Native and introduced range surveys for egg parasitoids of Agrilus auroguttatus,an invasive pest of native oaks in California

The invasive goldspotted oak borer, Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), has caused extensive mortality to indigenous oaks in southern California. This woodborer is native to southern Arizona where low population densities may, in part, be due to the presence of co-evolved natural enemies. Surveys were conducted in Arizona and California from June to August 2013 by deploying sentinel egg masses in an attempt to attract, collect and identify potential egg parasitoids of this beetle that could be used for a classical biological control programme in California. In total, 18,052 A. auroguttatus eggs were deployed throughout the native and introduced range. Parasitoids did not attack any eggs deployed in Arizona or California. The inability to detect egg parasitoids could be explained by the following: (1) host-specific egg parasitoids of A. auroguttatus do not exist, (2) surveying time did not coincide with peak activity of egg parasitoids or (3) surveying methods were insufficient at detecting egg parasitoids.     

Can the destruction of California’s oak woodlands be prevented? Potential for biological control of the goldspotted oak borer, <Emphasis Type="Italic">Agrilus auroguttatus</Emphasis>

The goldspotted oak borer (GSOB), Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), is an introduced and aggressive phloem/wood borer infesting native oaks in southern California. Elevated levels of oak mortality have occurred continually for the last nine years on three oak species in San Diego Co., California, USA. Biological control is being assessed as an option for long-term and widespread management of the invasive population of GSOB. Foreign exploration in the native ranges of GSOB and a related sibling species (Agrilus coxalis Waterhouse) was conducted to determine life history information, to assess the natural enemy complex, and to collect specimens for molecular analyses that could help to identify the area of origin of California’s introduced population. Two species of parasitoids, Calosota elongata Gibson (Eupelmidae) and Atanycolus simplex Cresson (Braconidae), were discovered with GSOB populations in Arizona and California. No insect natural enemies were found with populations of A. coxalis in southern Mexico. However, Quercus conzatti Trel. and Quercus peduncularis Nee in Oaxaca and Chiapas, respectively, were recorded as the first known hosts of A. coxalis. A comparative analysis of our understanding of the natural enemy complexes for other pestiferous Agrilus with that of GSOB suggests that more effort should be directed at uncovering potential egg parasitoids and microbial pathogens of GSOB. Analyses of mitochondrial and nuclear ribosomal DNA (rDNA) revealed that the California population of GSOB was more similar to the Arizona population. Specimens of A. coxalis from southern Mexico were confirmed as a separate species. Additional surveys and sampling are needed across the complete native range of the GSOB species complex to develop a comprehensive inventory of parasitoid species that could be considered for use in a classical biological control program in California and to delineate the area of origin of California’s population.     

Environmental suitability for Agrilus auroguttatus (Coleoptera: Buprestidae) in Mexico using MaxEnt and database records of four Quercus (Fagaceae) species

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Goldspotted oak borer effects on tree health and colonization patterns at six newly‐established sites

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Foliar nutrients explain goldspotted oak borer,Agrilus auroguttatus,adult feeding preference among four California oak species

Adults of the invasive goldspotted oak borer, Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), consumed foliar weight in no‐choice feeding tests of, in descending order, California black oak Quercus kelloggii Newb., Engelmann oak, Quercus engelmannii Greene, coast live oak, Quercus agrifolia Née, and canyon live oak, Quercus chrysolepis Liebm. (Fagaceae). Furthermore, significantly more foliar area was consumed of Q. kelloggii than of Q. chrysolepis. In dual‐choice feeding tests with isolated leaf disks, A. auroguttatus consumed significantly more foliar weight and area of Q. kelloggii relative to the other three oak species, and more foliar weight of Q. agrifolia than of Q. chrysolepis. In dual‐choice feeding tests with leaves on small branches, A. auroguttatus consumed more foliar weight of Q. kelloggii than of Q. engelmannii and Q. agrifolia. Thus, multiple experiments suggested that adults of A. auroguttatus preferred the foliage of Q. kelloggii over that of the other three oak species, and among the other three species they did not appear to have a strong feeding preference. Factor analysis reduced the quantities of 13 foliar nutrients into two new variables (factor 1 and factor 2). Factor 1 was weighted heavily on the quantities of nitrogen, sulfur, phosphorus, potassium, zinc, and copper, whereas factor 2 was weighted heavily on the quantities of zinc, iron, and aluminum. Factor 1 varied by oak species, with Q. kelloggii having a higher factor 1 nutrient content than the other three species. Factor 2 response was higher in Q. kelloggii, Q. agrifolia, and Q. engelmannii than in Q. chrysolepis. The collective effects of four macronutrients (nitrogen, sulfur, phosphorus, and potassium) and two micronutrients (zinc and copper) suggest that these might be the nutrients directing preferential feeding of A. auroguttatus adults on the foliage of Q. kelloggii. Leaf toughness might also play an important role in feeding preference. Female A. auroguttatus did not show an ovipositional preference among the four oak species.     

