Molecular Phylogenies of Fig Wasps: Partial Cocladogenesis of Pollinators and Parasites Carlos Lopez-Vaamonde,* Jean Yves Rasplus,† George D

Total Page:16

File Type:pdf, Size:1020Kb

Molecular Phylogenies of Fig Wasps: Partial Cocladogenesis of Pollinators and Parasites Carlos Lopez-Vaamonde,* Jean Yves Rasplus,† George D Molecular Phylogenetics and Evolution Vol. 21, No. 1, October, pp. 55–71, 2001 doi:10.1006/mpev.2001.0993, available online at http://www.idealibrary.com on Molecular Phylogenies of Fig Wasps: Partial Cocladogenesis of Pollinators and Parasites Carlos Lopez-Vaamonde,* Jean Yves Rasplus,† George D. Weiblen,‡ and James M. Cook*,1 *Department of Biology & NERC Centre for Population Biology, Imperial College, Silwood Park, Ascot, Berkshire SL5 7PY, United Kingdom; †INRA, Centre de Biologie et de Gestion des Populations, Campus International de Baillarguet, CS 30016, 34988 Montferrier-sur-Lez, France; and ‡Department of Zoology, 203 Natural Science Building, Michigan State University, East Lansing, Michigan 48824 Received October 24, 2000; revised April 13, 2001 derpins the idea (Fahrenholz’s rule) that host and par- Figs (Ficus spp., Moraceae) and their pollinating asite phylogenies should be mirror images (Lyal, 1986). wasps form an obligate mutualism, which has long Although cospeciation is not a new idea, the opportu- been considered a classic case of coevolution and co- nity offered by molecular techniques to test it has led to speciation. Figs are also exploited by several clades of a burst of interest and activity in this area (Hafner et nonpollinating wasps, which are parasites of the mu- al., 1994; Hafner and Page, 1995; Moran et al., 1995; tualism and whose patterns of speciation have re- Page, 1996a; Farrell and Mitter, 1998; Peek et al., ceived little attention. We used data from nuclear and 1998; Clark et al., 2000). Molecular markers may per- mitochondrial DNA regions to estimate the phylog- mit the estimation of phylogenies for parasites and enies of 20 species of Pleistodontes pollinating wasps their hosts by use of genes that (unlike some morpho- and 16 species of Sycoscapter nonpollinating wasps associated with Ficus species in the section Mal- logical traits) are not coadapted to the species interac- vanthera. We compare the phylogenies of 15 matched tion. Further, if cospeciation is inferred, molecular Pleistodontes/Sycoscapter species pairs and show that data can be used to compare rates of evolution in host the level of cospeciation is significantly greater than and parasite lineages (Hafner and Nadler, 1990; that expected by chance. Our estimates of the maxi- Hafner et al., 1994; Hafner and Page, 1995; Moran et mum level of cospeciation (50 to 64% of nodes) are very al., 1995; Page, 1996a; Paterson et al., 2000). similar to those obtained in other recent studies of To date few cospeciation studies have produced mo- coevolved parasitic and mutualistic associations. lecular phylogenies for both sets of taxa and insights However, we also show that there is not perfect con- are limited to a few systems. One interaction in par- gruence of pollinator and parasite phylogenies (for ticular—that between pocket gophers (geomyid ro- any substantial clade) and argue that host plant dents) and their chewing lice—has dominated both switching is likely to be less constrained for Sycoscap- methodological (Page, 1990, 1993a,b, 1994) and biolog- ter parasites than for Pleistodontes pollinators. There ical (Nadler et al., 1990; Demastes et al., 1998) debates is perfect correspondence between two terminal about cospeciation. The pioneering studies based on clades of two sister species in the respective phylog- enies, and rates of molecular evolution in these pairs allozyme (Hafner and Nadler, 1988) and DNA (Hafner et al., 1994) data provided evidence for a high degree of are similar. © 2001 Academic Press Key Words: Agaonidae; Pleistodontes; Sycoscapter; cospeciation in the gopher–louse