DOCSLIB.ORG

Department of Biology Imperial College at Silwood Park Ascot, Berks, SL5 7PY, UK

Total Page：16

File Type：pdf, Size：1020Kb

Download full-text PDF Read full-text
Department of Biology Imperial College at Silwood Park Ascot, Berks, SL5 7PY, UK EVOLUTIONARY ECOLOGY OF FIGS AND FIG WASPS by DANIEL BEAN September 2002 A Thesis submitted for the Degree of Doctor of Philosophy of the University of London Department of Biology Imperial College at Silwood Park Ascot, Berks, SL5 7PY, UK 1 i \ 1 'JA rvw. ABSTRACT Figs (Ficus spp., Moraceae) and their pollinating fig wasps (Hymenoptera: Agaonidae) are obligate mutualists, figs provide a site for larval development whereas wasps disperse pollen and fertilise flowers. Figs are attacked by non-pollinating wasps which oviposit in fig inflorescences (syconia) without benefitting their hosts. This study analyses data from Indoaustralian figs, principally from section Malvathera, and their associated wasps. Male non-pollinating wasps fight to the death for matings and display considerable morphological variation. In Sycoscapter australis and Philotrypesis sp. disproportionately large mandibles are selected for in males, consistent with their role as weapons. Mandible size varies up to sevenfold, and can be either continuous or discrete. The frequency of serious or fatal fights among S. australis males is positively correlated with the ratio of the number of sexually active males to the number of females available for mating, a product of the rate of female emergence. Fights occur in Philotrypesis sp. between males that are less similar in mandible size than average, indicating that fights occur because large males can attack small males with impunity. Figs are in conflict with their pollinators over the fate of flowers. Flowers provide wasps with a site for larval development but figs produce viable seeds by protecting their flowers. Long term stability in the mutualism relies on the ability of figs to control wasp oviposition behaviour. Phlyogenetically controlled comparative analyses of Malvanthera figs and their associated wasps show that figs can reduce investment in by fostering in their pollen production active pollination wasps -a suite of morphological and behavioural traits that facilitate efficient pollen transmission. Active pollination may have arisen more than once in pollinating fig wasps. Figs can protect their flowers by reducing flower size. This results in smaller wasps, which are able to carry fewer eggs. I DECLARE THAT THIS THESIS IS ENTIRELY MY OWN WORK, EXCEPT FOR PART OF THE DATA ANALYSED IN CHAPTERS 4 AND 5, WHICH WERE COLLECTED BY JAMES COOK, STUART WEST AND E. ALLEN HERRE 2 TABLE OF CONTENTS Title Page 1 Abstract 2 Table of Contents 3 List of Figures 7 List of Tables 12 Chapter 1- Reproductive Conflict in the Fig and Fig Wasp Mutualism 14 Introduction 14 Fatal Fighting 15 Mechanisms of mate competition 15 Occurrence of fatal fighting 17 Game theory models 19 Kin selection models 20 Encounter rate and resource value models 22 Sex ratio models 24 Mutualistic Coevolution and Coadaptation 27 Definition and occurrence 27 Phylogenetic congruence between figs and wasps 29 Conflicts between figs and wasps 32 Flower protection mechanisms 34 Syconium protection mechanisms 36 Pollination mode and its impact on the mutualism 37 3 Non-pollinating Chapter 2- Male Mating Tactics and Lethal Combat in the 40 Fig Wasp Sycoscapter australis Abstract 40 Introduction 41 Background Biology 43 44 Methods and Analysis Data Collection 44 Dimorphism in S. australis males 45 46 Serious injury in S. australis males Emergence in S. australis males 47 Results 48 Population Data 48 Dimorphism in S. australis males 48 Serious injury in S. australis males 50 Emergence in S. australis males 51 Discussion 52 Morphology and mating tactics in S. australis 52 Factors affecting serious injury in S. australis 53 Emergence 54 Chapter 3- Victimisation and Morphology in Fighting Male Fig Wasps 57 Abstract 57 Introduction 58 Methods 61 Study species 61 Data collection 62 4 Allometry 63 Size matching of pairs of fighting males 65 Results 66 Allometry 66 Size matching