Reassessment of the Species in the Euwallacea Fornicatus (Coleoptera: Curculionidae: Scolytinae) Complex After the Rediscovery of the “Lost” Type Specimen
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insects Article Reassessment of the Species in the Euwallacea Fornicatus (Coleoptera: Curculionidae: Scolytinae) Complex after the Rediscovery of the “Lost” Type Specimen Sarah M. Smith 1, Demian F. Gomez 2 , Roger A. Beaver 3, Jiri Hulcr 2 and Anthony I. Cognato 1,* 1 Department of Entomology, Michigan State University, East Lansing, MI 48824, USA 2 School of Forest Resources and Conservation, University of Florida, 136 Newins-Ziegler Hall, Gainesville, FL 32611, USA 3 161/2 Mu 5, Soi Wat Pranon, T. Donkaew, A. Maerim, Chiangmai 50180, Thailand * Correspondence: [email protected]; Tel.: +1-51-7432-2369 Received: 22 July 2019; Accepted: 19 August 2019; Published: 22 August 2019 Abstract: Ambrosia beetles of the Euwallacea fornicatus (Eichhoff, 1868) species complex are emerging tree pests, responsible for significant damage to orchards and ecosystems around the world. The species complex comprises seven described species, all of which are nearly identical. Given that the morphology-defined species boundaries have been ambiguous, historically, there has been much disagreement on species validity, which was compounded by the presumed loss of the type series of E. fornicatus. The species complex was recently reviewed using morphometrics to associate the type specimens to the clades delineated with molecular data under the assumption of the lost type series. We rediscovered a syntype of Xyleborus fornicatus, and reevaluated the species in the complex using morphometrics. We propose the following taxonomic changes to the species complex: Euwallacea fornicatus (=E. tapatapaoensis (Schedl, 1951); = E. whitfordiodendrus (Schedl, 1942)) syn. res.); E. fornicatior (Eggers, 1923) (=E. schultzei (Schedl, 1951) syn. nov.); E. kuroshio (Gomez and Hulcr, 2018) and E. perbrevis (Schedl, 1951) stat. res. These taxonomic changes shift the species name associated with the widely used common names for two taxa, namely: Euwallacea fornicatus should be used for the “Polyphagous Shot Hole Borer”, and E. perbrevis for the “Tea Shot Hole Borer clade a”. A lectotype is designated for X. fornicatus in order to stabilize the use of the name. Keywords: ambrosia beetle; invasive species; taxonomy; species delineation 1. Introduction Xyleborine ambrosia beetles (Coleoptera: Curculionidae: Scolytinae: Xyleborini) are among the most damaging and invasive organisms, with impacts ranging from economic to ecological. Ecological and biological features, such as fungus-farming, haplodiploidy, and a wide host range, make the Xyleborini one of the most successful colonizers [1]. With ~1160 recognized species, this is the most species-rich tribe of Scolytinae, including some of the most widespread species [2]. The Euwallacea fornicatus species complex comprises seven species described from Asia and Oceania, namely: E. fornicatus (Eichhoff, 1868) (Sri Lanka), E. fornicatior (Eggers, 1923) (Sri Lanka), E. whitfordiodendrus (Schedl, 1942) (Malaysia), E. schultzei (Schedl, 1951) (Philippines), E. perbrevis (Schedl, 1951b) (Philippines), E. tapatapaoensis (Schedl, 1951) (Samoa), and E. kuroshio Gomez and Hulcr, 2018 (Japan). This species complex is arguably one of the most challenging in scolytine systematics, because the species are nearly identical. Given the blurred species boundaries, scolytine taxonomists disagreed on the validity of these species (see Stouthamer et al. [3] for a detailed synopsis). Wood [4] Insects 2019, 10, 261; doi:10.3390/insects10090261 www.mdpi.com/journal/insects Insects 2019, 10, x FOR PEER REVIEW 2 of 12 Insectstaxonomists2019, 10, 261disagreed on the validity of these species (see Stouthamer et al. [3] for a detailed2 of 11 synopsis). Wood [4] considered E. schultzei, E. perbrevis, E. tapatapaoensis, and E. whitfordiodendrus to be synonyms of E. fornicatus, and later, Wood and Bright [5] placed E. fornicatior in synonymy, leaving considered E. schultzei, E. perbrevis, E. tapatapaoensis, and E. whitfordiodendrus to be synonyms of E. only E. fornicatus as valid. fornicatus, and later, Wood and Bright [5] placed E. fornicatior in synonymy, leaving only E. fornicatus Xyleborus fornicatus was described from an unknown number of specimens from Sri Lanka [6]. as valid. Eichhoff’s collection and types were deposited in the Zoological Museum in Hamburg, Germany Xyleborus fornicatus was described from an unknown number of specimens from Sri Lanka [6]. (UZHM), and were destroyed when the museum was bombed during World War II. Approximately Eichhoff’s collection and types were deposited in the Zoological Museum in Hamburg, Germany a dozen Eichhoff types were saved by K.E. Schedl, and now reside at the Naturhistorisches Museum (UZHM), and