assemblage. How- Ficus; fig wasps; Malvanthera; coevolution; cospecia- ever, both studies are based on small phylogenies and tion; 28S; ITS2; cytochrome b. may not be typical of mammals and lice, or other ecto- parasites, in general (Taylor and Purvis, 2001). A re- cent study of pinworms and primates (Hugo, 1999) INTRODUCTION suggests that at least some endoparasites have under- gone considerable cospeciation with their hosts. More The speciation of parasites is often thought to be predictably, high cospeciation levels have been in- driven by the speciation of their hosts, such that, when ferred between invertebrates and endosymbiotic algae an ancestral host species splits into two daughter spe- or bacteria that contribute significantly to host metab- cies, its parasite also speciates. This assumption un- olism (Peek et al., 1998; Funk et al., 2000). Finally, whereas endogenous retroviruses generally track host 1 To whom correspondence and reprint requests should be ad- phylogeny, occasional host shifts do occur, even be- dressed. Fax (ϩ44) 207 594 2339. E-mail: [email protected]. tween mammals and birds (Martin et al., 1999). 55 1055-7903/01 $35.00 Copyright © 2001 by Academic Press All rights of reproduction in any form reserved. 56 LOPEZ-VAAMONDE ET AL. Lice, endosymbiotic bacteria, retroviruses, and fig logenetic studies of Panamanian pollinating and asso- wasps are not expected to have the same levels of ciated nonpollinating wasps (Machado et al., 1996) and cospeciation as they encompass a range of interactions, of figs and pollinators in general (Herre et al., 1996), with different transmission mechanisms. However, the were consistent with high levels of cospeciation. same methods can be used to analyze these different Australasia harbors great Ficus diversity with at systems, and, as case studies accumulate, it should least 130 species in New Guinea alone (Wiebes, 1982a). become possible to identify general features of interac- The section Malvanthera Corner comprises 22 species, tions (e.g., transmission mode) that promote or limit 14 of which are endemic to Australia and 8 to New cospeciation (Clayton et al., 2001). Guinea (Corner, 1965). There are two distinct groups: a Phytophagous insects can be regarded as parasites of lineage of stranglers and free-standing trees found in their host plants (Price, 1977) and may be generalists rainforests of eastern Australia and New Guinea and a or specialists. Some specialists attack just a single clade of lithophytic shrubs and trees found in more arid plant species, whereas many others feed on a few parts of Australia. Malvanthera species are all polli- closely related species. Interaction and coevolution nated by wasps in the genus Pleistodontes and are have probably been important during the radiations of exploited by a range of nonpollinating wasps (Boucek, plant lineages and their insect herbivores, with plant 1988; Cook and Power, 1996), of which Sycoscapter chemistry playing a key role (Ehrlich and Raven 1964); species are the most prevalent. Sycoscapter larvae are however, strict sense cospeciation seems generally un- thought to be either true parasitoids or phytophagous likely (Farrell and Mitter, 1990, 1998). More specific kleptoparasites of the pollinators (Boucek, 1993; interactions occur between certain groups of plants Compton et al., 1994). Although both are referred to as and their pollinators and perhaps the two best-known fig wasps, Pleistodontes and Sycoscapter are distantly cases are figs and fig wasps (Herre, 1996; Anstett et al., related genera from different families of chalcid wasps 1997) and yuccas and yucca moths (Pellmyr and Lee- (Rasplus et al., 1998). bens-Mack, 1999). Both of these systems are mutual- In this paper, we use nuclear and mitochondrial isms but also involve conflicts of interest, principally DNA sequences to infer phylogenies for 20 Pleistodon- over whether female flowers nourish developing seeds tes and 16 