SYSTEMATICS A New Species of Bark Beetle, … · SYSTEMATICS A New Species of Bark Beetle,...

SYSTEMATICS

A New Species of Bark Beetle Dendroctonus mesoamericanussp nov (Curculionidae Scolytinae) in Southern Mexico and

Central America

FRANCISCO ARMENDARIZ-TOLEDANO1 ALICIA NINO2 BRIAN T SULLIVAN3

LAWRENCE R KIRKENDALL4 AND GERARDO ZUNIGA15

Ann Entomol Soc Am 1ndash12 (2015) DOI 101093aesasav020

ABSTRACT The bark beetle Dendroctonus mesoamericanus sp nov is described from a population inParque Nacional Lagunas de Montebello La Trinitaria Chiapas Mexico This species belongs to theD frontalis complex which includes D adjunctus Blandford 1897 D approximatus Dietz 1890 D bre-vicomis LeConte 1876 D frontalis Zimmermann 1868 D mexicanus Hopkins 1905 and D vitei Wood1975 The new species can be distinguished from closely related species D frontalis D mexicanus andD vitei by four diagnostic morphological characters the presence of striations on the preepisternal areaof the prothorax (anterolateral pronotum) in both sexes seminal rod with an approximately linear poste-rior margin of the dorsal process (in lateral view) irregularly sized ornamentations on the distal edge ofsquamiform plates present on the eighth abdominal tergite in females and an inconspicuous pronotalcallus in females Furthermore the karyotype of D mesoamericanus sp nov (5AAthornXyp XX) differsfrom that of its sister species D frontalis (7AAthorn Xyp XX) Dendroctonus mesoamericanus sp nov is dis-tributed in Belize Guatemala El Salvador Honduras Nicaragua and in the Michoacan Oaxaca andChiapas states in Mexico In these countries this species has been collected at elevations above from 311to 2600 m on six pine species and it commonly occurs in syntopy with D frontalis

KEY WORDS bark beetle Dendroctonus mesoamericanus seminal rod spermatheca

Bark beetles of the genus Dendroctonus Erichson1836 are major natural disturbance agents of Holarcticconifer forests and include some of the most economi-cally significant forestry pests in the world They attackspecies in the genera Larix Mill Picea A Dietr andPseudotsuga Carriere but occur in greatest diversity inPinus L They complete their life cycle within the barkof their host trees where attacking adults pair ovipositwithin a single gallery excavated in the inner bark andboth the parent adults and developing larvae feed onphloem tissue (Wood 1982) They generally attack andkill trees that are prostrate or weakened by agedrought disease mechanical damage or other factorsHowever during very high populations associated withepidemics healthy trees can be colonized causing sig-nificant tree mortality and economic losses in the for-estry sector (Billings 2011)

In the first monograph of the genus DendroctonusHopkins (1909) recognized and integrated 24 valid

species arranged in divisions subdivisions sectionssubsections series and subseries according to morpho-logical characteristics Due to the high intra- and in-terspecific morphological variation of these species thegenus was subject to taxonomic revisions (Wood 19631982) Some species were synonymized (Wood 19631974 Lanier and Wood 1968) and new species or sub-species were described (Thomas and Bright 1970 Fur-niss 2001)

The genus currently includes 19 valid species (Wood1982) one of them with two subspecies (Furniss 2001Ruiz et al 2009) arranged into five to six groups orspecies complexes supported by morphological behav-ioral ecological chromosomal and molecular charac-teristics (Hopkins 1909 Wood 1963 1982 Lanier1981 Kelley and Farrell 1998 Zuniga et al 2002)

The Dendroctonus frontalis complex is composed ofsix species D adjunctus Blandford 1897 D approxi-matus Dietz 1890 D brevicomis LeConte 1876D frontalis Zimmermann 1868 D mexicanus Hopkins1905 and D vitei Wood 1974 (Wood 1982 Lanieret al 1988) Three species within the complexmdashD frontalis D mexicanus and D viteimdashare particu-larly problematic because they are very similarmorphologically (Wood 1974 1982 Vite et al 19741975 Lanier et al 1988) and their populations may co-exist in space and time in pine forests of Mexico andCentral America (Zuniga et al 1995 1999 Salinas-Moreno et al 2004 Moser et al 2005 Armendariz-Toledano et al 2014ab)

1 Instituto Politecnico Nacional Escuela Nacional de Ciencias Bio-logicas Prol de Carpio y Plan de Ayala Col Santo Tomas Del Mi-guel Hidalgo Mexico DF CP 11340

2 El Colegio de la Frontera Sur (ECOSUR) Carretera AntiguoAeropuerto km 25 Tapachula Chiapas Mexico CP 30700

3 USDA Forest Service Southern Research Station 2500 Shreve-port Hwy Pineville LA 71360

4 University of Bergen Department of Biology Postbox 7803 Ber-gen N-5006 Norway

5 Corresponding author e-mail capotezuhotmailcom

VC The Authors 2015 Published by Oxford University Press on behalf of Entomological Society of AmericaAll rights reserved For Permissions please email journalspermissionsoupcom

Annals of the Entomological Society of America Advance Access published April 3 2015

From early 2000 to 2002 gt25000 ha of mature pinestands in the Mountain Pine Ridge Forest Reserve ofBelize were devastated by an outbreak of what was ini-tially identified as D frontalis (Haack et al 2000 Ma-cıas-Samano et al 2001 Billings et al 2004) Duringstudies of this outbreak Midtgaard and Thunes (2002)collected a second apparently undescribed Dendrocto-nus species from infested trees that these authorscalled the ldquocaribbean pine beetlerdquo The caribbean pinebeetle species was distinguished from D frontalis bythe presence in both sexes of fine ridges on the pree-pisternal area of the prothorax (anterolateral pronotum)sensu Hopkins (1909 Fig 17) oriented roughly parallelto the anterior margin and by the width of the episto-mal process which appeared to be wider in caribbeanpine beetle than D frontalis (Thunes et al 20052011) These authors assigned the name ldquoDendroctonuswoodirdquo to identify caribbean pine beetle (Thunes et al2005 2011) however according to the InternationalCode of Zoological Nomenclature (ICZN 1999) thisbinomen is a nomen nudum because a formal descrip-tion of the species was not published

Because insufficient biological data supported sepa-ration of the D frontalis and caribbean pine beetle intotwo species Sullivan et al (2012) and Armendariz-Toledano et al (2014a) carried out studies to clarify thetaxonomic status of this bark beetle using crossing ex-periments as well as biochemical morphological karyo-logical and molecular data These studies consistentlysupported the separation of these species Thereforein this paper we describe Dendroctonus mesoamerica-nus sp nov which corresponds to ldquoD frontalis mor-photype Brdquo from Sullivan et al (2012) and Armendariz-Toledano et al (2014a) and to caribbean pine beetlefrom Midtgaard and Thunes (2002)

