Assessing the stand size of bay trees (Persea spp.) after exposure to laurel wilt disease in a North Florida Preserve

A. Rossi, Christopher Bentzien
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While a signifi cantly higher percent (76%) of swampbay trees exhibited signs of laurel wilt disease compared to redbay trees (62%); redbay had more of its canopy damaged by the disease (41% vs. 32% for redbay vs. swampbay respectively); this resulted in a signifi cantly smaller stem diameter for P. borbonia compared to swampbay, both species are experiencing signifi cant declines due to the disease. Both species exhibited a Type III survivorship curve in which the vast majority of individuals were in the smallest size class (average stem diameter was only 2.5 and 3.6 cm for redbay and swampbay respectively). Although traditionally, population age (or size) structure that is heavily biased toward younger or smaller size classes suggests that the population is likely to expand in the future, for these bay trees high mortality rate due to beetle/fungal infestation of larger size classes is responsible for this trend; the smallest size classes are largely free from beetle infestation and laurel wilt disease because the stem diameter is likely insuffi cient to support beetle development. Results from this study suggest that swampbay is also highly susceptible to laurel wilt disease and its populations are likely to exhibit a similar (albeit slower) decline in Florida’s wetland and mesic ecosystems. Research Article Assessing the stand size of bay trees (Persea spp.) after exposure to laurel wilt disease in a North Florida Preserve Anthony M Rossi* and Christopher Bentzien Department of Biology, University of North Florida, Jacksonville, Florida, USA *Address for Correspondence: Anthony M Rossi, Department of Biology, University of North Florida, Jacksonville, Florida, 32224, USA, Tel: (904) 620-1934; (904)-620-2830; Email: arossi@unf.edu Submitted: 09 May 2019 Approved: 03 June 2019 Published: 04 June 2019 Copyright: © 2019 Rossi AM, 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 How to cite this article: Rossi AM, Bentzien C. Assessing the stand size of bay trees (Persea spp.) after exposure to laurel wilt disease in a North Florida Preserve. J Plant Sci Phytopathol. 2019; 3: 042-049. https://doi.org/10.29328/journal.jpsp.1001030 Introduction Laurel wilt disease is a generally fatal infection caused by a non-native fungus (Raffaelea lauricola) that is spread from tree to tree by the exotic redbay ambrosia beetle (RAB) (Xyleborus glabratus). Trees are inoculated with the fungus during gallery formation by adult beetles; females lay eggs in these galleries and the beetle larvae feed on the fungal hyphae eventually fungal growth plugs the water-conducting xylem vessels of the plant causing the leaves to brown and wilt; typically, the tree will eventually die from the infection [1,2]. Unlike leaf abscission in which the leaves develop a characteristic abscission zone and change color (as chlorophyll production ceases and secondary pigments such as carotenoids and anthocyanins become visible) and they are retained on the stem by the vascular tissue until wind or abrasion causes them to drop, laurel wilt kills the leaf by restricting its water supply causing the leaf to brown and die on the stem [3]. Infected trees respond to the fungus by producing tyloses and gums, which may reduce the spread of inections thoughout the plant, but they also inhibit the ability of the vessels to transport water [2]. For instance, tyloses are ballon-like swellings produced by parenchyma cells that project though pit cavities Assessing the stand size of bay trees (Persea spp.) after exposure to laurel wilt disease in a North Florida Preserve Published: June 04, 2019 043 in the vessel wall and partially or completely obstruct the vessel [3]. Progression of the disease is easily recognizable by the whole-sale browning of leaves that remain attached to the affected branches as well as the appearance of frass (referred to as sawdust tubes) created by beetle feeding which forms plugs or sticks visible on the external