were destroyed when the museum was bombed during World War II. Approximately a Wien (NHMW) [5] (p. 3). Given that the type specimens were not present in either UZHM or NHMW, dozenthe type Eichho seriesff typeswas listed were as saved ”lost” by by K.E. Wood Schedl, and and Bright now [5] reside in their at the catalog Naturhistorisches of bark and ambrosia Museum Wienbeetles. (NHMW) However, [5] (p.a syntype 3). Given was that listed the type in specimensa checklist wereof scolytine not present and in platypodine either UZHM types or NHMW, in the theZoological type series Museum, was listed Museum as ”lost” and by In Woodstitute and of Zoology, Bright [5 ]Polish in their Academy catalog of of bark Science, and ambrosiaWarsaw, Poland beetles. However,(MIIZ) [7], abut syntype this checklist was listed was in not a checklistlisted in the of scolytinecatalog [5], and and platypodine was subsequently types in largely the Zoological ignored. Museum,This is one Museum of several and errors Institute pertaining of Zoology, to type Polish specimen Academy deposition of Science, listed Warsaw, in Wood Poland and Bright’s (MIIZ) [[5]7], butcatalog this (see checklist [8,9]). was not listed in the catalog [5], and was subsequently largely ignored. This is one of severalBeginning errors pertaining in the 1980 to type′s populations specimen depositionof what was listed presumed in Wood to and be Bright’sEuwallacea [5] fornicatus catalog (see became [8,9]). Euwallacea fornicatus establishedBeginning outside in the Asia 1980 0sin populations Central and of whatNorth was America, presumed Israel, to be and South Africa [4,10–13].became establishedStouthamer outside et al. [3] Asia sampled in Central E. fornicatus and North s.l. America, from multiple Israel, andlocalities South across Africa [Asia,4,10– 13as]. well Stouthamer as from E. fornicatus s.l. etintroduced al. [3] sampled populations around thefrom world. multiple Their molecular localities phylogeny across Asia, revealed as well that as from E. fornicatus introduced was E. fornicatus populationsa complex consisting around the of world. three Theirclades, molecular presumed phylogeny species, which revealed were that termed tea shotwas hole a complex borer consisting(TSHB), Kuroshio of three clades,shot hole presumed borer (KSHB), species, and which po werelyphagous termed shot tea shothole holeborer borer (PSHB). (TSHB), Gomez Kuroshio et al. shot[14] expanded hole borer the (KSHB), Stouthamer and polyphagous et al. [3] dataset shot an holed recognized borer (PSHB). the following Gomez etfour al. species: [14] expanded KSHB and the StouthamerPSHB, tea shot et al. hole [3] datasetborer a and(TSHBa), recognized and tea the shot following hole borer four b species: (TSHBb). KSHB These and clade PSHB, names tea shot are holefrequently borer aused (TSHBa), in the and literature, tea shot but hole they borer do b not (TSHBb). have official These cladetaxonomic names status are frequently and they usedare not in theofficially literature, recognized but they by dothe not Entomological have official Societ taxonomicy of America status andcommittee they are of notcommon officially names. recognized Gomez byet al. the [14] Entomological delineated species Society in of the America complex committee using morphometrics of common names. to associate Gomez molecular et al. [14] delineated data with speciestype specimens. in the complex With usingthe assumption morphometrics that tothe associate type series molecular of E. fornicatus data with was type lost specimens. [5], Gomez With etthe al. assumption[14] based their that thespecies type seriesconcept of E.on fornicatus the specimenswas lost sequenced [5], Gomez from et al. the [14 ]type based locality their species of Sri conceptLanka, onwhich the specimenscorresponded sequenced to TSHBa from clade. the type locality of Sri Lanka, which corresponded to TSHBa clade. InIn thisthis review, review, we we examined examined the rediscoveredthe rediscoveredXyleborus Xyleborus fornicatus fornicatussyntype syntype (Figure1 )(Figure and additional 1) and non-typeadditional specimens non-type ofspecim individualsens of inindividuals the Euwallacea in the fornicatus Euwallaceacomplex fornicatus so as complex to revise so diagnoses as to revise and providediagnoses new and and provide revised new distribution and revised information. distribution information. Figure 1. Syntype of Xyleborus fornicatus , female, from left to right: dorsal dorsal view, view, lateral lateral view, view, posterior oblique view ofof declivity,declivity, andand labellabel information.information. BodyBody lengthlength ofof 2.622.62 mm.mm. Insects 2019, 10, 261 3 of 11 2. Materials and Methods We follow the clade names for the Euwallacea fornicatus species complex proposed by Stouthamer et al. [3], and those subsequently followed by Gomez et al. [14]. The syntype of Xyleborus