of pairs of fighting males 68 Discussion 69 Morphological patterns in Philotrypesis sp. males 69 Fighting behaviour in Philotrypesis sp. males 71 Chapter 4- The Evolution and Maintenance of Active Pollination 74 Abstract 74 Introduction 76 Methods 78 Study taxon 78 Data collection 79 Phylogenetic analysis 80 Character evolution 81 Comparative analysis 81 Results 82 Phylogenetic analysis 82 Character evolution 82 Comparative analysis 84 Discussion 87 Phylogenetic analysis 87 Character evolution 88 Comparative analysis 90 5 Chapter 5- The Role Of The Fecundity Hypothesis in Protecting Fig Flowers 93 Abstract 93 Introduction 94 Methods 98 Study taxon 98 Data collection 98 Phylogenetic analysis 100 Comparative analysis 100 Do larger Malvathera figs produce more or larger female flowers? 101 Do smaller female flowers produce smaller pollinating wasps? 101 Results 103 Data collection 103 Do larger Malvathera figs produce more or larger female flowers? 104 Do smaller female flowers produce smaller pollinating wasps? 107 Discussion 109 Chapter 6- Conclusions 114 Introduction 114 Fatal Fighting in Non-pollinating Fig Wasps 114 Coevolution and Coadaptation between figs and their pollinating wasps 117 Acknowledgements 120 Bibliography 121 6 LIST OF FIGURES Chapter 1 Figure 1 Sycoscapter sp. from F. benjamina collected in Papua New Guinea, 19 a) dorsal view of head and thorax illustrating piercing mandibles, b) lateral view illustrating femoral spikes. Reproduced from Boucek (1988), not to scale. Chapter 2 Figure 1 Plot showing the mean wasp emergence per hour across all syconia 49 for P. froggatti (broken lines) and S. australis (solid lines) males (thick lines) and females (thin lines). Figure 2 Plot showing the regression slopes of the relationship between 50 mandible and head length in S. australis males for (a) values of head length less than 0.6mm, and (b) for value head length greater than 0.6mm. Figure 3 a) Plot showing data points and the regression slope of the 51 proportion of seriously injured S. australis males per syconium against the mean S. australis OSR per syconium. b) Plot showing data points and the regression slope of the proportion of seriously injured S. australis males per syconium against the mean S. australis male mandible size per syconium. Figure 4 a) Plot showing the proportion of emerged S. australis males per 52 syconium against the mean number of S. australis males per syconium with a fitted curve estimated from a linear model of male emergence - 0.72 - 0.08 x number of males + 0.002 x number of 7 malest. P value of squared term <0.005, d.f. = 1, adjusted r2 = 0.46. b) Plot showing the proportion of emerged S. australis males per syconium against the proportion of seriously injured S. australis males per syconium with a fitted curve estimated from a linear 3.55 model of male emergence - 0.56 - 2.55 x injury proportion + x injury proportion2. P value of squared term <0.001, d.f. = 1, adjusted r2 = 0.38. Chapter 3 Figure 1 Relationship between mandible length and head length in 67 Philotrypesis sp. males. Solid line represents a quadratic model of mandible vs head allometry. Dashed line indicates dimorphic model of mandible vs head allometry. Dotted lines indicate mandible and head values for the proposed switch point in the dimorphic model. Figure 2 Histogram showing the means for Philotrypesis sp. of the mean size 68 difference between pairs of fighting males per syconium (white bars) and the mean size difference between all pairs of males per syconium (grey bars) from 30 syconia, using mandible, head and hind tibia body measurements(t bars indicate standard errors). Chapter 4 Figure 1 50% majority rule tree based on Bayesian analysis of combined 28S 83 and ITS alignment for 14 Pleistodontes species. Analysis run for 3 million generations with GTR evolutionary model with gamma rates. Values indicate posterior probability support for adjacent node. White terminal boxes indicate active pollination, black terminal boxes indicate passive. 