surface of the trees. Xyleborus glabratus transports the fungus in its mycangia, which are small specialized pouches that are associated with the mandibles [2,4]. Recent studies comparing fungi collected from the mycangia of RAB using molecular data and culturing medium that is selective to members of the Ophiostomatales, including R. lauricola, are consistent with populations from Southeast Asia [4,5]. Speci ically, fungal isolates from the mycangia of X. glabratus populations trapped in U.S. were similar to those from Taiwan and Japan; moreover, bacterial communities from X. glabratus mycangia collected in the U.S. were distinctly different from those of native species of xyleborine beetles, but similar to those from Asia [4,6]. These results are consistent with the hypothesis that within the U.S. both the beetle and its fungal symbionts were introduced into the U.S. from Asia; most likely via wooden pallets or packing materials that entered through Port Wentworth, which is part of the greater Savannah, Georgia metropolitan area, in the late 1990’s or early 2000’s [7-10]. Not surprisingly, introduced laurel species such as camphortree (Cinnamomum camphora), which are susceptible to laurel wilt, but are indigenous to southeast Asia and whose native distribution overlaps with that of RAB, have developed resistance to the disease [10,11]. For instance, camphor tree, which has become naturalized in the U.S., has shown signs of laurel wilt disease (e.g. browning of leaves and branch dieback), but affected trees typically recover from laurel wilt disease [10]. Laurel wilt has the potential to cause both ecological and economic damage because the laurel family of trees includes both non-commercial native trees such as redbay (Persea borbonia) and swampbay (P. palustris), but also economically important species such as avocado (P. americana). While loss of a commercial product such as avocado is relatively easy to estimate, ecological impacts of declining native bay trees is more problematic; redbay in particular is an important food plant for native animals such as the Palamedes swallowtail butter ly [5,12,13]. In the approximately two decades since its introduction to the U.S. laurel wilt disease has spread rapidly throughout the Southeastern U.S., especially along its primary east-coast interstate highway (I-95) which likely facilitates spread of the beetle and fungus via commercial deliveries along this busy traf ic corridor [14]. During 2005 RAB and the fungus were irst reported causing laurel wilt disease in redbay trees in north Florida from Timucuan Ecological and Historic Preserve, Duval County which includes the Jacksonville metropolitan region [5,8]. Fraedrich et al. [15], conducted a detailed study of redbay trees with a dbh > 2.5 cm within the preserve and found that percentage of cumulative mortality increased from approximately 10 to over 90% during a 16-month period and larger diameter trees died more quickly than those from smaller size class. Additionally, a 2008 study conducted in St. Johns County (just south of Duval County) of the state-listed endangered laurel tree, pondspice (Litsea aestivalis), reported 85% of the pondspice trees were already dead or dying from the disease [16]. Although southward spread of the beetle and the concomitant fungus was estimated at 20-30 miles (32-48 km) per year; the beetle was detected in central Florida coastal counties including Indian River and Brevard counties in 2005 and 2006 respectively and south Florida counties including Dade (which includes the Miami metropolitan area) in 2010 some ive years sooner than predicted [9,14]. Goals of this project were: 1) determine the extent of the disease affecting the two most common species of bay trees (i.e. P. borbonia and P. palustris) on the University of North Florida (UNF) campus (as part of the university’s land management strategy); 2) determine if the tree is infesting swampbay as well as redbay; 3) determine the effect of the disease on tree survival and size class. Assessing the stand size of bay trees (Persea spp.) after exposure to laurel wilt disease in a North Florida Preserve