Zealand 47. Manaaki lona. Whenua Press, New Zealand. 108 pp. Cruz Rodríguez, J. de la. 1999. Polinización por Leschen, R. A. B., and P. E. Skelley. 2002. coleópteros en Ceratozamia norstogii Steven- Languriidae Wiedeman, 1823. pp. 343-347. In son (Zamiaceae) en el ejido de La Sombra de La Arnett, R.H. Jr., M. C. Thomas, P. E. Skelley Selva, Municipio Villaflores, Chiapas. Tesis de and J. Howard Frank [eds]. American beetles. Licenciatura (director M.A. Pérez-Farrera). Vol. 2. Polyphaga: Scarabaeoidea through Cur- Escuela de Biología, Universidad del Estado de culionoidea. CRC Press, Florida. Chiapas. 87 pp. Mound, L. A., and I. Terry. 2001. Thrips pollina- Donaldson, J. S. 1997. Is there a floral parasite tion of the central Australian cycad, Macrozamia mutualism in cycad pollination? The pollina- macdonnellii (Cycadales). International Jour- tion biology of Encephalartos villosus (Zami- nal of Plant Sciences 162: 147-154. aceae). American Journal of Botany 84: 1398- Norstog, K.J., and T. J. Nicholls. 1997. The 1406. biology of the cycads. Cornell Univ. Press, Ith- Donaldson, J. S. 2003. [ed.]. Cycads. Status Sur- aca. 363 pp. vey and Conservation Action Plan. IUCN/SSC Norstog, K.J., D. W. Stevenson, and K. J. Ni- Cycad Specialist Group. IUCN, Gland, Switzer- klas. 1986. The role of beetles in the pollination land and Cambridge, UK. ix + 86 pp. of Zamia furfuracea L. fil. (Zamiaceae). Biotro- Donaldson, J.S., I. Nänni, and J.D. Bösen- pica 18: 300-306. berg. 1995. The role of insects in the pollina- Norstog, K. J., P. K. S. Fawcett, and A. P. tion of Encephalartos cycadifolius. pp. 423-434. Vovides. 1992. pollination of two spe- In Vorster, P. [ed.]. Proceedings of the Third cies of Zamia: evolutionary and ecological con- International Conference on Cycad Biology. siderations. Palaeobotanist 41: 149-158. 150 Volume 19, No. 3, September, 2005, INSECTA MUNDI

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