8 Figure 2 Pleistodontes phylogeny showing unordered character state 84 reconstruction of pollination mode, boxes by character names indicate character statesfor each species. White shading indicates active pollination, black shading indicates passive pollination, barred shading indicates equivocal ancestral character state. Black circles indicate nodes at which contrasts were calculated for flower sex ratio using CAIC (Purvis & Rambaut 1995). Figure 3 Pleistodontes phylogeny showing character state reconstruction of 85 pollination mode, represented as in Fig 2. Tree (a) illustrates the most parsimonious character reconstruction when active pollination is hard to acquire but easy to lose (scenario 1 in the methods), tree (b) indicates the most parsimonious character state reconstruction when active pollination is equally easy to acquire and lose (scenario 2 in the methods). Figure 4 Histogram comparing the mean flower sex ratios (FSR) for actively 86 and passively pollinated fig species from section Malvanthera. Bars indicate standard errors. Figure 5 Reconstruction of the evolution of pollination behaviour across 15 86 pollinating fig wasp genera using a maximum likelihood tree calculated by Machado et al. (2001). All genera are represented by one species except Ceratosolen, which is represented by four. Branch shading indicates pollination mode, white = active, black = passive, grey = absent, hatched = polymorphic. Figure reproduced from Machado et al. (2001). 9 Chapter 5 Figure 1 Variation in the size and shape of the ripe syconia of malvantheran 100 fig species. Syconia are drawn to scale. Seven species have essentially spherical syconia and are shown as such. The other six have syconia of varying shapesas depicted. Species are labelled as follows: a) F. obliqua, b) F. lilliputiana, c) F. brachypoda, d) F. rubiginosa, e) F. platypoda, f) F. macrophylla, g) F. hesperidiformis, h) F. crassipes, i) F. pleurocarpa, j) F. watkinsiana, k) F. xylosycia, 1) F. triradiata, m) F. destruens. Figure 2 50% majority rule tree based on Bayesian analysis of combined 28S 102 and ITS alignment for 14 Pleistodontes species. Analysis run for 3 million generations of GTR evolutionary model with gamma rates. Values indicate posterior probability support for adjacent node.
Load more

Recommended publications
  • Comparative Anatomy of the Fig Wall (Ficus, Moraceae)
    Botany Comparative anatomy of the fig wall (Ficus, Moraceae) Journal: Botany Manuscript ID cjb-2018-0192.R2 Manuscript Type: Article Date Submitted by the 12-Mar-2019 Author: Complete List of Authors: Fan, Kang-Yu; National Taiwan University, Institute of Ecology and Evolutionary Biology Bain, Anthony; national Sun yat-sen university, Department of biological sciences; National Taiwan University, Institute of Ecology and Evolutionary Biology Tzeng, Hsy-Yu; National Chung Hsing University, Department of Forestry Chiang, Yun-Peng;Draft National Taiwan University, Institute of Ecology and Evolutionary Biology Chou, Lien-Siang; National Taiwan University, Institute of Ecology and Evolutionary Biology Kuo-Huang, Ling-Long; National Taiwan University, Institute of Ecology and Evolutionary Biology Keyword: Comparative Anatomy, Ficus, Histology, Inflorescence Is the invited manuscript for consideration in a Special Not applicable (regular submission) Issue? : https://mc06.manuscriptcentral.com/botany-pubs Page 1 of 29 Botany Comparative anatomy of the fig wall (Ficus, Moraceae) Kang-Yu Fana, Anthony Baina,b *, Hsy-Yu Tzengc, Yun-Peng Chianga, Lien-Siang Choua, Ling-Long Kuo-Huanga a Institute of Ecology and Evolutionary Biology, College of Life Sciences, National Taiwan University, 1, Sec. 4, Roosevelt Road, Taipei, 10617, Taiwan b current address: Department of Biological Sciences, National Sun Yat-sen University, 70 Lien-Hai road, Kaohsiung, Taiwan.Draft c Department of Forestry, National Chung Hsing University, 145 Xingda Rd., South Dist., Taichung, 402, Taiwan. * Corresponding author: [email protected]; Tel: +886-75252000-3617; Fax: +886-75253609. 1 https://mc06.manuscriptcentral.com/botany-pubs Botany Page 2 of 29 Abstract The genus Ficus is unique by its closed inflorescence (fig) holding all flowers inside its cavity, which is isolated from the outside world by a fleshy barrier: the fig wall.