Materials and Methods

Specimens A total of 289 adult Dendroctonus spnov were examined from 40 collection sites (Table 1)Specimens reported by Armendariz Toledano et al(2014a) corresponding to Dendroctonus sp nov (ieD frontalis morphotype B) were included in the analy-ses Additionally adult specimens of Dendroctonus spnov were collected by the authors from naturallyinfested Pinus sp or graciously loaned by the followinginstitutions Comision Forestal del Estado de Michoa-can Morelia Michoacan El Colegio de la Frontera SurTapachula Chiapas (ECOSUR) Laboratorio de Varia-cion Biologica y Evolucion Escuela Nacional de Cien-cias Biologicas Instituto Politecnico Nacional MexicoColeccion Cientıfica de Entomologıa Forestal Divisionde Ciencias Forestales Universidad Autonoma Cha-pingo (UACH) Estado de Mexico United States ForestService Southern Research Station Pineville LA

Morphology Sex of specimens was determined bya combination of dimorphic characters the size andabundance of large tubercles on the fronts the stridula-tory apparatus in males and genitalia (Lyon 1958Mendoza-Correa and Zuniga 1991) Species identifica-tion of specimens within the D frontalis complex wasperformed using the key of Lanier et al (1988)

Specimens of Dendroctonus sp nov were distinguishedby means of diagnostic characters reported by Armen-dariz-Toledano et al (2014a) for D frontalis morpho-type B ie the presence of striations on preepisternalarea (PEA) the irregular size of ornamentations of theplates of the eighth tergite of females (AP) and theshape of seminal rod (SSR) (Armendariz et al 2014aFig 1)

External morphology of adults was exhaustivelyexamined and photographed with a NIKON SMZ800dissecting microscope and a Nikon DXM1200F digitalcamera Body measurements were taken with this samemicroscope with a micrometer eye piece (at 20ndash63)For some photos of the holotype and paratype 2(Fig 1) images were recorded at successive focaldepths and automatically combined into a single sharpimage (ie focus stacking) using CombineZM software(Hadley 2008) Morphological details of some speci-mens were photographed with an environmental scan-ning electron microscope ESEM (Evo 40 VP ZeissJena Germany)

The elytra and genitalia from 100 specimens of bothsexes were removed These structures were cleared byincubating them at 50C in 10 KOH for 120ndash180 minfor elytra and for 10ndash15 min for male and female geni-talia After incubation structures were immersed in20 acetic acid solution to neutralize the KOH andsubsequently rinsed with 100 ethanol For both malesand females the elytra were cut into two parts (elytraldisc and declivity) and then mounted on the same slideThe eighth abdominal tergite and spermatheca fromindividual females were mounted in the same slideand the seminal rod and genital capsule of individualmales were separated and mounted in the same slide

All structures were mounted semipermanently inglycerol and observed at 100ndash400 with phase contrastmicroscopy Elements of elytral sculpture (puncturesgranules crenulations) were measured and quantifiedwith a phase contrast microscope with an ocularmicrometer (400) Photographs of male and femalegenitalia were taken with a Nikon DXM1200F digitalcamera mounted in a phase contrast microscopeMorphological details of genitalia were photographedwith an ESEM (Evo 40 VP Zeiss)

From the entire series (nfrac14 289) 98 specimens from26 geographic localities were selected to constitute thetype series (Table 1) In the type series six specimensused by Armendariz-Toledano et al (2014a) in the kar-yological analysis were included The genitalia of theholotype allotype and 18 paratypes were extractedphotographed and preserved in a glycerol-filled micro-vial affixed to the same pin as the respective specimenHolotype was indicated with H1 label allotype wasindicated with A1 label and individual paratypes weredesignated from P1 to P96

Dendroctonus mesoamericanusArmendariz-Toledano amp Sullivan new species

(Figs 1ndash3)

Type Material HOLOTYPE male MexicoChiapas La Trinitaria Parque Nacional Lagunas de

2 ANNALS OF THE ENTOMOLOGICAL SOCIETY OF AMERICA

Montebello (CTP) 16 0804200 N 91 4302900 W1800 m 20-October-2011 on Pinus maximinoi H EMoore A Nino (A N) and F Armendariz-Toledano(F A T) coll in the Coleccion Nacional de Insectosdel Instituto de Biologıa Universidad Nacional Auton-oma de Mexico (CNIN)

ALLOTYPE female from type locality same coordi-nates elevation date host collectors in CNIN

PARATYPES 96 40 (P1-P40) type locality (CTP)(4 3) 16 0804200 N 91 4302900 W 1587 m 9-IX-2009 (15 19) 16 0604500 N 91 4305600 W 1494 m20-X-2011 on P maximinoi A N and F A T collnine (P41-P49) (5 4) Guatemala ChimaltenangoMisco Club Campestre de Alta Montana (GMC) 14

470900 N 90 5502200 W 2410 m 26-VI-2011 on P max-iminoi A N and F A T coll two (P50-P51) ()Mexico Chiapas Teopisca Area Forestal de Teopisca

(CTT) 16 3402500 N 92 2703000 W 2321 m 5-XI-2003 on ex Pinus trapped J Macıas coll two(P52-P53) () Mexico Chiapas Comitan PredioSanta Cruz (CCC) 16 1204700 N 92 1400000 W1801 m 5-XII-2003 on P oocarpa Schiede ex Schlech-tendal C A Gallegos coll four (P54-P57) (2 2)Mexico Chiapas Jitotol de Zaragoza Ejido las Maravil-las (CJM) 17 0504800 N 92 5302800 W 1677 m 5-X-2005 trapped J Macıas coll two (P58-P59) ()Mexico Oaxaca Santiago Yosondua Santa CatarinaCuanana Paraje la Guacamaya (OSY) 16 4804900 N97 2405500 W 2142 m 2005 on P oocarpa J Cruzcoll four (P60-P63) (2 2) Mexico MichoacanChinicuila El Volantin (MCV) 18 3802400 N 103

2105200 W 1180 m 14-XI-2005 Comision Forestal delEstado de Michoacan one (P64) () Mexico Michoa-can Villa Madero Balcones (MVB) 19 150000 N 101

Table 1 Key country locality coordinates and host of the populations of Dendroctonus mesoamericanus sp nov

C Key Localitie Latitude N Longitude W Altitude(m)

Host

Mexico MTO Michoacan Tancitaro Ojo de Agua de los Sauces 19 1400000 102 1700000 1803 MVB Michoacan Villa Madero Balconesa 19 150000 101 104800 1813 Pinus pringleiMCV Michoacan Chinicuila El Volantina 18 3802400 103 2105200 1180 OSM Oaxaca Santa Marıa Albarradas San Pablo Villa de Mitla 16 5903300 96 1101500 2106 P pringleiOSY Oaxaca Santiago Yosondua Santa Catarina Cuanana

Paraje la Guacamayaa16 480490 0 97 240550 0 2142 P oocarpa

OHT Oaxaca Heroica Cd de Tlaxiaco Paraje Cerro Negro 17 2002300 97 4503600 2454 P ocarpa ampP pringleii