Published: June 04, 2019 044 Methods","PeriodicalId":93470,"journal":{"name":"Journal of plant science and phytopathology","volume":" ","pages":""},"PeriodicalIF":0.0000,"publicationDate":"2019-06-04","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":"0","resultStr":null,"platform":"Semanticscholar","paperid":null,"PeriodicalName":"Journal of plant science and phytopathology","FirstCategoryId":"1085","ListUrlMain":"https://doi.org/10.29328/JOURNAL.JPSP.1001030","RegionNum":0,"RegionCategory":null,"ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":null,"EPubDate":"","PubModel":"","JCR":"","JCRName":"","Score":null,"Total":0}
引用次数: 0

Abstract

Although laurel wilt disease was fi rst reported in the United States in 2002 from redbay trees (Persea borbonia) around Savannah, Georgia it has rapidly spread throughout the southeastern coastal plain including Georgia and Florida. In the current study, transects were used to assess the spread and impact of the disease on two native bay trees redbay (P. borbonia) and swampbay (P. palustris) from north Florida in a semi-naturalized ecological preserve. Although tree size and mortality rates have been reported previously, this study provides the fi rst size-based static life tables for both species. While a signifi cantly higher percent (76%) of swampbay trees exhibited signs of laurel wilt disease compared to redbay trees (62%); redbay had more of its canopy damaged by the disease (41% vs. 32% for redbay vs. swampbay respectively); this resulted in a signifi cantly smaller stem diameter for P. borbonia compared to swampbay, both species are experiencing signifi cant declines due to the disease. Both species exhibited a Type III survivorship curve in which the vast majority of individuals were in the smallest size class (average stem diameter was only 2.5 and 3.6 cm for redbay and swampbay respectively). Although traditionally, population age (or size) structure that is heavily biased toward younger or smaller size classes suggests that the population is likely to expand in the future, for these bay trees high mortality rate due to beetle/fungal infestation of larger size classes is responsible for this trend; the smallest size classes are largely free from beetle infestation and laurel wilt disease because the stem diameter is likely insuffi cient to support beetle development. Results from this study suggest that swampbay is also highly susceptible to laurel wilt disease and its populations are likely to exhibit a similar (albeit slower) decline in Florida’s wetland and mesic ecosystems. Research Article Assessing the stand size of bay trees (Persea spp.) after exposure to laurel wilt disease in a North Florida Preserve Anthony M Rossi* and Christopher Bentzien Department of Biology, University of North Florida, Jacksonville, Florida, USA *Address for Correspondence: Anthony M Rossi, Department of Biology, University of North Florida, Jacksonville, Florida, 32224, USA, Tel: (904) 620-1934; (904)-620-2830; Email: arossi@unf.edu Submitted: 09 May 2019 Approved: 03 June 2019 Published: 04 June 2019 Copyright: © 2019 Rossi AM, 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 How to cite this article: Rossi AM, Bentzien C. Assessing the stand size of bay trees (Persea spp.) after exposure to laurel wilt disease in a North Florida Preserve. J Plant Sci Phytopathol. 2019; 3: 042-049. https://doi.org/10.29328/journal.jpsp.1001030 Introduction Laurel wilt disease is a generally fatal infection caused by a non-native fungus (Raffaelea lauricola) that is spread from tree to tree by the exotic redbay ambrosia beetle (RAB) (Xyleborus glabratus). Trees are inoculated with the fungus during gallery formation by adult beetles; females lay eggs in these galleries and the beetle larvae feed on the fungal hyphae eventually fungal growth plugs the water-conducting xylem vessels of the plant causing the leaves to brown and wilt; typically, the tree will eventually die from the infection [1,2]. Unlike leaf abscission in which the leaves develop a characteristic abscission zone and change color (as chlorophyll production ceases and secondary pigments such