    [Show full text]
  • Ficus Cf. Platypoda Port Jackson Fig Moraceae
    Ficus cf. platypoda Port jackson fig Moraceae Forest Starr, Kim Starr, and Lloyd Loope United States Geological Survey--Biological Resources Division Haleakala Field Station, Maui, Hawai'i January, 2003 OVERVIEW There are no Ficus species native to Hawai'i. F. cf. platypoda is one of about 60 species of Ficus that is cultivated in Hawai'i (Wagner et al. 1999). About 39,000 F. cf. platypoda trees were planted in the state of Hawai'i during the 1920's and 1930's as a forestry tree (Skolmen 1960). On Maui, F.cf. platypoda were planted in plantations along the Hana Hwy. from Ha'iku to Hana and in Fleming Arboretum on West Maui. The pollinator wasp for F. cf. platypoda, Pleistondontes imperialis Saunders, was introduced to Hawai'i in 1922 (Wagner et al. 1999) to facilitate the spread of this tree species as each Ficus species needs a specific pollinating wasp in order to reproduce and spread (Ramirez 1970). As a result, F. cf. platypoda is reproducing sexually in Hawai'i today. It was first reported by Nagata (1995) under the name F. rubiginosa Desf as naturalized on O'ahu. It was then later reported as naturalized on both West and East Maui (Wagner et al. 1999, Oppenheimer and Bartlett 2000). Two other Ficus species that have had their associated pollinator wasps introduced are also spreading on Maui. These include F. microcarpa and F. macrophylla (Wagner et al. 1999, Oppenheimer and Bartlett 2000). All three species invade both disturbed and native ecosystems. F. cf. platypoda is capable of germinating in native host trees, such as koa (Acacia koa) and ohia (Metrosideros polymorpha), growing as epiphytes, sending down aerial roots, and eventually destroying the host tree.
    [Show full text]
  • Ficus Burkei
    International Scholarly Research Network ISRN Zoology Volume 2012, Article ID 908560, 6 pages doi:10.5402/2012/908560 Research Article Spatial Stratification of Internally and Externally Non-Pollinating Fig Wasps and Their Effects on Pollinator and Seed Abundance in Ficus burkei Sarah Al-Beidh,1 Derek W. Dunn,2 and James M. Cook2 1 Division of Biology, Imperial College London, Ascot, Berkshire SL5 7PY, UK 2 School of Biological Sciences, University of Reading, Reading, Berkshire RG6 6AS, UK Correspondence should be addressed to James M. Cook, [email protected] Received 30 November 2011; Accepted 19 December 2011 Academic Editors: M. Kuntner and S. Van Nouhuys Copyright © 2012 Sarah Al-Beidh et al. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Fig trees (Ficus spp.) are pollinated by tiny wasps that enter their enclosed inflorescences (syconia). The wasp larvae also consume some fig ovules, which negatively affects seed production. Within syconia, pollinator larvae mature mostly in the inner ovules whereas seeds develop mostly in outer ovules—a stratification pattern that enables mutualism persistence. Pollinators may prefer inner ovules because they provide enemy-free space from externally ovipositing parasitic wasps. In some Australasian Ficus, this results in spatial segregation of pollinator and parasite offspring within syconia, with parasites occurring in shorter ovules than pollinators. Australian figs lack non-pollinating fig wasps (NPFW) that enter syconia to oviposit, but these occur in Africa and Asia, and may affect mutualist reproduction via parasitism or seed predation.