OPZ Oaxaca San Pedro Ayutla Zacatepec Distrito Mixe 17 070120 0 96 440560 0 1770 P pringleiCJM Chiapas Jitotol de Zaragoza Ejido Las Maravillasa 17 050480 0 92 530280 0 1677 CTT Chiapas Teopisca Area Forestal de Teopiscaa 16 3402500 92 2703000 2321 CTC Chiapas Teopisca Bienes Comunales de Teopiscaa 16 3003000 92 2901500 1809 CBS Chiapas Reserva de la Biosfera la Sepultura 16 2500000 93 2000000 598 CCC Chiapas Comitan Predio Santa Cruza 16 1204700 92 1400000 1347 P oocarpaCTP Chiapas La Trinitaria PNLMb 16 080420 0 91 430290 0 1557 P oocarpa amp

PmaximinoiCTPI Chiapas La Trinitaria PNLMa 16 060450 0 91 430560 0 1497 Pmaximinoi

Belize BMP1 Cayo Mountain pine ridgea 17 10 88 180 PcaribaeaBMP2 Cayo Mountain pine ridgea 16 590 88 490 901 P oocarpaBMP Cayo Mountain pine ridge 16 590 88 460 928 BMP3 Cayo Mountain pine ridgea 17 000 88 510 857 P oocarpa

Guatemala GHC Huehuetenango Canselaj 15 160100 0 91 250530 0 2065 P oocarpaGHM Huehuetenango Malacatacinto Cacum 15 150090 0 91 420030 0 2067 P montezumaeGQQ El Quiche Santa Cruz del Quiche Chinique de las Floresa 15 10 91 80 2024 P montezumaeGQS Quetzaltenango San Carlos Sija Estancia de la Virgena 14 59080 0 91 300290 0 2612 P oocarpaGST Solola Colonia Maria Tecun 14 4805000 91 120200 0 2409 P oocarpaGMC Mixco Club Campestre Alta montanaa 14 47090 0 90 550220 0 2410 P maximinoiGCT Chimaltenango Tecpan Carretera Interamericana 13 km 14 4505000 90 5901400 2288 P maximinoiGSS Solola Finca Socorro 14 4501000 91 0802000 1771 PpseudostrobusGSC Solola Finca Chuchiya 14 4400000 91 0705000 2022 PpseudostrobusGM Guatemala Mixcoa 14 38000 0 9036000 0 1660 P maximinoii amp

P oocarpaGMS Guatemala Mixco San Cristobala 14 3601400 90 3504000 1582 P pseudostrobus

Honduras HYY Yoro Yorito Luquiguea 14 590000 0 87 360000 0 311 HGA Guaimaca 14 320080 0 86 490000 0 862 P oocarpaHCS Comayagua Sihuatepequea 14 310490 0 87 320130 0 1717 PoocarpaHT Distrito Central Tegucigalpaa 14 80520 0 87 100210 0 1168 P oocarpaHMT Francisco Morazan Tatumblaa 14 00580 0 87 60160 0 1374 P oocarpa

ES SSC San Salvador El Carmena 13 45000 0 89 12000 0 561 P caribaeaNicaragua NJP Jalapa 13 55010 0 86 070380 0 697 P caribaea

NSF Nueva Segovia San Fernandoa 13 400380 0 86 180550 0 734 P oocarpaNMZ Madriz Somotoa 13 250140 0 86 340490 0 1366 PoocarpaNEE Esteli Estelia 13 120000 0 86 180000 0 1290 P oocarpaNPT El Picacho Cerro Tomabu 13 020000 0 86 180000 0 1238 P oocarpa

C country ES El Salvador PNLM Parque Nacional Lagunas de Montebelloa Localities included in the type seriesb Type locality

2015 ARMENDARIZ-TOLEDANO ET AL D mesoamericanus sp nov DESCRIPTION 3

104800 W 1813 m 14-XI-2009 on P pringlei GR Shawex Sargent Comision Forestal del Estado de Michoa-can coll two (P65-P66) () Mexico Chiapas Teo-pisca Bienes comunales de Teopisca (CTC) 16

3003000 N 92 2901500 W 1809 m 2003 on ex Pinustrapped J Macıas coll six Belize Cayo MountainPine Ridge (BMP1-3) (P67-P68) (1 1) 17 10 N88 180 W 9-VII-2013 on P caribaea Morelet B TSullivan coll (P69-P70) (1 1) 16 590 N 88 490

W 901 m 14-VII-2009 on P oocarpa S R Clarkecoll (P71-P72) (1 1) 17 000 N 88 510 W 857 m16-VII-2009 on P oocarpa S R Clarke coll two(P73-P74) (1 1) Guatemala El Quiche SantaCruz del Quiche Chinique de las Flores (GQQ) 15 10

N 91 80 W 2224 m 4-VII-2013 on P montezumaeLambert P Ortiz coll two (P74-P75) (1 1) Gua-temala Quetzaltenango San Carlos Sija Estancia de la

Virgen (GQS) 14 590800 N 91 3002900 W 2612 m7-III-2007 P oocarpa P Ortiz coll two (P77-P78)(1 1) Guatemala Mixco (GM) 14 380 N 90 360

W 1660 m P maximinoi and P oocarpa 25-I-2013 POrtiz coll two (P79-P80) (1 1) GuatemalaMixco San Cristobal (GMS) 14 3601400 N 90 3504000

W 1582 m 23-IV-2013 on P pseudostrobus Lindley POrtiz coll two (P81-P82) (1 1) Honduras YoroYorito Luquigue (HYY) 14 5900000 N 87 3600000 W311 m 31-V-2007 R F Billings coll two (P83-P84)(1 1) Honduras Comayagua Sihuatepeque(HCS) 14 3104900 N 87 3201300 W 1717 m 9-XI-2011 on P oocarpa R F Billings coll two (P85-P86)(1 1) Honduras Distrito Central Tegucigalpa(HT) 14 805200 N 87 1002100 W 1168 m 10-VIII-2011 on P oocarpa R F Billings coll two (P87-P88)(1 1) Honduras Francisco Morazan Tatumbla(HMT) 14 005800 N 87 601600 W 1374 m 8-VIII-2011 on P oocarpa R F Billings coll two (P89-P90)(1 1) El Salvador San Salvador El Carmen(SCC) 13 450 N 89 120 W 561 m 22-XI-2003 onP caribaea R F Billings coll two (P91-P92) (11) Nicaragua Nueva Segovia San Fernando (NSF)13 4003800 N 86 1805500 W 734 m 12-III-2010 onP oocarpa R F Billings coll two (P93-P94) (11) Nicaragua Madriz Somoto (NMZ) 13 2501400

N 86 3404900 W 1366 m 13-III-2010 on P oocarpaR F Billings coll two (P95-P96) (1 1) NicaraguaEsteli Esteli (NEE) 13 120 N 86 180 W 4-XII-13 onP oocarpa R Billings coll