Suitability of native and ornamental oak species in California for Agrilus auroguttatus

Goldspotted oak borer, Agrilus auroguttatus Schaeffer (Coleoptera: Buprestidae), is a new invasive species in southern California, USA. The extent of the host range of this insect is not known, but this knowledge will have a major impact on assessment of the risks that this pest poses to oaks [Quercus spp. (Fagaceae)]. We conducted laboratory tests to determine the potential suitability of native and ornamental oak species for larvae and adults of A. auroguttatus. We infested 179 cut logs (from 163 different trees) with eggs or larvae, measured neonate survival and, after 5 months, counted feeding galleries, and noted the proportion of galleries with late instars. Initial larval survival was generally high when larvae penetrated the phloem (range: 62–98% among oak species), and low by the time larvae began to feed at the phloem/xylem interface (range: 0–25% among oak species). The majority of larvae that established a visible feeding gallery survived to the fourth instar (total of 73% for all oak species). Larval galleries were established with greater frequency in red oaks (Section Lobatae) compared with other oaks (19 vs. 7 or 4%). All red oaks tested (Q. agrifolia Née, Q. kelloggii Newberry, and Q. wislizeni A. DC.) were likely suitable hosts for larvae. Larvae were apparently able to feed on some of the other oaks (Q. chrysolepis Leibmann, Q. suber L., Q. lobata Née, and Q. douglasii Hook & Arn), although it remains unclear whether these species would be preferred hosts under natural conditions. Adult longevity and fecundity varied little by species of oak foliage fed to adults. The host range of A. auroguttatus is likely limited by suitability of oak species for the larval rather than the adult life stage. Results support published field observations that red oaks are more suitable hosts than white oaks.     

Another look at Hymenoxys subgenus Plummera (Asteraceae: Heliantheae: Gaillardiinae) from Arizona and New Mexico

The recent merger ofPlummera intoHymenoxys is followed here. The two previously recognized species ofPlummera are here treated as a single polymorphic species ofHymenoxys, H. ambigens, consisting of three varieties: var.ambigens from the Mescal, Santa Teresa, and Pinaleno Mountains of southeastern Arizona; var.floribunda from the Chiricahua, Dos Cabezas, Dragoon, Little Dragoon, and Mule Mountains of southeastern Arizona; and var.neomexicana described here from the Animas and Peloncillo Mountains of southwestern New Mexico.     

Inadvertent biological control: an Australian thrips killing an invasive New Zealand tree in California

Transport hubs of international trade and tourism are sites of unprecedented long-distance dispersal of species and novel ecological interactions. In cases of invasive plants released from their specialist natural enemies, novel interactions with both resident enemies and new arrivals can accumulate and potentially reduce weed competitiveness. I present here one dramatic example of this, where an invasive woody weed in southern California is being rapidly controlled by an accidentally introduced genus-specialist herbivorous insect. The New Zealand native shrub/small tree, Myoporum laetum, is a long-time popular ornamental plant in California and has become an invasive woody weed. In 2005, a Myoporum-specific thrips, Klambothrips myopori, was discovered (and described) in California feeding on M. laetum leaves. Several searches have failed to find K. myopori in New Zealand and a population has recently been discovered in Tasmania, Australia, feeding on Myoporum insulare. In 5 years, K. myopori has killed off about half of southern Californian M. laetum with almost all surviving individuals being gradually defoliated. Inadequate border biosecurity has resulted in inadvertent biological control, in a rapid timeframe, caused by a novel enemy. Unfortunately, K. myopori has subsequently been accidentally transported from California to Hawaii where it is now killing off Hawaiian native Myoporum sandwicense. Transport hubs can both connect weeds with natural enemies and disperse those enemies more widely.     