    [Show full text]
  • Zoologische Mededelingen
    MINISTERIE VAN ONDERWIJS KUNSTEN EN WETENSCHAPPEN ZOOLOGISCHE MEDEDELINGEN UITGEGEVEN DOOR HET RIJKSMUSEUM VAN NATUURLIJKE HISTORIE TE LEIDEN DEEL XXXVIII, No. 19 18 november 1963 INDO-MALAYAN AND PAPUAN FIG WASPS (HYMENOPTERA, CHALCIDOIDEA) 2. THE GENUS PLEISTODONTES SAUNDERS (AGAONIDAE) by J. T. WIEBES Rijksmuseum van Natuurlijke Historie, Leiden Until now, the genus Pleistodontes Saunders is known from the Australian continent and from Lord Howe Island. Two species were introduced into the Hawaiian Islands. Samples of fig wasps from New Guinea and from the Solomon Islands were sent to me by Mr. E. J. H. Corner. They proved to consist of new species of Pleistodontes, which are described below. In addition, some new records of Australian species, mostly taken from the collection of the Ha• waiian Sugar Planters' Association ("H.S.P.A."), Honolulu, are given, and the type species of the genus is redescribed. The records and "descriptions" by Girault are not considered in the present paper, as the typical material is being studied by Mr. E. F. Riek. A discussion on the host records follows the descriptions of the species. Pleistodontes blandus spec. nov. Material. — Eight immature 9, 14 $, ex Ficus glandifera Summerh. (det. E. J. H. Corner), Solomon Is., leg. Kajewski, no. 3494; coll. Museum Leiden, no. 438; holotype, S, slide no. 438b, allotype, 9, slide 438a, paratypes, 9 $, slides 438c, d. Description. — Male. Head (fig. 10) distinctly longer than its maximum width, and nearly twice as long as wide anteriorly; with the usual pubescence next to the antennal groove and behind the hypostomal margin. Eyes large. Mandible, fig.
    [Show full text]
  • Investigations Into Stability in the Fig/Fig-Wasp Mutualism
    Investigations into stability in the fig/fig-wasp mutualism Sarah Al-Beidh A thesis submitted for the degree of Doctor of Philosophy of Imperial College London. Declaration I hereby declare that this submission is my own work, or if not, it is clearly stated and fully acknowledged in the text. Sarah Al-Beidh 2 Abstract Fig trees (Ficus, Moraceae) and their pollinating wasps (Chalcidoidea, Agaonidae) are involved in an obligate mutualism where each partner relies on the other in order to reproduce: the pollinating fig wasps are a fig tree’s only pollen disperser whilst the fig trees provide the wasps with places in which to lay their eggs. Mutualistic interactions are, however, ultimately genetically selfish and as such, are often rife with conflict. Fig trees are either monoecious, where wasps and seeds develop together within fig fruit (syconia), or dioecious, where wasps and seeds develop separately. In interactions between monoecious fig trees and their pollinating wasps, there are conflicts of interest over the relative allocation of fig flowers to wasp and seed development. Although fig trees reap the rewards associated with wasp and seed production (through pollen and seed dispersal respectively), pollinators only benefit directly from flowers that nurture the development of wasp larvae, and increase their fitness by attempting to oviposit in as many ovules as possible. If successful, this oviposition strategy would eventually destroy the mutualism; however, the interaction has lasted for over 60 million years suggesting that mechanisms must be in place to limit wasp oviposition. This thesis addresses a number of factors to elucidate how stability may be achieved in monoecious fig systems.