Thirty-five paratypes were deposited in CNIN Mex-ico (P1-P7 P41- P44 P50 P52 P54-P55 P58 P60-P61 P64 P66 P68 P70 P72 P74 P76 P78 P80 P82P84 P86 P88 P90 P92 P94 P96) 22 paratypes in theMuseo de Historia Natural de la Ciudad y CulturaAmbiental Mexico (P18-P28 P45 P46 P49 P56 P62P67 P73 P79 P85 P91 P95) 20 paratypes are in theSmithsonian National Museum of Natural History(P13-P17 P29-P34 P47 P51 P57 P63 P69 P75 P81P87 P93) and 19 in the American Museum of NaturalHistory (P8-P12 P35-P40 P48 P53 P59 P65 P71P77 P83 P89)

Diagnosis Dendroctonus mesoamericanus sp novis distinguished from closely related species D frontalisZimmermann D mexicanus Hopkins and D viteiWood by the presence of striations on the preepisternalarea in both sexes running parallel to the anterior mar-gin of the prothorax (Figs 1c d and 2j k m) which areabsent in the other three species It is also distin-guished by the seminal rod which has an approxi-mately triangular-shaped laterally compressed dorsalprocess with a nearly straight posterior margin in lateralview (ie arching inward or outward by less thanapproximately 15 of the length of the posterior mar-gin Figs 2s and 3a b) In D frontalis the seminalrod in lateral view has a hemicircular or lobed dorsalprocess with a distinctly convex posterior margin inD mexicanus and D vitei the dorsal process has astrongly concave posterior margin in lateral view andis not compressed laterally (Vite et al 1975 Figs 4ndash6Armendariz-Toledano et al 2014a Fig 5a)In D mesoamericanus females the squamiform plates

Fig 1 Dendroctonus mesoamericanus sp nov maleadult Holotype dorsal view (a) lateral view (b)preepisternal area (c) paratype one preepisternal area (d)The arrows indicate the striations on preepisternal area


Fig 2 Dendroctonus mesoamericanus sp nov adult general anatomy (a) male head (b) male epicranial surface (c)male front surface (d) male epistomal process surface (e) female head (f) female epicranial surface (g) female front surface(h) female epistomal process surface (i) anterior view of preepisternal area (j) lateral surface of pronotum (k) dorsal surfaceof pronotum (l) preepisternal area (m) lateral posterior surface of pronotum (n) episternal surface (o) elytral declivity (p)three first interspaces on the distal area of elytral disc (q r) lateral-dorsal view of seminal rod (s) plates on eight tergite infemales (t) ap acute projections cr crenulations dp dorsal projection dpu distance between punctures ft frontal tuberclesgr granules in 1 interspace one in 2 interspace two in 3 interspace three pu punctures st striations on preepisternal areast 1 striae one st 2 striae two st 3 striae three vp ventral process The letters in (a) indicate the figures in which themorphological structure is showed


of the eighth tergite possess ornamentations along theirdistal edge of highly variable size (Figs 2t and 3c d)whereas in D frontalis these ornamentations are uni-formly short In addition females of D mexicanus andD frontalis can be readily distinguished from those ofD mesoamericanus sp nov by the inconspicuous pro-notal callus in the latter (Fig 1c f in Sullivan et al2012) Further it can be distinguished from its sisterspecies D frontalis by male meiotic formula 5AAthornXyp

(Fig 6 in Armendariz-Toledano et al 2014a)Holotype Male (Fig 1a b c) Total length 41 mm

(27ndash49 mm Nfrac14 50) 27 times longer than widehead pronotum and elytra medium brown to blackwith head darker than pronotum and pronotum darkerthan elytra

Head Length of head 07 mm surface of epicra-nium and vertex covered with small punctures (Fig 2bc) vertex and frons truncated convex or completelyconvex (in lateral view) Frons armed by a pair of trans-verse elevations surrounded by eight prominenttubercles (paratypes 4ndash8 tubercles Nfrac14 50) and gran-ules just below upper level of eyes (Fig 2b) length ofhighest frontal tubercles 004 mm (0015ndash006 mmNfrac14 50) frontal tubercles separated by a mediangroove extending from just above epistomal process toupper level of eyes (Fig 2b) surface of median grooveis principally shining and smooth (some specimens withsmall granules between frontal transverse elevations)surface between eyes with punctures and granules dis-tributed around median grove (Fig 2b d) puncturesare less impressed in upper and lateral areas of episto-mal process (Fig 2b e) area immediately above

epistomal process flat (some specimens flat or slightlyconcave) shining finely granulate and with sparsesmall punctures (some specimens with or withoutpunctures Fig 2b e) Width of epistomal process058 mm (033ndash075 mm Nfrac14 50) 23 of distancebetween eyes (09 mm 06ndash10 mm Nfrac14 50) arms ofepistomal process thick elevated and oblique approxi-mately 26 from horizontal (Fig 2b e) transversemidsection of the epistomal process absence or weakseparating the prominent arms by a space equal orlarger than the length of the one arm surface smoothand shining with small punctures underside with adense brush of conspicuous yellowish setae pubes-cence of head yellowish of three different sizes sparsedistributed around frons and less abundant on areaimmediately above epistomal process (Figs 1b and 2bd) shortest setae distributed on vertex those ofmedium size on vertex and frons and largest setae sur-rounding epistomal process

Pronotum Length 092 mm (064ndash13 mm Nfrac14 50)width 15 mm (08ndash17 mm Nfrac14 50) widest on poste-rior third anterior region slightly constricted dorsalsurface with rather coarse moderately deep closepunctures and interspaces smooth and shiny (Figs 1aand 2k l n) Punctures largest on anterior dorsal areawhereas those on preepisternal and episternal areas(lateral surfaces on anterolateral pronotum) shallowerless abundant and reduced in diameter (Fig 2o) sur-face of preepisternal area with a zone of striations run-ning parallel to anterior margin of pronotum (Figs 1cd and 2j k m) pubescence yellowish longer and moresparse on dorsal posterior area (Figs 1b and 2l)

Fig 3 Dendroctonus mesoamericanus sp nov genital organs Male seminal rod (Sr) in dorsal and lateral view holotype(a) paratype one (b) female spermatheca (Sp) and eight tergite (et) allotype (c) paratype six (d) ap acute projections cocornu dp dorsal process no nodulus rsp ridges of squamiform plates sv seminal valve vp ventral process


pubescence of preepisternal and episternal areasshorter and less abundant than dorsal area (Fig 2k mo)