as carotenoids and anthocyanins become visible) and they are retained on the stem by the vascular tissue until wind or abrasion causes them to drop, laurel wilt kills the leaf by restricting its water supply causing the leaf to brown and die on the stem [3]. Infected trees respond to the fungus by producing tyloses and gums, which may reduce the spread of inections thoughout the plant, but they also inhibit the ability of the vessels to transport water [2]. For instance, tyloses are ballon-like swellings produced by parenchyma cells that project though pit cavities Assessing the stand size of bay trees (Persea spp.) after exposure to laurel wilt disease in a North Florida Preserve Published: June 04, 2019 043 in the vessel wall and partially or completely obstruct the vessel [3]. Progression of the disease is easily recognizable by the whole-sale browning of leaves that remain attached to the affected branches as well as the appearance of frass (referred to as sawdust tubes) created by beetle feeding which forms plugs or sticks visible on the external surface of the trees. Xyleborus glabratus transports the fungus in its mycangia, which are small specialized pouches that are associated with the mandibles [2,4]. Recent studies comparing fungi collected from the mycangia of RAB using molecular data and culturing medium that is selective to members of the Ophiostomatales, including R. lauricola, are consistent with populations from Southeast Asia [4,5]. Speci ically, fungal isolates from the mycangia of X. glabratus populations trapped in U.S. were similar to those from Taiwan and Japan; moreover, bacterial communities from X. glabratus mycangia collected in the U.S. were distinctly different from those of native species of xyleborine beetles, but similar to those from Asia [4,6]. These results are consistent with the hypothesis that within the U.S. both the beetle and its fungal symbionts were introduced into the U.S. from Asia; most likely via wooden pallets or packing materials that entered through Port Wentworth, which is part of the greater Savannah, Georgia metropolitan area, in the late 1990’s or early 2000’s [7-10]. Not surprisingly, introduced laurel species such as camphortree (Cinnamomum camphora), which are susceptible to laurel wilt, but are indigenous to southeast Asia and whose native distribution overlaps with that of RAB, have developed resistance to the disease [10,11]. For instance, camphor tree, which has become naturalized in the U.S., has shown signs of laurel wilt disease (e.g. browning of leaves and branch dieback), but affected trees typically recover from laurel wilt disease [10]. Laurel wilt has the potential to cause both ecological and economic damage because the laurel family of trees includes both non-commercial native trees such as redbay (Persea borbonia) and swampbay (P. palustris), but also economically important species such as avocado (P. americana). While loss of a commercial product such as avocado is relatively easy to estimate, ecological impacts of declining native bay trees is more problematic; redbay in particular is an important food plant for native animals such as the Palamedes swallowtail butter ly [5,12,13]. In the approximately two decades since its introduction to the U.S. laurel wilt disease has spread rapidly throughout the Southeastern U.S., especially along its primary east-coast interstate highway (I-95) which likely facilitates spread of the beetle and fungus via commercial deliveries along this busy traf ic corridor [14]. During 2005 RAB and the fungus were irst reported causing laurel wilt disease in redbay trees in north Florida from Timucuan Ecological and Historic Preserve, Duval County which includes the Jacksonville metropolitan region [5,8]. Fraedrich et al. [15], conducted a detailed study of redbay trees with a dbh > 2.5 cm within the preserve and found that percentage of cumulative mortality increased from approximately 10 to over 90% during a 16-month period and larger diameter trees died more quickly than those from smaller size class. Additionally, a 2008 study conducted in St. Johns County (just south of Duval County) of