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Comparative Anatomy of the Fig Wall (Ficus, Moraceae)

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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; 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)
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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. 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.

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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. The fig wall is the main structure of the fig giving its shape but the wall has also important ecological functions such as protection of fig seeds and fig wasp larvae. Nevertheless, the fig wall anatomy is poorly understood. This study aims to examine the fig wall anatomy of 22 Ficus taxa (21 species, one species having two varieties) in Taiwan in order to reveal the diversity in anatomy of the fig wall. We found that these 21 fig species exhibited a great variety in fig wall anatomy, from the simplest parenchymatic wall to complex fig walls. Fig walls of 12 sampled taxa developed aerenchyma and sclerenchyma formations whereas seven taxa had fig walls containing tanniferous cells. Five anatomical types of fig walls have been identified according to the presence or absence of the different differentiated tissues. These types are distributed among the Ficus subgenera. Further studies on tissue differentiations of the fig wall should be investigated in other Ficus species as well as the ecological functions of the fig wall. Keywords: Comparative Anatomy; Ficus; Histology; Inflorescence

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1. Introduction

The Moraceae family comprises about 38 genera and 1180 species (Christenhusz and Byng 2016) displaying a great diversity of life history traits with few synapomorphic characters such as laticifers, anatropous ovules, and apical placentation (Sytsma et al. 2002). Among the six Moraceae tribes (Clement and Weiblen 2009), the Ficeae tribe is monogeneric with the single genus Ficus (Berg 2001) defined by a common feature: a closed inflorescence with a single tight orifice (ostiole) closed by bracts called fig (or syconium) (Berg and Corner 2005).

The Moraceae phylogeny shows that the Ficeae tribe is not basal and the inflorescence morphology within the Moraceae family has evolved in many directions (Clement and Weiblen 2009). This diversity is also present within the tribes. For instance, figs display many shapes from the ellipsoid figs of Ficus dammaropsis measuring about 10cm in diameter with conspicuous lateral bracts to the globose minute figs of F. caulocarpa measuring half a centimetre (Berg and Corner 2005) or to the Australian banana fig, F. pleurocarpa (Dixon 2003). In addition to the fig morphology, Ficus species are mostly famous for being pollinated by minute wasps from the Agaonidae family (Hymenoptera: Chalcidoidea) and solely by the wasps of this family. Together, the fig tree and the pollinating wasp are mutualistic partners: the fig provides oviposition sites

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for the agaonid wasp which bring pollen inside the fig in order to pollinate the enclosed flowers (Kjellberg et al. 2005). This nursery pollination mutualism is targeted by many nonpollinating wasp (NPFW) species parasitizing either the fig ovules or fig wasp larvae (Bronstein 1991; Tzeng et al. 2008). These NPFWs can drastically reduce the number of seeds and pollinating wasps produced in a single fig (Cardona et al. 2013). The specificity of these NPFWs is their way to reach the fig ovules or the fig wasp larvae inside the figs. Indeed, they are laying eggs from outside the figs, using their long ovipositor (Ghara et al. 2011). They are not using the fig ostiole to penetrate the fig but pass through the fig wall.The fig wall is the tissue between the ovules and the outside world (Fig. 1). The thickness of the fig wall increases during the fig development with the size of the fig (Galil et al. 1970). Moreover, some figs of Ficus erecta var. beecheyana, in Taiwan, have a fig wall thick enough to protect their flowers from nonpollinating fig wasp oviposition (Tzeng et al. 2014). Other than laticifers (Marinho et al. 2017), the anatomy of the fig wall is poorly understood. To our knowledge, the general description is “the fig wall

generally parenchymatic and contains some 30-40 cell layers. Various sclerified cell layers may occur […] and contains many laticifers, tannin cells, and a vascularization”

(Verkerke 1989). This general description based on information from different reviewed studies but the fig wall can contain more cell layers than in the Verkerke’s description reaching up to 48 cell layers in F. ingens in South Africa (Baijnath and Naicker 1989).