Elytra Length 24 mm (16ndash27 mm Nfrac14 50) 15times longer than wide 26 times longer than prono-tum sides straight on basal two-thirds rather broadlyrounded behind (Fig 1a) declivity convex with striaeweakly impressed (Fig 2p) punctures of striations witha diameter ranging from 16 to 28mm (Fig 2p r) sur-face between punctures smooth with a distancebetween punctures of 10 to 31mm (Nfrac14 15) diameterof punctures roughly averaging size of interspaces(Fig 2p) declivital interstria (measured at mid-interstria) 2 narrower (X frac14 72mm Nfrac14 15) than 1(X frac14 93mm Nfrac14 15) and 3 (X frac14 100mm Nfrac14 15) decli-vital interstria 2 not constricted apically (mildly con-stricted or not Fig 2p) surface of interstriae withslightly marked rugae punctures and crenulationsPunctures on declivital interstriae less well definedwith some slightly larger than those of striae averagediameter of strial punctures 11 times diameter ofpunctures on interstriae (Fig 2p r) crenulations ininterstriae of variable size more defined and abundanttoward anterior part (Fig 2q) average width of crenu-lations on proximal area of elytral declivity 05 timeswidth of interspaces Setae in interstriae 1ndash3 of decliv-ity originating from center of punctures more than onesize class setae shortest about one-third length of lon-gest and longer ones show a gradual progression ofsizes color yellow to amber in mounted specimens(Fig 1b)

Genitalia Composed of four sclerotized structurestegmen spicule penis and accessory apparatus (semi-nal rod and anchor) General anatomy of tegmen spi-cule and penis similar to Dendroctonus ponderosaeHopkins (frac14 D monticolae Hopkins) (Cerezke 1964)

Anchor consisting of a tubular bag supported by twoanchor branches and located within penis Seminal rodconsisting of a sclerotized structure connected to ejacu-latory duct and situated within anchor Proximal regionof seminal rod consisting of a seminal valve with distalportion divided into dorsal and ventral processes(Figs 2s and 3ab) Ventral process consisting of anacute spine-shaped projection forming an approxi-mately 90 angle with posterior margin of dorsal proc-ess Dorsal process consisting of a laterally compressedbroadly triangularly shaped projection with ventralprocess forming a continuation of ventral side of trian-gle (Figs 2s and 3ab) Ventral process extends furtherfrom seminal valve (ie more distally) than dorsal proc-ess dorsal process somewhat narrower than dorsalprocess in ventral view and posterior margin of ventralprocess varies only slightly from straight in lateral view(Fig 3a b)

Allotype Female Total length 42 mm (22ndash46 mmNfrac14 48) 28 times longer than wide body color similarto male

Head Length 06 mm surface of epicranium andvertex covered with small punctures (Fig 2f g) lessabundant and smaller than in male shape of vertex andfrons in lateral view similar to male Frons with slighttransverse elevations covered with small granules

tubercles much less prominent than those on male(Fig 2f) transverse elevations separated by a mediangroove extending from just above epistomal process toupper level of eyes (Fig 2f) median groove weak vary-ing from scarcely perceptible to distinct only occasion-ally stretching all the way from vertex to epistomamainly visible between elevations surface of mediangroove shining and largely impunctate punctures andgranules distributed around of median grove less abun-dant and smaller than in male (Fig 2f h) distributionof punctures and granules between eyes is similar tomales (Fig 2d h) area immediately above epistomalprocess flat (some specimens concave) shining lessgranulate and with more sparse and smaller puncturesthan in males (some specimens without punctures)(Fig 2f i) epistomal process (width 052 mm033ndash069 mm Nfrac14 48) narrower than males 34 thedistance between eyes (07 mm 048ndash097 mmNfrac14 48) arms of epistomal process elevated obliqueand about 21 from horizontal (Fig 2f i) surfacesmooth and shining with punctures less abundant andsmaller than in male arms of the epistomal processcontinuous or nearly so (not clearly separated as inmale) underside with a less dense brush of setae thanin male pubescence of head slightly more abundant(Fig 2f i g)

Pronotum Length 10 mm (07ndash12 mm Nfrac14 48)width 15 mm (09ndash18 mm Nfrac14 48) widest on poste-rior third anterior region slightly constricted pronotalconstriction possessing a transverse callus that is visiblebut very inconspicuous both laterally and dorsally ves-titure and sculpture of pronotum similar to males

Elytra Length 25 mm (10ndash29 mm Nfrac14 48) 16times longer than wide 23 times longer than prono-tum sides parallel on basal two-thirds rather broadlyrounded behind declivity convex with striae weaklyimpressed vestiture and sculpture of elytral declivitysimilar to males

Eighth Tergite Structure characterized by an ovaldistal edge and two oval proximal lateral edges converg-ing on an anterior median line Distal surface of tergiteis covered by abundant setae of two different sizesLength of large setae 012 times width of tergite andtwo times longer and wider than small setae Eighthtergite displays 18 parallel lines of squamiform plates ofsclerotized cuticle (13ndash22 Nfrac14 20 Fig 3c d) Linesbetter defined in anterior area and fade in posteriorarea Squamiform plates possess acute distal projec-tions of variable size (Figs 2t and 3c d)

Spermatheca Reniform and divided into two partsnodulus and cornu Nodulus transversely covered by 18striae (10ndash23) becoming confused in proximal region ofnodulus Cornu is longer than width of middle constric-tion displaying an oval shape in lateral view (Fig 3c d)

Variation Despite wide morphological variation ofD mesoamericanus this species has characters in addi-tion to those in the diagnosis which produce good (butless reliable) discrimination from sibling species Theseinclude body measurements sculpture of elytral decliv-ity and female terminalia

Body proportions can be used for discriminationbetween D mesoamericanus sp nov and D frontalis


when long series are available Numerous body meas-urements are on average greater in D mesoamericanusepistomal process width (EPW 051 6 SEX 001 mm)distance between eyes (DE 083 6 01 mm Nfrac14 289)eye width (WE 024 6 0003 mm Nfrac14 289) width ofpronotum at widest point (PW 139 6 003 mmNfrac14 289) pronotum length (PL 097 6 0014 mm Nfrac14289) and elytral length (EL 226 6 007 mm Nfrac14 289)in contrast these measurements in D frontalis areEPW 038 6 005 mm Nfrac14 289 DEmdash067 60006 mm Nfrac14 289 WEmdash026 0001 mm Nfrac14 289PWmdash118 6 001 mm Nfrac14 289 PLmdash081 6 001 mmNfrac14 289 ELmdash206 001 mm Nfrac14 289 Of these epis-tomal process and eye width have the lowest overlap inSEX between species and consequently size of thesefeatures are better characters than other external meas-urements for separating these two species by size(EPW and WE are 30 and 20 respectively larger inD mesoamericanus sp nov) Frontal tubercles are rel-atively more prominent in male D frontalis than inmale D mesoamericanus sp nov

Strial punctures of the elytral declivity of D mesoa-mericanus sp nov are larger (diameter Dfrac14 172612mm Nfrac14 40) and more sparse (distance betweenpunctures DBPfrac14 264 6 11mm Nfrac14 40) than inD frontalis (Dfrac14 1506 05mm Nfrac14 40 DBPfrac14 187606mm Nfrac14 40) In some specimens the number ofgranules (G) and punctures (P) on the interstriae 3 ofthe elytral declivity is apparently greater in D frontalis(Gfrac14 2676 42 Nfrac14 100 Pfrac14 1476 17 Nfrac14 100) thanD mesoamerianus sp nov (Gfrac14 2136 20 Nfrac14 100Pfrac14 12 6 20 Nfrac14 100) Because the variation in thenumber and size of these structures is wide in bothspecies these characters are not recommended foridentification