Molecular evidence for a single genetic clone of invasive Arundo donax in the United States

Arundo donax (giant reed) is an aggressive invasive weed of riparian habitats throughout the southern half of the United States from California to Maryland. Native to Asia, the species is believed to have been initially introduced into North America from the Mediterranean region although subsequent introductions were from multiple regions. To provide insight into the potential for biological control of A. donax, genetic variation in plants sampled from a wide geographical area in the United States was analyzed using Sequence Related Amplification Polymorphism (SRAP) and transposable element (TE)-based molecular markers. Invasive individuals from 15 states as well as four populations in southern France were genetically fingerprinted using 10 SRAP and 12 TE-based primer combinations. With the exception of simple mutations detected in four plants, A. donax exhibited a single multilocus DNA fingerprint indicating a single genetic clone. The genetic uniformity of invasive A. donax suggests that classical biological control of the species could be successful. A lack of genetic diversity in the invaded range simplifies identification of native source populations to search for natural enemies that could be used as biocontrol agents.     

A revision of the salvia dorrii complex (Lamiaceae)

Expressions of theSalvia dorrii (Kellogg) Abrams complex, a member of subg.Audibertia Benth., are distributed over much of the western United States. The complex consists of two species,S. dorrii andS. pachyphylla Epling ex Munz.Salvia dorrii is divided into subsp.dorrii and subsp.mearnsii (Britton) McClintock, the latter endemic to central Arizona. Subspeciesdorrii is composed of four varieties: var.carnosa (Douglas ex Greene) Cronq., a large-leaved erect shrub of Washington and Idaho southward through Oregon into extreme north-central California: var.dorrii, a small-leaved erect shrub of southern Oregon and Idaho southward through the Great Basin of Nevada and western Utah to southeastern California and northern Arizona: var.pilosa (A. Gray) Strachan & Reveal, a small-leaved erect shrub differing from var.dorrii by pilose bracts and calyx, in southern California and western Arizona with disjunct populations in the Lahontan Basin of northwestern Nevada and northestern California; and var.clokeyi Strachan, var. nov., a low mat-forming subshrub of the high mountains of Clark Co., Nevada.Salvia pachyphylla ranges from southern California southward to the high mountains of Baja California Norte, Mexico, and eastward across extreme southern Nevada to (as a disjunct) northeastern Arizona. Each entity is described in detail, mapped and illustrated.     

Comparative morphometric and chemical analyses of phenotypes of two invasive ambrosia beetles (Euwallacea spp.) in the United States

The polyphagous shot hole borer (PSHB), Euwallacea sp., was first detected in 2003 in Los Angeles County, California, USA. Recently, this invasive species has become a major pest of many hardwood trees in urban and wildland forests throughout southern California. PSHB is nearly identical in morphology and life history to the tea shot hole borer (TSHB), Euwallacea fornicatus, an invasive pest of hardwoods in Florida, USA and many other parts of the world. However, molecular studies have suggested that the taxa are different species. We conducted morphometric and chemical analyses of the phenotypes of Euwallacea sp. collected in southern California (Los Angeles County) and E. fornicatus collected in Florida (Miami‐Dade County). Our analyses indicated that PSHB has 3 larval instars. The third larval instar was separated from the first 2 instars by head capsule width with 0 probability of misclassification. The body length, head width, and pronotal width of PSHB adult males were significantly less than those of females. Head width and pronotal width of female PSHB were significantly less than those of female TSHB. In contrast, body length, and ratio of body length to pronotal width of female PSHB were significantly greater than those of female TSHB. However, females of these 2 species could not be separated completely by these 4 measurements because of the overlapping ranges. Cuticular hydrocarbons detected in both species were exclusively alkanes (i.e., n‐alkanes, monomethylalkanes, dimethylalkanes, and trimethylalkanes). Cuticular hydrocarbon profiles of PSHB males and females were similar, but they both differed from that of TSHB females. Cuticular hydrocarbons of PSHB were predominantly internally branched dimethylalkanes with backbones of 31 and 33 carbons, whereas cuticular hydrocarbons of TSHB females were dominated by internally branched monomethylalkanes and dimethylalkanes with backbones of 28 and 29 carbons. Multiple compounds within these classes appear to be diagnostic for PSHB and TSHB, respectively.     