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One of the common feature seems to be that, when it occurs, sclerenchyma becomes a larger part of the fig wall during the fig development (Galil et al. 1970; Verkerke 1986; Baijnath and Naicker 1989) but not all the fig species have a hardened fig wall (Verkerke 1988). This statement leads to the question why some fig species have developed a hardened fig wall. Considering the cost of the NPFWs (Cardona et al. 2013) and the adaptation of these NPFWs to the oviposition through the fig wall (Ghara et al. 2011), the fig wall may be an important organ for the fig trees to manage the parasitism by NPFWs. For instance, some closely related fig species, such as F. caulocarpa and F. subpisocarpa, have a very a different NPFW community (Bain et al. 2015): 20 species for F. subpisocarpa and only two species for F. caulocarpa. Then, is the fig wall of these two species very different? Another example is F. erecta var. beecheyana which can have a very thick fig wall excluding any NPFWs. What does the fig wall of F. erecta var. beecheyana consist of? And more generally, what are the fig walls made of? Other than the simple description from Verkeke’s work (1988), we have little information except that fig walls display some anatomical diversity. Moreover, what happens in dioecious fig species? In these species, the male trees bear figs with pollinating fig wasps and are the targets of many NPFWs whereas the female trees are producing only seeds and NPFWs are rarely breeding in female figs (Wu et al. 2013). If the fig wall is ecologically linked to the presence of the NPFWs, we can expect to see morphological differences

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between male and female trees. Thus, this study aims to investigate the fig wall anatomy of most of the Taiwan Ficus species (21 species, one of which has two varieties, from six subgenera) and tentatively link the anatomical structures to ecological functions such as the parasitism from NPFWs. We also discuss the effect of phylogeny on the distribution of the anatomical structures within the genus Ficus.

2. Methods

2.1. Fig development and reproductive biology of pollinating fig wasps

The pollinating fig wasps grow as larva inside the figs and mate shortly after the eclosion of their gall (pupal case) when they are still inside their natal fig (Kjellberg et al. 2005). Only the winged female pollinating fig wasps will leave the fig to find another fig of the same species because pollinating fig wasps pollinate only one fig tree species (few exceptions to this rule have been found (Compton et al. 2009)). Once the female pollinating wasp has found a receptive fig (see below for the descriptions of fig developmental phases), it enters inside the fig to pollinate it and lay eggs. In the figs, pollinating and nonpollinating fig wasps have a very similar life cycle but, instead of entering to figs to lay eggs, NPFWs lay eggs from outside of the fig and have to penetrate

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the fig wall with their ovipositors (Weiblen et al. 2002).
Based on the definition by Galil and Eisikowitch (1968), fig development is divided into five phases. The A phase (prefemale phase) is the earliest stage, in which all the flowers are immature. In the B phase (female phase), the female flowers are receptive, and pollinating fig wasps can pollinate them, thus entering the figs through the ostiole. During the C phase (interfloral phase), both the fig wasp larvae and seeds develop. The D phase (male phase) begins when the first male wasp hatches. Subsequently, the wasps mate, and the males dig an exit for the female winged wasps, which leave the fig to find another receptive fig. During the E phase (post-floral phase), the figs ripen and attract frugivores to disperse the seeds.
During their larval stage, NPFWs exhibit a great diversity of hosts and diets: for instance, some larvae feed on the plant tissue growing from a gall whereas other NPFW species feed on the larva of another fig wasp species (they are parasitoid). Most NPFWs do not feed as adult. Thus, depending on their larval diet, NPFWs will appear on figs to lay their eggs during two of the developmental phases: During the late B phase or early C phase, if the NPFW species are gall makers; and later during the C phase if they are inquiline (gall thief) or parasitoid (Kerdelhué and Rasplus 1996a). Thus, the parasitic pressure from NPFWs is almost exclusively during the C phase.
The developmental process of dioecious fig tree species is slightly different. In these

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species, male figs produce pollen and wasps but no seeds (no E phase), and female figs produce seeds but no wasps (no D phase) (Harrison and Yamamura 2003). Thus, the male figs are the target of the NPFWs and female figs only rarely (Wu et al. 2013) which could lead to anatomical differences between male and female figs. In addition, female trees produce usually less figs than male trees (Bain et al. 2014) providing less oviposition sites for NPFWs. Also, dioecious fig species host a reduced diversity of NPFW species compared to monoecious fig species (Kerdelhué and Rasplus 1996b).