    [Show full text]
  • Pollinated by Pleistodontes Imperialis. (Ficus Carica); Most
    The Port Jackson, or Rusty Fig, is yet another handsome Australian native fig, common in eastern Australia, from near the NSW/Victorian border in the south, to the very tip of Cape York in far north Queensland. This fig can grow on its own roots, but often starts life as a lithophyte (growing on rock) or as a hemiparasite (growing on other plants). Port Jackson Fig, as the name suggests, can be seen on many rocky headlands of Sydney Harbour. It commonly occurs on the margins of rainforests, in vine thickets and in riverine vegetation. In appearance, it’s not unlike a smaller version of the Moreton Bay Fig, Ficus macrophylla and, like the Moreton Bay Fig, is not only popular for planting in urban parks and gardens, but in miniature, as a bonsai plant. Most fig species are pollinated by just one species of fig wasp. In this case, the Port Jackson Fig is pollinated by Pleistodontes imperialis. There are perhaps 750 species of Ficus worldwide, including the edible fig (Ficus carica); most occur in tropical and sub-tropical regions, Typical habitat for Ficus with some species rubiginosa – on rocky sandstone headlands of Sydney Harbour. (Ficus carica, the edible fig, for example) occur in temperate parts of the world. Figs (Ficus spp.) belong to the plant family Moraceae, which also includes Mulberries (Morus spp.), Breadfruit and Jackfruit (Artocarpus spp.). Think of a mulberry, and imagine it turned inside out. This might perhaps bear some resemblance to a fig. Ficus rubiginosa growing on a sandstone platform adjoining mangroves. Branches of one can be seen in the foreground, a larger one at the rear.
    [Show full text]
  • The Dark Spreading Crowns of Moreton Bay Fig Trees Are a Glorious
    The dark spreading crowns of Moreton Bay Fig Trees are a glorious feature of many Sydney parklands, for example Mrs Macquaries Chair, Hyde Park, the Domain and Moore Park. In their native habitat, these handsome trees can be found in eastern Australia, from south-eastern Queensland to Wollongong in southern NSW. Moreton Bay Figs, and many other fig species, can start life as seed that germinates in the branches of another tree, sending roots down to the ground. These roots and the trunk then thicken, enveloping the original host and eventually killing it. Figs with this growth habit are often referred to as banyans although in Australia we usually call them strangler figs. The fruit is edible, but not very palatable. However, they are particularly interesting and both structure and reproductive strategies are complex! Unlike most flowers with which we are familiar, fig flowers, both male and female, are extremely small and are produced on the INSIDE of the fruit which is technically referred to as a synconium. The flowers are pollinated by tiny fig wasps (Pleistodontes froggatti), with which the fig trees share an obligate mutual relationship. The figs depend on wasps for pollination, and the wasps can reproduce only in the female flowers. It had been thought that each fig species was pollinated by one specific wasp species but recent molecular studies have shown that some fig wasp species have been known to cheat, so the story isn’t quite as straight forward as originally envisaged. Pollination works like this: 1. Female wasps enter the fig and lay eggs in maturing female flowers.
    [Show full text]
  • Terrestrial Arthropod Surveys on Pagan Island, Northern Marianas
    Terrestrial Arthropod Surveys on Pagan Island, Northern Marianas Neal L. Evenhuis, Lucius G. Eldredge, Keith T. Arakaki, Darcy Oishi, Janis N. Garcia & William P. Haines Pacific Biological Survey, Bishop Museum, Honolulu, Hawaii 96817 Final Report November 2010 Prepared for: U.S. Fish and Wildlife Service, Pacific Islands Fish & Wildlife Office Honolulu, Hawaii Evenhuis et al. — Pagan Island Arthropod Survey 2 BISHOP MUSEUM The State Museum of Natural and Cultural History 1525 Bernice Street Honolulu, Hawai’i 96817–2704, USA Copyright© 2010 Bishop Museum All Rights Reserved Printed in the United States of America Contribution No. 2010-015 to the Pacific Biological Survey Evenhuis et al. — Pagan Island Arthropod Survey 3 TABLE OF CONTENTS Executive Summary ......................................................................................................... 5 Background ..................................................................................................................... 7 General History .............................................................................................................. 10 Previous Expeditions to Pagan Surveying Terrestrial Arthropods ................................ 12 Current Survey and List of Collecting Sites .................................................................. 18 Sampling Methods ......................................................................................................... 25 Survey Results ..............................................................................................................