Variability in essential oils and needle resin canals of Pinus longaeva from Eastern California and Western Nevada in relation to other members of subsection Balfourianae

Mono- and sesquiterpenoids from wood and leaves of Pinus longaeva trees collected in various localities of eastern California and western Nevada were quantitatively analyzed by GLC. Concurrently the number and size of resin canals in foliage of the same trees and as well as Pinus aristata trees from Arizona, Colorado and New Mexico were determined. While some monoterpenes (mainly α-pinene and 3-carene) suggest that P. longaeva trees from the Panamint Mountains of California are P. aristata-like, the total chemical data and morphological data indicate that Panamint trees should be grouped with P. longaeva. A chemical latitudinal gradient is evidenced by the appearance of trees producing 3-carene in large amounts along the line joining the Panamint, Inyo and White Mountains of California. Pinus rzedowskii, well separated geographically and morphologically from the accepted members of subsection Balfourianae, is shown to be sufficiently distinct chemically, as to leave its suggested membership in the subsection an open question.     

Optimization of sampling methods for within-tree populations of red oak borer, Enaphalodes rufulus (Haldeman) (Coleoptera: Cerambycidae)

In the Ozark Mountains of northern Arkansas and southern Missouri, an oak decline event, coupled with epidemic populations of red oak borer (Enaphalodes rufulus Haldeman), has resulted in extensive red oak (Quercus spp., section Lobatae) mortality. Twenty-four northern red oak trees, Quercus rubra L., infested with red oak borer, were felled in the Ozark National Forest between March 2002 and June 2003. Infested tree boles were cut into 0.5-m sample bolts, and the following red oak borer population variables were measured: current generation galleries, live red oak borer, emergence holes, and previous generation galleries. Population density estimates from sampling plans using varying numbers of samples taken randomly and systematically were compared with total census measurements for the entire infested tree bole. Systematic sampling consistently yielded lower percent root mean square error (%RMSE) than random sampling. Systematic sampling of one half of the tree (every other 0.5-m sample along the tree bole) yielded the lowest values. Estimates from plans systematically sampling one half the tree and systematic proportional sampling using seven or nine samples did not differ significantly from each other and were within 25% RMSE of the "true" mean. Thus, we recommend systematically removing and dissecting seven 0.5-m samples from infested trees as an optimal sampling plan for monitoring red oak borer within-tree population densities. This optimal sampling plan should allow for collection of acceptably accurate within-tree population density data for this native wood-boring insect and reducing labor and costs of dissecting whole trees.     

Genetic diversity does not affect the invasiveness of fountain grass (Pennisetum setaceum) in Arizona, California and Hawaii

Pennisetum setaceum (Poaceae) is a perennial bunch grass that invaded the United States during the 20th century and is highly invasive in Hawaii, moderately invasive in Arizona, and not yet invasive in southern California. Pennisetum setaceum is apomictic, a condition that is normally associated with low genetic variation within populations, but even moderate levels of genetic variation among populations could account for differences in invasiveness. To determine whether genetic factors are causing the variable invasion success, we used Inter‐Simple Sequence Repeat markers (ISSRs) to examine genetic variation in populations from the three areas. Screening of 16 primers revealed no genetic variation within any population or between any geographical areas, a pattern consistent with complete apomixis. Variation in invasion success appears unrelated to genetic differences among populations. Differences in the seasonal timing of rainfall among the regions may be the cause of variable invasiveness of fountain grass. Alternatively, differences in timing of introduction or duration of lag phase may have limited invasiveness in Arizona and southern California.     