2.2 Plant material

Twenty-six species (29 taxa in total counting the varieties and subspecies) of native or introduced (one single species: F. altissima) fig trees have been recorded in Taiwan (Bain et al. 2015). The proportion of dioecious species is particularly high (20 of the 26 species) while worldwide about half of the Ficus species are dioecious. In Taiwan, fig tree species are widely distributed from the coastal areas (F. pedunculosa var. mearnsii) to the montane forests up to 1000 m above sea level (F. erecta var. beecheyana), with several

species, such as F. benjamina var. bracteata and F. pedunculosa, restricted to the

southern tropical part of the island (Tzeng 2004).
More than 200 figs from 21 fig tree species (one species with two varieties: sample sizes per species in Fig. 2 and 3) from six subgenera were collected on Taiwan Island and

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Orchid Island (southeast of Taiwan Island) from September 2013 to September 2015 (Table S1 in supplementary material). In Northern Taiwan, an exotic species, F. altissima, which has a pollinating wasp in Taiwan, was sampled on the National Taiwan University campus and in Da’an Forest Park in Taipei City. Figs of F. caulocarpa, F. subpisocarpa, and F. microcarpa were sampled at various stages of development in order to study the developmental changes in detail whereas figs at the C phase were sampled for all the remaining studied fig species.

2.3. Sample preparation

In order to measure the fig diameter and fig wall thickness, the figs were cut longitudinally through the ostiole and peduncle (Fig. 1). For ideal fixation, the figs were dissected from the widest part into small blocks (less than 4 mm long and 5 mm wide).
For paraffin sections, the samples were first fixed either in FPGA solution
(formalin/propionic acid/glycerol/95% ethanol/distilled water at 1:1:3:7:8) or 0.5% glutaraldehyde in 0.1 M phosphate buffer. In order to reduce the potential for the formation of artefacts, samples collected in the field were directly fixed in 0.5% glutaraldehyde in 0.1 M phosphate buffer. The tissue blocks were then washed in 50% alcohol or 0.1 M phosphate buffer and dehydrated in a tert-butyl alcohol series. After dehydration, the samples were infiltrated and embedded in paraffin (Paraplast; Leica

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Biosystems), and then sectioned using a rotary microtome (HM320, Microm) mounted with disposable microtome blades (Leica Biosystems). The thickness of the sections was 10–14 μm depending on the hardness of the tissues and was 20 μm for histochemical tests.

2.4 Staining and histochemical tests

The following staining methods were performed on deparaffinized sections: (a) Toluidine Blue O in 1% borax as the metachromatic stain (Mercer 1963), (b) Fast Green–Safranin O for differentiating the tissue structure (Sass 1958), (c) phloroglucinol in 6 N HCl for lignin localization (O’Brien and McCully 1981), and (d) ferric chloride solution with sodium carbonate to highlight tannins (Johansen 1940). The freehand sections fixed with 70% ethanol were treated with Sudan black B for staining the latex (Johansen 1940). Positive control staining using locally known plant species samples was conducted for the aforementioned staining methods.

2.5 Imaging procedure and analyses

To measure the thickness of the fig wall (from the outer epidermis to the inner epidermis at the “equator” of the fig (Fig. 1)), the sections were photographed using a digital camera (650D, Canon) mounted on a stereomicroscope (S8 APO, Leica Microsystems). For other

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freehand and paraffin sections, the slides were photographed using a Nikon D3 digital camera mounted on a light microscope (DMRB, Leica Microsystems). The thickness of the fig walls and each tissue type were measured using Fiji image processing software (open source). Spearman rank correlation test was used to test the correlation between fig wall thickness and fig diameter using SYSTAT 13 (Systat Software®, San Jose, CA, USA).