the state-listed endangered laurel tree, pondspice (Litsea aestivalis), reported 85% of the pondspice trees were already dead or dying from the disease [16]. Although southward spread of the beetle and the concomitant fungus was estimated at 20-30 miles (32-48 km) per year; the beetle was detected in central Florida coastal counties including Indian River and Brevard counties in 2005 and 2006 respectively and south Florida counties including Dade (which includes the Miami metropolitan area) in 2010 some ive years sooner than predicted [9,14]. Goals of this project were: 1) determine the extent of the disease affecting the two most common species of bay trees (i.e. P. borbonia and P. palustris) on the University of North Florida (UNF) campus (as part of the university’s land management strategy); 2) determine if the tree is infesting swampbay as well as redbay; 3) determine the effect of the disease on tree survival and size class. Assessing the stand size of bay trees (Persea spp.) after exposure to laurel wilt disease in a North Florida Preserve Published: June 04, 2019 044 Methods
北佛罗里达保护区月桂枯萎病暴露后月桂树(Persea spp.)林分大小的评估
虽然桂冠枯萎病最早是2002年在美国乔治亚州萨凡纳附近的红杉树(Persea borbonia)上报道的,但它已经迅速蔓延到包括乔治亚州和佛罗里达州在内的东南沿海平原。在本研究中,采用样带方法评估了该病害在佛罗里达州北部半自然生态保护区的两种本地月桂树红月桂(P. borbonia)和沼泽月桂树(P. palustris)上的传播和影响。尽管之前已经报道过树木的大小和死亡率,但这项研究首次为这两个物种提供了基于大小的静态生命表。而与红杉树(62%)相比,沼泽湾树(76%)表现出月桂枯萎病的迹象;红杉的冠层受病害的破坏较多(41%比32%);这导致P. borbonia的茎直径明显小于swampbay,这两个物种都因疾病而经历了显着的下降。两个物种均表现为III型生存曲线,绝大多数个体在最小级(红湾和沼泽湾的平均茎粗分别仅为2.5和3.6 cm)。虽然传统上,种群年龄(或大小)结构严重倾向于年轻或较小的种群,这表明种群可能在未来扩大,但对于这些月桂树来说,由于甲虫/真菌侵染的高死亡率是造成这一趋势的原因;最小的种类基本上没有甲虫侵扰和月桂枯萎病,因为茎直径可能不足以支持甲虫的发育。这项研究的结果表明,swampbay也极易受到月桂枯萎病的影响,其种群可能在佛罗里达州的湿地和mesic生态系统中表现出类似的(尽管速度较慢)下降。研究文章:北佛罗里达自然保护区月桂树(Persea spp.)暴露于月桂树枯萎病后的林分大小评估Anthony M Rossi*和Christopher Bentzien北佛罗里达大学生物系,佛罗里达州杰克逊维尔,美国*通讯地址:Anthony M Rossi,北佛罗里达大学生物系,佛罗里达州杰克逊维尔,32224,美国,电话:(904)620-1934;(904) 620 - 2830;邮箱:arossi@unf.edu提交时间:2019年5月9日批准时间:2019年6月3日发布时间:2019年6月4日版权所有:©2019 Rossi AM, et al。这是一篇在知识共享署名许可下发布的开放获取文章,该许可允许在任何媒介上不受限制地使用、分发和复制,前提是原始作品被正确引用。如何引用这篇文章:Rossi AM, Bentzien C.在北佛罗里达自然保护区评估月桂枯萎病后月桂树(Persea spp.)的林分大小。植物化学学报,2019;3: 042 - 049。https://doi.org/10.29328/journal.jpsp.1001030介绍月桂枯萎病是由一种非本地真菌(Raffaelea lauricola)引起的一种通常致命的感染,这种真菌由外来的红月桂甲虫(Xyleborus glabratus)在树到树之间传播。成虫在廊道形成过程中给树木接种真菌;雌性甲虫在这些通道中产卵,幼虫以真菌菌丝为食,最终真菌的生长堵塞了植物的木质部导管,导致叶子变成褐色和枯萎;通常情况下,树最终会死于感染[1,2]。与叶子脱落不同的是,叶子形成一个典型的脱落区并改变颜色(当叶绿素停止产生,次生色素如类胡萝卜素和花青素变得可见),它们被维管组织保留在茎上,直到风或磨损导致它们脱落,月桂枯萎通过限制其水分供应而杀死叶子,导致叶子变成棕色并死在茎上。受感染的树木对真菌的反应是产生tylose和树胶,这可能会减少感染在整个植物中的传播,但它们也会抑制血管运输水的能力。例如,tyloss是由薄壁细胞产生的气球状肿胀,这些细胞通过坑腔突出。在北佛罗里达保护区暴露于月桂枯萎病后,评估月桂树(Persea spp.)的林分大小,并部分或完全阻塞血管壁。这种疾病的进展很容易通过附着在受影响树枝上的叶子的整体褐变以及甲虫进食产生的草(称为锯末管)的外观来识别,这些草在树木的外表面形成可见的塞或棒。Xyleborus glabratus在其mycangia中运输真菌,mycangia是与下颌骨相关的小型专用袋[2,4]。 最近的研究比较了从RAB菌丝中收集的真菌,使用分子数据和对Ophiostomatales成员(包括R. lauricola)有选择性的培养基,结果与来自东南亚的群体一致[4,5]。其中,从美国捕获的秃毛天牛种群菌丝中分离出的真菌与台湾和日本相似;此外,在美国采集到的毛缕甲(X. glabratus mycangia)的细菌群落与本土木leborine甲虫有明显差异,但与亚洲的相似[4,6]。这些结果与假设一致,即在美国,甲虫和它的真菌共生体都是从亚洲引入美国的;最有可能是通过木制托盘或包装材料进入温特沃斯港,这是大萨凡纳,乔治亚州大都市区的一部分,在20世纪90年代末或21世纪初[7-10]。毫不奇怪,引进的桂树物种,如樟树(Cinnamomum camphora),对月桂枯萎病敏感,但原产于东南亚,其本地分布与RAB重叠,已经对该疾病产生了抗性[10,11]。例如,在美国已经归化的樟树已经显示出月桂枯萎病的迹象(例如叶子变褐和树枝枯死),但受影响的树木通常会从月桂枯萎病中恢复过来。月桂枯萎病有可能造成生态和经济损害,因为月桂科树木既包括非商业的本地树木,如红月桂(Persea borbonia)和沼泽月桂(P. palustris),也包括经济上重要的物种,如鳄梨(P. americana)。虽然像鳄梨这样的商业产品的损失相对容易估计,但本地月桂树数量下降的生态影响更成问题;尤其是红月桂,是Palamedes swallowtail butter等本地动物的重要食物植物[5,12,13]。在传入美国的大约20年里,月桂枯萎病在美国东南部迅速蔓延,特别是沿着主要的东海岸州际公路(I-95),这可能促进了甲虫和真菌在繁忙的交通走廊[14]上的商业运输的传播。2005年,RAB和真菌首次被报道在佛罗里达州北部的杜瓦尔县Timucuan生态和历史保护区引起月桂枯萎病,该保护区包括杰克逊维尔大都市区[5,8]。freedrich et al. b[15]对保护区内胸径为2.5 cm的红杉树进行了详细研究,发现在16个月的时间内,累积死亡率从约10%增加到90%以上,直径较大的树木比尺寸较小的树木死亡得更快。此外,2008年在圣约翰县(就在杜瓦尔县的南部)进行的一项研究表明,国家列出的濒临灭绝的月桂树,塘香树(Litsea aestivalis), 85%的塘香树已经死亡或死于疾病bbb。虽然据估计,这种甲虫和伴随的真菌每年向南传播20-30英里(32-48公里);这种甲虫分别于2005年和2006年在佛罗里达中部沿海县(包括印度河县和布里瓦德县)和2010年在佛罗里达南部县(包括戴德县(包括迈阿密市区))被发现,比预测的时间提前了大约5年[9,14]。该项目的目标是:1)确定影响北佛罗里达大学(UNF)校园内两种最常见的月桂树(即P. borbonia和P. palustris)的疾病程度(作为该大学土地管理战略的一部分);2)确定该树是否同时侵扰沼泽湾和红湾;3)确定病害对树木成活率和大小等级的影响。北佛罗里达保护区月桂枯萎病暴露后月桂树(Persea spp.)的林分大小评估发表:June 04, 2019
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