Sycoscapter species from Malvanthera figs. or insect larvae. We test two null hypotheses of host–parasite associa- Figs (Ficus spp.) and their pollinating wasps (family tions (Fig. 1; see Huelsenbeck et al., 1997, 2000; Clark Agaonidae sensu Rasplus et al., 1998) provide a classic et al., 2000). H1 is that host and parasite phylogenies example of an obligate mutualism. Most Ficus species have been produced by independent random-branching have their own unique pollinating wasp species, whose processes. Acceptance of H1 infers that cospeciation larvae feed on the fig seeds. Each partner relies on the has been rare or absent, whereas rejection of H1 sug- other to complete its life cycle and many morphological gests that cospeciation (or another process imparting characters of the partners are thought to be coadapted congruence) has occurred. In the latter case we can (Herre, 1989; Van Noort and Compton, 1996). The bi- then test H2 that Pleistodontes and Sycoscapter data ology of figs and their pollinators has led to a general sets are consistent with the same underlying tree to- acceptance of the idea that they have coevolved and pology (Peek et al., 1998; Clark et al., 2000; Huelsen- cospeciated extensively. However, rigorous testing of beck et al., 2000, 2001; Johnson et al., 2001). Accep- this idea has been hindered by the fact that the polli- tance of H2 would provide stronger evidence for strict nator taxonomy has developed under the influence of cospeciation and permit a test of H3 that the rates of existing fig taxonomy and a strong cospeciation as- molecular evolution and speciation times are the same sumption (Ramirez, 1974, 1977; Wiebes, 1982b). in the two clades. Fig-based invertebrate communities lend them- selves to cospeciation studies because, in addition to pollinating wasps, there are diverse nonpollinating MATERIAL AND METHODS wasps, mites, drosophilids (Harry et al., 1996), and Taxon Sampling parasitic nematodes (Poinar and Herre, 1991). Many (perhaps
Recommended publications
  • 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]
  • 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]
  • Fig Parrot Husbandry
    Made available at http://www.aszk.org.au/Husbandry%20Manuals.htm with permission of the author AVIAN HUSBANDRY NOTES FIG PARROTS MACLEAY’S FIG PARROT Cyclopsitta diopthalma macleayana BAND SIZE AND SPECIAL BANDING REQUIREMENTS - Band size 3/16” - Donna Corporation Band Bands must be metal as fig parrots are vigorous chewers and will destroy bands made of softer material. SEXING METHODS - Fig Parrots can be sexed visually, when mature (approximately 1 year). Males - Lower cheeks and centre of forehead red; remainder of facial area blue, darker on sides of forehead, paler and more greenish around eyes. Females - General plumage duller and more yellowish than male; centre of forehead red; lower cheeks buff - brown with bluish markings; larger size (Forshaw,1992). ADULT WEIGHTS AND MEASUREMENTS - Male - Wing: 83-90 mm Tail : 34-45 mm Exp. cul: 13-14 mm Tars: 13-14 mm Weight : 39-43 g Female - Wing: 79-89 mm Tail: 34-45 mm Exp.cul: 13-14 mm Tars: 13-14 mm Weight : 39-43 g (Crome & Shield, 1992) Orange-breasted Fig Parrot Cyclopsitta gulielmitertii Edwards Fig Parrot Psittaculirostris edwardsii Salvadori’s Fig Parrot Psittaculirostris salvadorii Desmarest Fig Parrot Psittaculirostris desmarestii Fig Parrot Husbandry Manual – Draft September2000 Compiled by Liz Romer 1 Made available at http://www.aszk.org.au/Husbandry%20Manuals.htm with permission of the author NATURAL HISTORY Macleay’s Fig Parrot 1.0 DISTRIBUTION Macleay’s Fig Parrot inhabits coastal and contiguous mountain rainforests of north - eastern Queensland, from Mount Amos, near Cooktown, south to Cardwell, and possibly the Seaview Range. This subspecies is particularly common in the Atherton Tableland region and near Cairns where it visits fig trees in and around the town to feed during the breeding season (Forshaw,1992).