3. Results

3.1. General fig wall structure

Fig wall thickness was positively correlated with fig size (Fig. S1 in supplementary material). Among the 22 taxa (mean = 1640 ± 166 µm; n = 132), the figs of F. pumila var. pumila had the thickest wall (mean = 8472 µm; n = 7), and those of F. ampelas had the thinnest wall (mean = 529 µm; n = 4) (Fig. 2). Cross sections of the fig walls demonstrated that they were composed of dermal (outer and inner epidermis), ground, and vascular tissues (Fig. S2 in supplementary material). The outer epidermis may differentiate into trichomes or lenticels. Laticifers and druses were common in the ground tissue of the fig wall and were generally associated with vascular bundles.

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3.2. Fig wall classification

The fig walls (at C phase) were categorized into five types (Table 1) according to the ground tissue composition of the C phase fig wall (Fig. 4). The species and their fig wall types are summarized in Table 1. Type I: The ground tissue was composed of only the regular parenchyma. This type was observed in the dioecious fig species (nine species from four subgenera). However, in two species (F. septica and F. benguetensis), only the female figs belong to the type I. On the other hand, the male figs of these two species belong to the type III. Type II: The ground tissue was characterized by aerenchyma (tissue with enlarged air spaces: Evans 2003) and a high density of laticifers. The aerenchyma was distributed between the parenchyma tissues near the outer and the inner epidermis. This type was observed only in two of the Taiwanese species (Table 1). Type III: The ground tissue was characterized by sclerenchyma formation. Three subtypes were recognized according to the distribution of sclerenchyma cells: one-layered sclerenchyma (type IIIa), two-layered sclerenchyma (type IIIb), and scattered sclerenchyma cells (type IIIc). Type IIIc was observed only in F. nervosa figs. Type IV: The parenchyma was characterized by abundant tannin deposition in the cells. This type was observed only in three species belonging to two dioecious subgenera (Table

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1). Type V: The parenchyma was characterized by the presence of both sclerenchyma and tanniferous cells. This type was found mostly in monoecious species (75% of the species) and once in the male figs of the dioecious species, F. pedunculosa var. mearnsii.

3.3. Developmental progress of fig walls in F. caulocarpa, F. subpisocarpa, and F.

microcarpa

The fig walls of F. caulocarpa and F. subpisocarpa were of type IIIb and their

developmental progress was similar (Table 1, Fig. 3). During A phase, the fig walls were composed of only parenchyma. Tanniferous cells were present at the beginning of the A phase, then quickly degraded after the fig peduncle elongated. At the end of the A phase, a thin sclerenchyma layer developed near the outer epidermis, and a second thin sclerenchyma layer formed when the figs became receptive (B phase). This two-layered sclerenchyma structure was observed toward the end of maturation.
The fig wall of F. microcarpa was of type V and exhibited a distinct wall structure until maturation. In comparison to F. caulocarpa and F. subpisocarpa, the young fig walls (A phase) were composed of only parenchyma without tannins. At the end of A phase, a thick tanniferous cell layer formed adjacent to the outer epidermis. In the B phase, a sclerenchyma layer formed on the inner side of the tanniferous cell layer. After pollination

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(C phase), a second layer of sclerenchyma developed within the tanniferous cells, and the first sclerenchyma cell layer thickened. The two-layered sclerenchyma pattern was observed until the end of maturation. Nevertheless, the tannins disappeared in E phase.