The eighth tergite in D mesoamericanus sp nov hason average a greater number of rows of squamiformplates than D frontalis (18 vs 16 respectively) Thespermatheca of Dendroctonus mesoamericanus sp novhas the edge of the cornu ovate striations coveringmore than half of the spermatheca body and conglom-erate striations in the proximal region of nodulus Thespematheca of D frontalis females tend to has theedge of the cornu round striations covering one fourthof the spermatheca body and a nodulus with conglom-erate striations in the proximal region

Although the seminal rod shape varies intraspecifi-cally associated to the evagination of the posterior mar-gin of the dorsal process and in the relative size ofventral process (Fig 4) this structure can be used reli-ably for discriminating D mesoamericanus sp nov andD frontalis In D mesoamericanus sp nov the poste-rior margin of the dorsal process is either straight(Fig 3a b) or slightly convex or concave (eg Fig 4populations MTO OHG GMS NPT) Additionally theventral process is invariably longer than the dorsal one(see Fig 1a in Armendariz-Toledano et al 2014b) inall specimens In contrast D frontalis has a dorsal proc-ess that is strongly convex and is equal to or greaterthan ventral process in length (Fig 5 in Armendariz-Toledano et al 2014a) The overall shape of the dorsalprocess in D frontalis is semicircular or bulbous

whereas in D mesoamericanus sp nov it is broadlytriangular

Distribution The type locality is the ParqueNacional Lagunas de Montebello La Trinitaria Chia-pas Mexico which is part of the Sierra de ChiapasThe climate of this region is temperate and humid withrain all seasons and the dominant vegetation is mixedpine-oak and sweetgum (Comision Nacional de AreasNaturales Protegidas [CONANP] 2007) Six pine spe-cies are present in this locality Pinus caribeae P leio-phylla P maximinoi P montezumae P oocarpaP pseudostrobus and P tenuifolia (frac14 P maximinoi)(Gonzalez-Espinosa et al 1997) however Dendrocto-nus mesoamericanus has been recorded in this areaonly in P maximinoi and P oocarpa

Identifications based on seminal rod shape the pres-ence of striations on the preepisternal area and identi-fication based on DNA sequencing indicate thatD mesoamericanus occurs within Nicaragua Guate-mala El Salvador Honduras Belize and states ofChiapas Oaxaca and Michoacan in Mexico Withinthis geographical area the distribution of this speciesappears to be entirely sympatric with D frontalis

Biology Dendroctonus mesoamericanus has beenobserved at elevations above 311 m on P caribaeaP devoniana P hartwegii (frac14 P rudis) P oocarpaP maximinoi P montezumae P pringlei and P pseu-dostrobus Generally this species has been encounteredinfesting hosts simultaneously with D frontalis (ieoccurring in syntopy) In this situation D mesoameri-canus colonizes primarily the bottom 1ndash3 meters of thestem whereas D frontalis is concentrated in the mid-to upper-bole However there is extensive spatialoverlap by these species in the bark and D mesoameri-canus brood have been reared in at least small num-bers from the entire length of the bole as well as thecrown of co-infested trees (Moreno 2008) Midtgaardand Thunes (2002) likewise observed that this speciescan colonize the entire bole All brood stages appear tobe present at any time of year (as with D frontalis)

The aggressiveness of D mesoamericanus and itsregional importance as a killer of pines has not beenadequately established Midtgaard and Thunes (2002)reported this species to be the major mortality agentduring the catastrophic Dendroctonus outbreak in Bel-ize in 2000ndash2002 However we have only occasionallyobserved D mesoamericanus initiating attacks on appa-rently vigorous trees in the absence of other barkbeetle species Furthermore its habit of colonizing pre-dominantly the basal portions of host trees coinfestedwith another species is a behavior typical for secondary(ie relatively less aggressive) bark beetles (Wood1982) In trees colonized by both D frontalis andD mesoamericanus the authors have always observedthe former species greatly outnumbering the latterD mesoamericanus has not been observed infestingdowned trees

Pitch tubes of D mesoamericanus are conspicuouslylarger than those of D frontalis Parent galleries areS-shaped frequently intercross and cannot be readilydistinguished from those of D frontalis (as displayed inWood 1982 Fig 60G) As with D frontalis larvae


Fig

4

Geo

grap

hic

ald

istr

ibu

tion

ofD

endr

octo

nus

mes

oam

eric

anus

sp

nov

T

he

Key

sof

loca

litie

sar

esh

owed

inT

able

1B

el

Bel

ize

ES

E

lS

alva

dor

G

ua

Gu

atem

ala

Hon

H

ond

ura

sM

exM

exic

oN

icN

icar

agu

a


mine within the inner bark roughly perpendicularly tothe parent gallery and form a progressively broadenedfrass-filled feeding chamber typically one to a few cen-timeters distance from the parent gallery Pupationoccurs in the outer bark Peak emergence of D mesoa-mericanus from logs derived from coinfested treesoccurs three to four weeks later than D frontalisalthough it is not known whether this is due to laterattack by D mesoamericanus a longer developmenttime for this species or both factors

D mesoamericanus differs from other species in theD frontalis complex in its pheromone and cuticularhydrocarbon compositions A pheromone produced byD mesoamericanus females strongly attracted conspe-cific males in a walking olfactometer while being largelyunattractive to D frontalis males (unpublished data)Additionally synthetic trap lures that are highlyattractive to flying D frontalis catch very few D mesoa-mericanus (Sullivan et al 2012) Female D mesoameri-canus produce the combination of frontalin endo-brevicomin and ipsdienol (Sullivan et al 2012)whereas this blend does not occurs in females of otherspecies in the D frontalis group that have so far beeninvestigated (ie D adjunctus D brevicomis D fron-talis and D vitei Renwick et al 1975 Hugheset al 1976 Byers et al 1984) Additionally the compo-sition of the cuticular hydrocarbon blend of D mesoa-mericanus differs significantly from that of D frontalis(Sullivan 2012) In particular the ratio of 5 - to 9-methylheptacosane is a biochemical character that pro-vides gt999 reliable separation of D mesoamericanusand D frontalis specimens collected in Belize andChiapas Mexico

Etymology The specific epithet mesoamericanus isa substantive in genitive that refers to the geographicregion called Mesoamerica It is the region in theAmerican continent that include the meridional regionfrom Mexico and the territories from Guatemala ElSalvador y Belize the west of Honduras Nicaraguaand a part of Costa Rica This region was defineddelimited and characterized by Kirchhoff (1943) Thisname was chosen for this species because all collectionsare in the central and southern region of MesoamericaMichoacan Oaxaca Chiapas Guatemala Belize ElSalvador Honduras and Nicaragua The commonname to propose for this species is Mesoamerican pinebeetle (acronym MAPB)