    [Show full text]
  • Evolution of Tbe Mandibular Appendage in Figwasps (Hymenoptera: Agaonidae)
    Rev. Biol. Trop., 39 (1): 87-95. 1991 Evolution of tbe mandibular appendage in figwasps (Hymenoptera: Agaonidae) William Ramírez B. Escuela de Fitotecnia, Universidadde Costa Rica. (Ree. 20-IX-I990. Acep. 15-X-I990) Abstract: The phylogenetic value of the conformation of the mandibular appendages, the number of mandibular glands andother head characters in the Agaonidae are examined.The phylogenetic arrangement suggests thatthe pre­ agaonid had a normal bidentate mandible with two glands, and a undistinct facial groove as in sorneDiazi ella (Sycoecinae), Sycophaga (Sycophaginae) and in related chalcidoid non-pollinating fig wasps. lt also had thirteen-seg­ mented antennae;'a long scape, several times ¡onger than wide a long triangular pedicel, two or three anelli andeight flagellomeres withsensilla (as uniquely found in Tetrapus). The mandibularappendage apparently co-evolved with the development oí the ostiolum of thesyconium andthe firstmandibular appendage was fixed andhad ridges or lameUae. A flexible hinge evolved laterat its base.The polygamous males were wingless with extendible (solenogastrus) abdo­ men and mate inside the galls. These characters are also found in most Sycophaginae.The ancestor of Aganoidae was probably a primary sycophilous wasp, withdorsoventral depressed head, thorax and abdomen, that oviposit through thestylar channel as stiU Sycophaga sycomori does. Sycophaga wasps withits apterousand polyg amous males seem to be the sister group of Agaonidae. The Agaonidae females are characterized by their mandibular appendage and the antennalprocess. They have a prognathous head with gula. Themales are wingless polygamous andsolenogastrous. Key words: Agaonidae(Hymenoptera), mandibular appendage, evolution, phylogeny. Ficus is pollinated by small chalcidoid wasps and two or three antennal anelli.
    [Show full text]
  • Ficus Rubiginosa 1 (X /2)
    KEY TO GROUP 4 Plants with a milky white sap present – latex. Although not all are poisonous, all should be treated with caution, at least initially. (May need to squeeze the broken end of the stem or petiole). The plants in this group belong to the Apocynaceae, Euphorbiaceae, Moraceae, and Sapotaceae. Although an occasional vine in the Convolvulaceae which, has some watery/milky sap will key to here, please refer to Group 3. (3.I, 3.J, 3.K) A. leaves B. leaves C. leaves alternate opposite whorled 1 Leaves alternate on the twigs (see sketch A), usually shrubs and 2 trees, occasionally a woody vine or scrambler go to Group 4.A 1* Leaves opposite (B) or whorled (C), i.e., more than 2 arising at the same level on the twigs go to 2 2 Herbs usually less than 60 cm tall go to Group 4.B 2* Shrubs or trees usually taller than 1 m go to Group 4.C 1 (All Apocynaceae) Ficus obliqua 1 (x /2) Ficus rubiginosa 1 (x /2) 2 GROUP 4.A Leaves alternate, shrubs or trees, occasional vine (chiefly Moraceae, Sapotaceae). Ficus spp. (Moraceae) Ficus, the Latin word for the edible fig. About 9 species have been recorded for the Island. Most, unless cultivated, will be found only in the dry rainforest areas or closed forest, as in Nelly Bay. They are distinguished by the latex which flows from all broken portions; the alternate usually leathery leaves; the prominent stipule (↑) which encloses the terminal bud and the “fig” (↑) or syconia. This fleshy receptacle bears the flowers on the inside; as the seeds mature the receptacle enlarges and often softens (Think of the edible fig!).