    [Show full text]
  • Weiblen, G.D. 2002 How to Be a Fig Wasp. Ann. Rev. Entomol. 47:299
    25 Oct 2001 17:34 AR ar147-11.tex ar147-11.sgm ARv2(2001/05/10) P1: GJB Annu. Rev. Entomol. 2002. 47:299–330 Copyright c 2002 by Annual Reviews. All rights reserved ! HOW TO BE A FIG WASP George D. Weiblen University of Minnesota, Department of Plant Biology, St. Paul, Minnesota 55108; e-mail: [email protected] Key Words Agaonidae, coevolution, cospeciation, parasitism, pollination ■ Abstract In the two decades since Janzen described how to be a fig, more than 200 papers have appeared on fig wasps (Agaonidae) and their host plants (Ficus spp., Moraceae). Fig pollination is now widely regarded as a model system for the study of coevolved mutualism, and earlier reviews have focused on the evolution of resource conflicts between pollinating fig wasps, their hosts, and their parasites. Fig wasps have also been a focus of research on sex ratio evolution, the evolution of virulence, coevolu- tion, population genetics, host-parasitoid interactions, community ecology, historical biogeography, and conservation biology. This new synthesis of fig wasp research at- tempts to integrate recent contributions with the older literature and to promote research on diverse topics ranging from behavioral ecology to molecular evolution. CONTENTS INTRODUCING FIG WASPS ...........................................300 FIG WASP ECOLOGY .................................................302 Pollination Ecology ..................................................303 Host Specificity .....................................................304 Host Utilization .....................................................305
    [Show full text]
  • TAXON:Ficus Macrophylla Pers. SCORE:9.0 RATING:High Risk
    TAXON: Ficus macrophylla Pers. SCORE: 9.0 RATING: High Risk Taxon: Ficus macrophylla Pers. Family: Moraceae Common Name(s): Australian banyan Synonym(s): Ficus magnolioides Borzì black fig Moreton Bay fig Assessor: Chuck Chimera Status: Assessor Approved End Date: 15 Oct 2019 WRA Score: 9.0 Designation: H(Hawai'i) Rating: High Risk Keywords: Strangler Fig, Naturalized, Environmental Weed, Bird-Dispersed, Resprouts Qsn # Question Answer Option Answer 101 Is the species highly domesticated? y=-3, n=0 n 102 Has the species become naturalized where grown? 103 Does the species have weedy races? Species suited to tropical or subtropical climate(s) - If 201 island is primarily wet habitat, then substitute "wet (0-low; 1-intermediate; 2-high) (See Appendix 2) High tropical" for "tropical or subtropical" 202 Quality of climate match data (0-low; 1-intermediate; 2-high) (See Appendix 2) High 203 Broad climate suitability (environmental versatility) y=1, n=0 n Native or naturalized in regions with tropical or 204 y=1, n=0 y subtropical climates Does the species have a history of repeated introductions 205 y=-2, ?=-1, n=0 y outside its natural range? 301 Naturalized beyond native range y = 1*multiplier (see Appendix 2), n= question 205 y 302 Garden/amenity/disturbance weed n=0, y = 1*multiplier (see Appendix 2) n 303 Agricultural/forestry/horticultural weed n=0, y = 2*multiplier (see Appendix 2) n 304 Environmental weed n=0, y = 2*multiplier (see Appendix 2) y 305 Congeneric weed n=0, y = 1*multiplier (see Appendix 2) y 401 Produces spines, thorns or burrs y=1, n=0 n 402 Allelopathic 403 Parasitic y=1, n=0 n 404 Unpalatable to grazing animals y=1, n=-1 n 405 Toxic to animals y=1, n=0 n 406 Host for recognized pests and pathogens y=1, n=0 y 407 Causes allergies or is otherwise toxic to humans 408 Creates a fire hazard in natural ecosystems y=1, n=0 n 409 Is a shade tolerant plant at some stage of its life cycle y=1, n=0 y Creation Date: 15 Oct 2019 (Ficus macrophylla Pers.) Page 1 of 20 TAXON: Ficus macrophylla Pers.