Systematics and Affinities The new speciesbelongs to the Dendroctonus frontalis complex thatincludes D adjunctus D approximatus D brevicomisD frontalis D mexicanus and D vitei (Lanier et al1988) The complex (Wood 1968 1982) is character-ized by a deep narrow median groove on the frons ofsexes the frons at the level of the upper eyes protuber-ant and armed by tubercles (most noticeable in males)with the exception of D adjunctus and a transverse ele-vated callus inconspicuous occurring laterally and dor-sally on the anterior region of pronotum in femalesThese species except D adjunctus were included inDendroctonus species group I according to morpholog-ical ecological and biological characteristics (Wood1963 1982) or group VI based on the small

chromosomal number that present these species withinDendroctonus genus with karyotypes of six seven oreight pairs of chromosomes (Lanier 1981 Zuniga et al2002)

Phylogenetic analysis of the Dendroctonus frontaliscomplex based on 786 base pairs of cytochrome oxidaseI of mtDNA indicated that the species group ofD mesoamericanus D frontalis D mexicanus andD vitei is monophyletic and further that D mesoamer-icanus and D frontalis are monophyletic within thisspecies group (Fig 7 in Armendariz-Toledano et al2014a)

D mesoamericanus and D frontalis are sibling spe-cies (sensu lato) and share morphological similaritiesin male genitalia (Sullivan et al 2012 Armendariz-Toledano et al 2014a) The seminal rod of these twospecies has a dorsal process that is laterally compressedand broadened in the sagittal plane (shape broadlytriangular hemicircular or lobed) and an acute spine-shaped ventral process (Fig 5a in Armendariz-Toledano et al 2014a) In contrast D mexicanus andD vitei have a dorsal process that is broadened perpen-dicularly to the sagittal plane and has an invaginatedposterior margin in lateral view (Vite 1975 Lanier et al1988 Armendariz-Toledano et al 2014 ab) Geometricmorphometric analyses of the seminal rods revealedquantitative differences among these four species(Armandariz-Toledano et al 2014ab)

Acknowledgments

We are grateful to Juan Jose Morrone David Cibrian andMiguel Angel Barrios for their comments on an earlier ver-sion of the manuscript We thank Juan Cruz (ComisionNacional Forestal Mexico) Ivon Lopez (Comision ForestalMichoacana) Roberto Castellanos (Parque Nacional deLagunas de Montebello Mexico) Jorge Macias Samano (Syn-ergy semiochemical Cooporation) Ronald Billings (TexasForest Service) Steve Clarke (USDA Forest Service) andFred Mitgaard and Karl H Thunes (Norwegian ForestResearch Institute) for their assistance in the field and labora-tory and for providing some specimens This study was parti-ally funded both by the Secretarıa de Investigacion yPosgrado IPN (20144216) and USDA Forest Service cooper-ative agreements FAT was a member of Programa Institu-cional de Formacion de Investigadores of InstitutoPolitecnico Nacional (PIFI-IPN) and of Consejo Nacional deCiencia y Tecnologıa fellow (CONACyT 267436)

References Cited

Armendariz-Toledano F A Nino B T Sullivan JMacıas-Samano J Victor S R Clarke and G Zuniga2014a Two species within Dendroctonus frontalis (Coleop-tera Curculionidae) Evidence from morphological karyolog-ical molecular and crossing studies Ann Entomol Soc Am107 11ndash27

Armendariz-Toledano F A Nino J Macıas-Samano andG Zuniga 2014b Review of the geographical distributionof Dendroctonus vitei (Curculionidae Scolytinae) based ongeometric morphometrics of the seminal rod Ann EntomolSoc Am 107 748ndash55

Billings R F 2011 Aerial detection ground evaluation andmonitoring of the southern pine beetle State perspectives


pp 245ndash261 In R N Coulson and K D Klepzig (eds)Southern Pine Beetle II USDA Forest Service SouthernResearch Station Gen Tech Rep SRS-140 Asheville NC

Billings R F S R Clarke V Espino-Mendoza P Cor-don-Cabrera B Melendez Figueroa J R Campos andG Baeza 2004 Bark beetle outbreaks and fire a devastat-ing combination for Central Americarsquos pine forests Unasylva55 15ndash21

Byers J A D L Wood J Craig and L B Hendry 1984Attractive and inhibitory pheromones produced in the barkbeetle Dendroctonus brevicomis during host colonizationRegulation of inter- and intraspecific competition J ChemEcol 10 861ndash877

Cerezke H F 1964 The morphology and functions of the re-productive systems of Dendroctonus monticolae Hopk(Coleoptera Scolytidae) Can Entomol 96 477ndash500

(CONANP) Comision Nacional de Areas Naturales Protegi-das 2007 Programa de conservacion y manejo del ParqueNacional Lagunas de Montebello Secretarıa de Medio Ambi-ente y Recursos Naturales (SEMARNAT) Comision Nacio-nal de Areas Naturales Protegidas (CONANP) Mexico

Furniss M M 2001 A new subspecies of Dendroctonus(Coleoptera Scolytidae) from Mexico Ann Entomol SocAm 94 21ndash25

Gonzalez-Espinosa M S Ochoa-Gaona N Ramırez-Mar-cial and P F Quintana-Ascencio 1997 Contexto vegeta-cional y florıstico de la agricultura pp 85ndash117 In M RParra-Vazquez and B M Dıaz-Hernandez (eds) Los Altosde Chiapas Agricultura y crisis rural Tomo I Los RecursosNaturales El Colegio de la Frontera Sur San Cristobal deLas Casas Chiapas Mexico

Haack R A C M Eckelmann and E Green 2000 South-ern pine beetle outbreak in Belize Newsl Mich EntomolSoc 45 13ndash15

Hadley A 2008 CombineZM version 1 (httpwwwhadley-webpwpblueyondercoukindexhtm)

Hopkins A D 1909 Contributions toward a monograph of thescolytid beetles I the genus Dendroctonus U S Dep AgricBureau Entomol Tech Ser 17 164ndash232

(ICZN) International Code of Zoological Nomenclature1999 International Trust for Zoological Nomenclature 4thed London

Hughes P R JAA Renwick and J P Vite 1976 Identifi-cation and field bioassay of chemical attractants in the round-headed pine beetle Environ Entomol 5 1165ndash1168

Kelley S T and B D Farrell 1998 Is specialization a didend The phylogeny of host use in Dendroctonus bark beetle(Scolytidae) Evolution 52 1731ndash1743

Kirchhoff P 1943 Mesoamerica sus lımites geograficos com-posicion etnica y caracteres culturales Acta Am 1 92ndash107

Lanier G N 1981 Cytotaxonomy of Dendroctonus pp 35ndash65In M W Stock (ed) Applications of genetics and cytology ininsects systematics and evolution Forest Wildlife and RangeExperiment Station University of Idaho Moscow Idaho

Lanier G N and S L Wood 1968 Controlled mating kary-ology morphology and sex ratio in Dendroctonus ponderosacomplex Ann Entomol Soc Am 61 517ndash516

Lanier G N J P Hendrichs and J E Flores 1988 Biosys-tematics of the Dendroctonus frontalis (Coleoptera Scolyti-dae) complex Ann Entomol Soc Am 81 403ndash418