    [Show full text]
  • Flying Foxes Jerry Copy
    Species Common Name Habit Flowering Fruit Notes Acacia macradenia Zigzag wattle Shrub August Possible pollen source Albizia lebbek Lebbek Tall tree summer Source of nectar. Excellent shade tree for large gardens. Alphitonia excelsa Red ash Tree October to March November to May Food source for Black and Grey- headed flying fox Angophora costata Smooth-barked Tall tree December to Source of nectar apple January Angophora Rough-barked apple Tall tree September to Source of nectar floribunda February Angophora costata Smooth-barked Tall tree November to Source of nectar subsp. leiocarpa apple, Rusty gum February Archontophoenix Alexander palm Tree-like November to January Food source for alexandrae December Spectacled flying fox. Good garden tree Archontophoenix Bangalow palm Tree-like February to June March to July Food source for cunninghamiana Grey-headed flying fox. Good garden tree Species Common Name Habit Flowering Fruit Notes Banksia integrifolia Coastal Shrub or small tree Recurrent, all year Food source for honeysuckle round Black and Grey- headed flying fox. Good garden tree Banksia serrata Old man banksia Shrub or small tree February to May Source of nectar. Good garden tree Buckinghamia Ivory curl tree Small tree December to Possible source of celsissima February nectar. Good garden tree Callistemon citrinus Crimson bottlebrush Shrub or small tree November and Source of nectar. March Good garden tree Callistemon salignus White bottlebrush Shrub or small tree spring Source of nectar. Good garden tree Castanospermum Moreton Bay Tall tree spring Source of nectar australe chestnut, Black bean Corymbia citriodora Lemon-scented gum Tall tree may flower in any Source of nectar season Corymbia Red bloodwood From mallee to tall summer to autumn Source of nectar gummifera tree Corymbia Pink bloodwood Tall tree December to March Source of nectar.
    [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]
  • Ficus Rubiginosa F. Rubiginosa Click on Images to Enlarge
    Species information Abo ut Reso urces Hom e A B C D E F G H I J K L M N O P Q R S T U V W X Y Z Ficus rubiginosa f. rubiginosa Click on images to enlarge Family Moraceae Scientific Name Ficus rubiginosa Desf. ex Vent. f. rubiginosa Ventenat, E.P. (1805) Jard. Malm. : 114. Type: New Holland; holo: FI. Fide Dixon et al. (2001). Leaves and figs. Copyright G. Sankowsky Common name Fig; Small-leaved Fig; Larger Small-leaved Fig; Larger Small Leaf Fig; Fig, Larger Small Leaf; Fig, Small-leaved Stem A strangling fig, hemi-epiphyte or lithophyte to 30 m. Leaves Figs. Copyright G. Sankowsky Petioles and twigs produce a milky exudate. Stipules about 2-6 cm long, usually smooth on the outer surface, occasionally hairy. Petioles to 4 cm long, channelled on the upper surface. Leaf blades rather variable in shape, about 6-10 x 2-3 cm; leaves hairy. Flowers Tepals glabrous. Male flowers dispersed among the fruitlets in the ripe figs. Anthers reniform. Stigma cylindric, papillose, often slightly coiled. Bracts at the base of the fig, two. Lateral bracts not present on the outside of the fig body. Fruit Scale bar 10mm. Copyright CSIRO Figs pedunculate, globose, about 10-18 mm diam. Orifice triradiate, +/- closed by inflexed internal bracts. Seedlings Cotyledons ovate-oblong, about 5 mm long. At the tenth leaf stage: leaves ovate, apex acute or bluntly acute, base obtuse or cordate, margin entire, glabrous, somewhat triplinerved at the base; oil dots not visible; stipules large, sheathing the terminal bud, about 20-40 mm long.
    [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]
  • 12 Laman St Figs Report 280910[1] Mckenzie
    Expert Witness Report Parks and Playgrounds Movement Inc v Newcastle City Council [2010] NSWLEC 40745 Laman Street, Cooks Hill (between Dawson Street and Darby Street) Prepared for Parks and Playgrounds Movement Inc (Contact: Mr Doug Lithgow, President) Prepared by Ian McKenzie – Consulting Arborist 28 September 2010 Table of Contents 1 Summary.....................................................................................................2 2 Introduction ................................................................................................4 2.1 Acknowledgement ................................................................................................... 4 2.2 Disclaimer ................................................................................................................ 4 2.3 Background .............................................................................................................. 4 2.4 Brief ......................................................................................................................... 5 2.5 Methodology ............................................................................................................ 6 3 Site Details..................................................................................................7 3.1 Site Description........................................................................................................ 7 3.2 Heritage Status ........................................................................................................
    [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]