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  • Developmental Anatomy of Ficus Ingens Syconia in Relation to Its Wasp Faunula

    Developmental Anatomy of Ficus Ingens Syconia in Relation to Its Wasp Faunula

    S.Afr.J. Bot. , 1989, SS( 4): 409- 421 409 Developmental anatomy of Ficus ingens syconia in relation to its wasp faunula H. Baijnath* and S. Naicker Department of Botany, University of Durban-Westville, Private Bag X54001, Durban, 4000 Republic of South Africa Accepted 6 February 1989 Syconial development in Ficus ingens (Miq.) Miq., extends over a period of 70-80 days and displays the same basic pattern reported by earlier workers. Developmental anatomical observations of the different components of the syconium are presented. Following pollination, structural changes of gall and seed flowers differ and the implications of some of these are considered. Since hairs occur at different stages of develop­ ment, speculations are presented on their role. In most instances, the anatomical changes could be related to wasp development and behaviour, thus highlighting the close mutualistic relationship that exists between figs and wasps. Die ontwikkeling van die sikonium van Ficus ingens (Miq.) Miq. duur vir 'n peri ode van 70-80 dae en vertoon dieselfde basiese patroon soos reeds vroeer deur ander werkers aangetoon. Waarnemings betreffende die ontwikkelingsanatomie van die verskillende dele van die sikonium word aangebied. Die strukturele verander­ ings in die gal- en saadvormende blomme nadat bestuiwing plaasgevind het verskil en die implikasies van sommige hiervan word bespreek. Weens die feit dat hare op verskeie stadiums van ontwikkeling ontstaan, word daar oor hulle rol gespekuleer. Die anatomiese veranderings hou in die meeste gevalle verband met die ontwikkeling en gedrag van wespe waardeur die noue mutualistiese verwantskap tussen vye en wespe beklemtoon word. Keywords: Anatomy, Ficus ingens, syconium 'To whom correspondence should be addressed Introduction Ficus ingens (Mig .) Mig., is a medium-sized spreading dse tree which belongs to the subgenus Urostigma and is fairly widespread in Africa.
  • Title Kitongwe Name of Plants: a Preliminary Listing Author(S)

    Title Kitongwe Name of Plants: a Preliminary Listing Author(S)

    Title Kitongwe Name of Plants: A Preliminary Listing Author(s) NISHIDA, Toshisada; UEHARA, Shigeo Citation African Study Monographs (1981), 1: 109-131 Issue Date 1981 URL http://dx.doi.org/10.14989/67977 Right Type Departmental Bulletin Paper Textversion publisher Kyoto University 109 KITONGWE NAME OF PLANTS: A PRELIMINARY LISTING Edited by Toshisada NISHIDA and Shigeo UEHARA Departnlent ofAnthropology, Faculty ofScience, University of Tokyo, Tokyo, Japan INTRODUCTION Field workers of Kyoto University Africa Primatological Expedition collected plants in western Tanzania. Experts of Japan International Cooperation Agency working as Game (Wildlife) Research Officers at Kasoje Chimpanzee Research Station (Mahale Mountains Wildlife Research Centre) have concentrated their collecting activities Inainly to the Mahale Mountains. The collection of plants with notes of kitongwe name not only has facilitated the ecological studies on wild chimpanzees (and other wild animals), but also will be of use in analyzing the traditional system of classification of plants among Batongwe, as well as in re­ cording for ever a rapidly-vanishing culture. This is a revised version, though still only preliminary one, of the manuscript entitled "Sitongwe-Latin Dictionary of Plants" edited by T. Nishida on 4 April, 1975. COLLECTION The researchers who have contributed to this work in the collection of the plants are listed below, with the reference number in the East African Herbarium'(Kenya Herbarium), the number of total specimens collected, and the specimen number in each collection. All the plants with known kitongwe nalne collected within the Tongwe (and Bende) territory are listed in this edition. Local emphasis is put on the Mahale Mountains and especially on Kasoje area.
  • Studies on New Guinea Moths. 1. Introduction (Lepidoptera)

    Studies on New Guinea Moths. 1. Introduction (Lepidoptera)