Inferring the invasion history of coral berry Ardisia crenata from China to the USA using molecular markers

Genetic comparisons between native and invasive populations of a species can provide insights into its invasion history information, which is useful for guiding management and control strategies. The coral berry Ardisia crenata was introduced to Florida last century as a cultivated ornament plant, and has since spread widely throughout the southern regions of the USA. Previously, the genetic variation among 20 natural populations of A. crenata across its distribution center in southern China was quantified using seven microsatellite markers. Here we expand on that work by additionally sampling individuals from four other native populations in Taiwan and Japan, and from five invasive populations in the USA. We also examined the results from one chloroplast intergenic spacer region (trnF-trnL) in all 29 populations. Our aim is to identify the invasion source and subsequent history of the species?? spread throughout the southern USA. We observed lower genetic diversity in the invasive populations based on both microsatellite and chloroplast markers. Our data show that the invasive populations can be clustered with native populations in southeastern China, inferring this region as the geographic origin of A. crenata cultivars invading the USA. We further classified invasive individuals into invasive I and invasive II clusters. Nantou in Taiwan and Xihu in mainland China are the most closely related populations to those, which identify the former as potential sources for host-specific control agents. Our results, combined with the known introduction records, suggest that A. crenata was first multiply introduced into Florida and then secondarily colonized Louisiana and Texas from Florida.     

Loss of microbial (pathogen) infections associated with recent invasions of the red imported fire ant Solenopsis invicta

Loss of natural enemies during colonization is a prominent hypothesis explaining enhanced performance of invasive species in introduced areas. Numerous studies have tested this enemy release hypothesis in a wide range of taxa but few studies have focused on invasive ants. We conducted extensive surveys for the presence of six microbes in recently established populations (California, Australia, New Zealand, Taiwan, and China) of the invasive fire ant Solenopsis invicta. These microbes include Wolbachia, two microsporidia (Kneallhazia solenopsae and Vairimorpha invictae) and three RNA viruses (SINV-1, -2 and -3), all of which previously have been reported in native South American populations of S. invicta. These surveys showed that the total number of enemy species is lower in the recently invaded areas compared with both South American and US populations. Only two microbes were found in any of these recently invaded areas: SINV-1 was detected in all surveyed populations except Australia and New Zealand, and SINV-2 was detected in California and Taiwan only. These results support the general prediction that invasive species lose many of their natural enemies during invasion. Further, the conspicuous absence of some of these microbes in these areas may result from strong selection against founders due to fitness costs associated with harboring detrimental infections rather than the alternative hypothesis that they simply were absent among the original founders. While the successful invasion of S. invicta in these recently invaded areas may be explained partly by the absence of natural enemies, other factors likely have been important as well.     

Beetles that live with ants (Carabidae,Pseudomorphini, Pseudomorpha Kirby, 1825): A revision of the santarita species group

The Western Hemisphere genus Pseudomorpha Kirby 1825 was last revised by Notman in 1925 based on only a few known species (22) and paltry few specimens (73); other authors have added an additional six species represented by 53 additional specimens since 1925. Baehr (1997) assigned three species from Australia to this genus, albeit in a new subgenus, Austropseudomorpha Baehr 1997. A recent study of collections from throughout the Americas (1757 specimens) has revealed numerous new species that can be arrayed across 19 species groups based on a suite of attributes, some used by Notman and others newly discovered. A taxonomic revision of the species contained in one of these species groups, santarita, is provided herein, as well as a distributional synopsis of the remaining 18 species groups. New species described herein are as follows, each with its type locality: Pseudomorpha huachineras p. n., Arroyo El Cocono, Sierra Huachinera, Sonora, México; P. patagonia sp. n., Madera Canyon, Santa Rita Mountains, Arizona; P.penablanca sp. n., Peña Blanca Lake, Arizona; P. pima sp. n., Madera Canyon (lower), Santa Rita Mountains, Arizona; P. santacruz sp. n., Madera Canyon, Santa Rita Mountains, Arizona; and P. santarita sp. n., Santa Rita Ranch, Santa Rita Mountains, Arizona.     