    [Show full text]
  • Molecular Markers Reveal Reproductive Strategies of Non-Pollinating Fig
    Ecological Entomology (2017), DOI: 10.1111/een.12433 Molecular markers reveal reproductive strategies of non-pollinating fig wasps JAMES M. COOK,1,2 CAROLINE REUTER,1,3 JAMIE C. MOORE4 andSTUART A. WEST5 1School of Biological Sciences, University of Reading, Reading, U.K., 2Hawkesbury Institute for the Environment, Western Sydney University, Penrith, Australia, 3Wolfson Institute of Preventive Medicine, Queen Mary, University of London, London, U.K., 4Department of Social Statistics and Demography, University of Southampton, Southampton, U.K. and 5Department of Zoology, University of Oxford, Oxford, U.K. Abstract. 1. Fig wasps have proved extremely useful study organisms for testing how reproductive decisions evolve in response to population structure. In particular, they provide textbook examples of how natural selection can favour female-biased offspring sex ratios, lethal combat for mates and dimorphic mating strategies. 2. However, previous work has been challenged, because supposedly single species have been discovered to be a number of cryptic species. Consequently, new studies are required to determine population structure and reproductive decisions of individuals unambiguously assigned to species. 3. Microsatellites were used to determine species identity and reproductive patterns in three non-pollinating Sycoscapter species associated with the same fig species. Foundress number was typically one to five and most figs contained more than one Sycoscapter species. Foundresses produced very small clutches of about one to four offspring, but one foundress may lay eggs in several figs. 4. Overall, the data were a poor match to theoretical predictions of solitary male clutches and gregarious clutches with n − 1 females. However, sex ratios were male-biased in solitary clutches and female-biased in gregarious ones.
    [Show full text]
  • PROCEEDINGS of the HAWAIIAN SOCIETY for 1975
    PROCEEDINGS of the HAWAIIAN ENTOMOLOGICAL SOCIETY for 1975 VOL. XXII NO. 3 December 1977 Information for Contributors Manuscripts for publication, proof, and other editorial matters should be addressed to: Editor: Hawaiian Entomological Society c/o Department of Entomology University of Hawaii 3050 Maile Way. Honolulu, Hawaii 96822 Manuscripts should not exceed 40 typewritten pages, including illustrations (approximately 20 printed pages). Longer manuscripts may be rejected on the basis of length, or be subject to additional page charges. Typing — Manuscripts must be typewritten on one side of white bond paper, 8-1/2 x II inches. Double space all text, including tables, footnotes, and reference lists. Margins should be a minimum of one inch. Underscore only where italics are intended in body of text, not in headings. Geographical names, authors names, and names of plants and animals should be spelled out in lull. Except for the first time they are used, scientific names of organisms may be abbreviated by using the first letter of the generic name plus the full specific name. Submit original typescript and one copy. Pages should be numbered consecutively. Place footnotes at the bottom of the manuscript page on which they appear, with a dividing line. Place tables separately, not more than one table per manuscript page, at end of manuscript. Make a circled notation in margin of manuscript at approximate location where placement of a table is desired. Use only horizontal lines in tables. Illustrations — Illustrations should be planned to fit the type page of 4-1/2x 7 inches, with appropriate space allowed for captions.
    [Show full text]