    PROC. ENTOMOL. SOC. WASH. 105(4), 2003, pp. 1034-1042 STUDIES ON NEW GUINEA MOTHS. 1. INTRODUCTION (LEPIDOPTERA) SCOTT E. MILLER, VOJTECH NOVOTNY, AND YVES BASSET (SEM) Department of Systematic Biology, National Museum of Natural History, Smith- sonian Institution, Washington, DC 20560-0105, U.S.A. (e-mail: [email protected]. edu); (VN) Institute of Entomology, Czech Academy of Sciences and Biological Faculty, University of South Bohemia, Branisovska 31, 370 05 Ceske Budejovice, Czech Republic; (YB) Smithsonian Tropical Research Institute, Apartado 2072, Balboa, Ancon, Panama Abstract.•This is the first in a series of papers providing taxonomic data in support of ecological and biogeographic studies of moths in New Guinea. The primary study is an extensive inventory of the caterpillar fauna of a lowland rainforest site near Madang, Papua New Guinea, from 1994•2001. The inventory focused on the Lepidoptera com- munity on 71 woody plant species representing 45 genera and 23 families. During the study, 46,457 caterpillars representing 585 species were sampled, with 19,660 caterpillars representing 441 species reared to adults. This introductory contribution is intended to provide background on the project, including descriptions of the study site, sampling methods, and taxonomic methods. Key Words: Malesia, Papua New Guinea, Lepidoptera, biodiversity, rearing, community ecology A very large portion of tropical biodi- 1992 and 1993 (Basset 1996, Basset et al. versity consists of herbivorous insects, and 1996). This paper represents the first in a among them, Lepidoptera are among the series of papers providing taxonomic doc- most amenable to study. To better under- umentation in support of the broader stud- stand the structure and maintenance of trop- ies, and is intended to provide general back- ical biodiversity, we undertook a series of ground, including descriptions of the study related inventories of Lepidoptera in New site, sampling methods, and taxonomic Guinea.
  • Pharmacognostical and Preliminary Phytochemical Studies on the Leaf Extract of Ficus Pumila Linn

    Pharmacognostical and Preliminary Phytochemical Studies on the Leaf Extract of Ficus Pumila Linn

    ISSN 2278- 4136 ZDB-Number: 2668735-5 IC Journal No: 8192 Volume 1 Issue 4 Online Available at www.phytojournal.com Journal of Pharmacognosy and Phytochemistry Pharmacognostical and Preliminary Phytochemical Studies on the Leaf Extract of Ficus pumila Linn. Jasreet Kaur1* 1. College of Pharmacy, Shoolini University, Solan, HP. India. [E-mail: [email protected]] Ficus pumila Linn. (Family: Moraceae), commonly known as climbing fig. It is widely used as an ethno medicine in china and India. It is prescribed for a wide variety of ailments like diarrhea, hemorrhoids, treating gastrointestinal, piles, uterine problems and other infections. However, detailed scientific information is not available to identify the plant material and to ascertain its quality and purity. In present communication, morphology anatomical and physico-chemical characters along with phytochemical screening and fluorescence analysis of powdered crude drug were carried out for systemic identification and authentication of leaves. This study provides referential information for identification and characterization of Ficus pumila leaf and its extracts. Keyword: Ficus pumila linn, Phytochemical, Morphological, Methanolic extract. 1. Introduction 1.1 Ficus pumila Linn. The genus Ficus represents an important group of Ficus pumila Linn. is a member of the Moraceae trees, not only for their immense value but also family. It is a root climbing evergreen vine for their growth habits. The genus Ficus is an attaching to rocks, walls, tree trunks by means of exceptionally large pan tropical genus with over exudations from the aerial roots. This species is 800 species and belongs to the family moraceae. native to East Asia- south China, Vietnam, The Ficus species are used as food and for Taiwan, New Zealand Nepal, India, Western medicinal properties in Ayurveda and Traditional Australia and Japan[3].
  • Molecular and Morphological Support for a Florida Origin of the Cuban