Modern pollen and vegetation relationships in the mountains of southern California,USA

The relationship between modern pollen assemblages and modern vegetation along two elevational transects within the Transverse and Peninsular Ranges of southern California, USA, is demonstrated using cluster analysis of the pollen data. Cluster analysis separates the Sonora Desert vegetation, Valley grassland/agricultural land and chaparral vegetation types on the San Jacinto Mountains transect. Chaparral is not easily separated on the San Bernardino Mountains transect, probably due to the presence of Quercus dumosa (scrub oak) there. The lower montane Quercus – Pinus (oak – pine) community is distinct from other forest types, and can be subdivided palynologically based upon relative importance of Quercus, Pinus and Cupressaceae [primarily Calocedrus decurrens (incense cedar)] pollen. Subdivisions include Quercus – Pinus – Cupressaceae, Quercus – Cupressaceae – Pinus and Quercus – Pinus assemblages. Higher elevation Pinus – Abies (pine – fir) and Pinus-dominated communities are also differentiated from one another, although the subalpine vegetation type only occurs on the San Bernardino Mountains transect. Though the study area presently straddles a transition between winter-wet and summer-wet climatic regimes, differences between the pollen assemblages in the two mountain ranges are minimal. Pollen assemblages from lower elevations document the effects of human activities, primarily agriculture, on the modern pollen rain of the region, with the occurrence of introduced citrus (Citrus sp.) and shade (Eucalyptus sp.) trees and weedy disturbance indicators (e.g., Brassicaceae).     

Acorns of invasive Northern Red Oak (<Emphasis Type="Italic">Quercus rubra</Emphasis>) in Europe are larval hosts for moths and beetles

In their first phase of expanding into new areas, invasive plants often take advantage of the inability of existing herbivores and pathogenic species to exploit them. However, in the longer term local enemies may adapt to using these invasive species as a food source. This study assesses the use of mature acorns of two oak species in Europe (the native Pedunculate Oak Quercus robur and the invasive Northern Red Oak Quercus rubra) by moths Cydia fagiglandana and Cydia splendana and beetles Curculio spp. We show that acorns of invasive oak species can be equally attractive to C. splendana but only partially so to C. fagiglandana where infestation rates where significantly lower (approximately half) compared to the native oak. The infestation by Curculio beetles of Northern Red Oak was marginal, less than 1% of the rate in the native oak species. The larval final weights did not differ significantly between host species, but emergence of C. splendana and Curculio spp. took significantly longer in acorns of Northern Red Oak. It is likely that C. fagiglandana and C. splendana have increased their niche breadths by exploiting invasive oak species and avoiding competition with the Curculio weevils. Furthermore, the occurrence of Northern Red Oak could stabilize food resources during years when native oak species have poor acorn crops.     

Genetic differentiation,speciation, and phylogeography of cactus flies (Diptera: Neriidae: Odontoloxozus) from Mexico and south‐western USA

Nucleotide sequences from the mitochondrial cytochrome c oxidase subunit I (COI) gene, comprising the standard barcode segment, were used to examine genetic differentiation, systematics, and population structure of cactus flies (Diptera: Neriidae: Odontoloxozus) from Mexico and south‐western USA. Phylogenetic analyses revealed that samples of Odontoloxozus partitioned into two distinct clusters: one comprising the widely distributed Odontoloxozus longicornis (Coquillett) and the other comprising Odontoloxozus pachycericola Mangan & Baldwin, a recently described species from the Cape Region of the Baja California peninsula, which we show is distributed northward to southern California, USA. A mean Kimura two‐parameter genetic distance of 2.8% between O. longicornis and O. pachycericola, and eight diagnostic nucleotide substitutions in the COI gene segment, are consistent with a species‐level separation, thus providing the first independent molecular support for recognizing O. pachycericola as a distinct species. We also show that the only external morphological character considered to separate adults of the two species (number of anepisternal bristles) varies with body size and is therefore uninformative for making species assignments. Analysis of molecular variance indicated significant structure among populations of O. longicornis from three main geographical areas, (1) Arizona, USA and Sonora, Mexico; (2) Santa Catalina Island, California, USA; and (3) central Mexico (Querétaro and Guanajuato), although widely‐separated populations from Arizona and Sonora showed no evidence of structure. A TCS haplotype network showed no shared haplotypes of O. longicornis among the three main regions. The potential roles of vicariance and isolation‐by‐distance in restricting gene flow and promoting genetic differentiation and speciation in Odontoloxozus are discussed. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 245–256.