    Molecular and Morphological Support for a Florida Origin of the Cuban

    Journal of Biogeography (J. Biogeogr.) (2011) SPECIAL Molecular and morphological support ISSUE for a Florida origin of the Cuban oak Paul F. Gugger* and Jeannine Cavender-Bares Department of Ecology, Evolution and ABSTRACT Behavior, University of Minnesota, 100 Ecology Aim The origins of the Cuban biota are of long-standing interest in Building, 1987 Upper Buford Circle, Saint Paul, MN, USA biogeography, and the source of a small live oak (Quercus series Virentes) population on Cuba remains unresolved. Based on morphological evidence, previous authors have hypothesized a Florida origin from either Q. geminata or Q. virginiana or both; a Mexican origin from Q. oleoides; or a hybrid origin from both sources. We use molecular data and taxonomically informative leaf morphology to identify the source species and timing of colonization. Location Cuba, Central America, Mexico and the south-eastern United States. Methods We collected representative samples of Cuban oaks and each putative source species and genotyped each sample at 12 nuclear microsatellites and two chloroplast DNA sequences. We estimated population structure using a Bayesian clustering analysis and F-statistics, pairwise migration rates among taxa, and divergence time using an isolation-with-migration model. We measured seven leaf traits and conducted an analysis of similarity (ANOSIM) to determine which putative source species was most similar to Cuban oaks. Results Cuban oak contains one chloroplast DNA haplotype, which is common in southern Florida. Bayesian clustering analysis of microsatellites revealed that the Cuban oak forms a distinct and pure population cluster, and F-statistics showed that Cuban oaks are differentiated least from Q. virginiana and most from Q.
  • DNA Barcoding Confirms Polyphagy in a Generalist Moth, Homona Mermerodes (Lepidoptera: Tortricidae)

    DNA Barcoding Confirms Polyphagy in a Generalist Moth, Homona Mermerodes (Lepidoptera: Tortricidae)

    Molecular Ecology Notes (2007) 7, 549–557 doi: 10.1111/j.1471-8286.2007.01786.x BARCODINGBlackwell Publishing Ltd DNA barcoding confirms polyphagy in a generalist moth, Homona mermerodes (Lepidoptera: Tortricidae) JIRI HULCR,* SCOTT E. MILLER,† GREGORY P. SETLIFF,‡ KAROLYN DARROW,† NATHANIEL D. MUELLER,§ PAUL D. N. HEBERT¶ and GEORGE D. WEIBLEN** *Department of Entomology, Michigan State University, 243 Natural Sciences Building, East Lansing, Michigan 48824, USA, †National Museum of Natural History, Smithsonian Institution, Box 37012, Washington, DC 20013-7012, USA, ‡Department of Entomology, University of Minnesota, 1980 Folwell Avenue, Saint Paul, Minnesota 55108–1095 USA, §Saint Olaf College, 1500 Saint Olaf Avenue, Northfield, MN 55057, USA,¶Department of Integrative Biology, University of Guelph, Guelph, Ontario, Canada N1G2W1, **Bell Museum of Natural History and Department of Plant Biology, University of Minnesota, 220 Biological Sciences Center, 1445 Gortner Avenue, Saint Paul, Minnesota 55108–1095, USA Abstract Recent DNA barcoding of generalist insect herbivores has revealed complexes of cryptic species within named species. We evaluated the species concept for a common generalist moth occurring in New Guinea and Australia, Homona mermerodes, in light of host plant records and mitochondrial cytochrome c oxidase I haplotype diversity. Genetic divergence among H. mermerodes moths feeding on different host tree species was much lower than among several Homona species. Genetic divergence between haplotypes from New Guinea and Australia was also less than interspecific divergence. Whereas molecular species identification methods may reveal cryptic species in some generalist herbivores, these same methods may confirm polyphagy when identical haplotypes are reared from multiple host plant families. A lectotype for the species is designated, and a summarized bibliography and illustrations including male genitalia are provided for the first time.