Lyon R L 1958 A useful secondary sex character in Dendroc-tonus bark beetles Can Entomol 90 582ndash584

Macıas-Samano J E E D Green O Sosa S Garcıa andL Hilje 2001 Brotes del descortezador del pino en BeliceRev For Centroam 34 90ndash92

Mendoza-Correa M G and G Zuniga 1991 Determina-cion del sexo en Dendroctonus mexicanus Hopkins (Coleop-tera Scolytidae) a traves del analisis de caracteres

morfologicos abdominales y cefalicos Acta Zool Mex (ns)46 13ndash21

Midtgaard F and K H Thunes 2002 Pine bark bee-tles in the Mountain Pine Ridge Forest Reserve BelizeDescription of the species and advice on monitoring andcombating the beetle infestations p 18 Norwegian ForestryGroup and the Inter-American Development Bank IsadaltoslashNorway

Moreno 2008 Diferenciacion morfologica y comportamentalde dos morfos de Dendroctonus frontalis Zimmermann(Coleoptera Scolytinae) en Montebello Chiapas MS thesisEl Colegio de la Frontera Sur Tapachula Mexico

Moser J C B A Fitzgibbon and K D Klepzig 2005The Mexican pine beetle Dendroctonus mexicanus firstrecord in the United States and co-occurrence with the south-ern pine beetle Dendroctonus frontalis (Coleoptera Scolyti-dae or Curculionidae Scolytinae) Entomol News 116235ndash243

Renwick JAA P R Hughes and J P Vite 1975 The ag-gregation pheromone system of a Dendroctonus bark beetlein Guatemala J Insect Physiol 21 1097ndash1100

Ruiz E J Victor J L Hayes and G Zuniga 2009 Molecu-lar and morphological analysis of Dendroctonus pseoudotsu-gae (Coleoptera Curculionidae Scolytinae) An assessmentof the taxonomic status of subspecies Ann Entomol SocAm 6 982ndash987

Salinas-Moreno Y M G Mendoza M A Barrios RCisneros J E Macıas-Samano and G Zuniga 2004Areography of the genus Dendroctonus (ColeopteraCurculionidae Scolytinae) in Mexico J Biogeogr 311163ndash1177

Sullivan B T A Nino B Moreno C Brownie J Macıas-Samano S R Clarke L R Kirkendall and G Zuniga2012 Biochemical evidence that Dendroctonus frontalis con-sists of two sibling species in Belize and Chiapas MexicoAnn Entomol Soc Am 105 817ndash831

Thunes K H F Midtgaard L R Kirkendall O Leveronand J V Espino 2005 Los gorgojos del pino en Hondurasdescripcion de especies asociaciones de hospederos y meto-dos de monitoreo y control p 32 Norwegian Forestry Groupand the Inter-American Development Bank IsadaltoslashNorway

Thunes K H F Midtgaard L R Kirkendall O Leveronand J V Espino 2011 Manejo sostenible y combate de ladeforestacion en plantaciones de pino en America Central -una alternativa para el triangulo destructivo de las plagas fore-stales incendios y tala Sustanaible management and combatof deforestation in pine plantations in Central America and al-ternative for the destructive triangle of forest pest and illegallogging pp 191ndash226 In D M Chacvarriaga (Ed) Proteccionfitosanitaria forestal Instituto Colombiano AgropecuarioColombia

Thomas J B and D E Bright 1970 A new species of Den-droctonus (Coleoptera Scolytidae) from Mexico Can Ento-mol 102 479ndash483

Vite J P S F Islas JAA Renwick P R Hughes and RA Kliefoth 1974 Biochemical and biological variation ofsouthern pine beetle population in North and Central Amer-ica Z Angew Entomol 75 422ndash435

Vite J P R F Hughes and JAA Renwick 1975 Pine bee-tles of the genus Dendroctonus Pest population in CentralAmerica FAO Bull Plant Prot 6 178ndash184

Wood S L 1963 A revision of the bark beetles genus Dendroc-tonus Erichson (Coleoptera Scolytidae) Great Basin NatMem 23 1ndash117

Wood S L 1974 New synonymy and records of Americanbark beetles (Coleoptera Scolytidae) Great Basin Nat Mem34 277ndash290


Wood S L 1982 The bark and ambrosia beetles of North andCentral America (Coleoptera Scolytidae) a taxonomic mono-graph Great Basin Nat Mem 6 1ndash1359

Zuniga G R Cisneros Y Y Salinas-Moreno 1995 Coexis-tencia de Dendroctonus frontalis Zimmerman y D mexicanusHopkins (Coleoptera Scolytinae) sobre un mismo hospederoActa Zool Mex 64 59ndash62

Zuniga G G Mendoza-Correa R Cisneros Y Y Salinas-Moreno 1999 Zonas de sobreposicion en las areasde

distribucion geografica de las especies mexicanasde Dendroctonus Erichson (Coleoptera Scolytidae) y susimplicaciones ecologico-evolutivas Acta Zool Mex 77 1ndash22

Zuniga G R Cisneros J L Hayes and J Macıas Samano2002 Karyology geographic distribution and origin of the ge-nus Dendroctonus Erichson (Coleoptera Scolytidae) AnnEntomol Soc Am 95 267ndash275Received 17 October 2014

accepted 17 February 2015


sav020-TF1

sav020-TF2

sav020-TF3

From early 2000 to 2002 gt25000 ha of mature pinestands in the Mountain Pine Ridge Forest Reserve ofBelize were devastated by an outbreak of what was ini-tially identified as D frontalis (Haack et al 2000 Ma-cıas-Samano et al 2001 Billings et al 2004) Duringstudies of this outbreak Midtgaard and Thunes (2002)collected a second apparently undescribed Dendrocto-nus species from infested trees that these authorscalled the ldquocaribbean pine beetlerdquo The caribbean pinebeetle species was distinguished from D frontalis bythe presence in both sexes of fine ridges on the pree-pisternal area of the prothorax (anterolateral pronotum)sensu Hopkins (1909 Fig 17) oriented roughly parallelto the anterior margin and by the width of the episto-mal process which appeared to be wider in caribbeanpine beetle than D frontalis (Thunes et al 20052011) These authors assigned the name ldquoDendroctonuswoodirdquo to identify caribbean pine beetle (Thunes et al2005 2011) however according to the InternationalCode of Zoological Nomenclature (ICZN 1999) thisbinomen is a nomen nudum because a formal descrip-tion of the species was not published

Because insufficient biological data supported sepa-ration of the D frontalis and caribbean pine beetle intotwo species Sullivan et al (2012) and Armendariz-Toledano et al (2014a) carried out studies to clarify thetaxonomic status of this bark beetle using crossing ex-periments as well as biochemical morphological karyo-logical and molecular data These studies consistentlysupported the separation of these species Thereforein this paper we describe Dendroctonus mesoamerica-nus sp nov which corresponds to ldquoD frontalis mor-photype Brdquo from Sullivan et al (2012) and Armendariz-Toledano et al (2014a) and to caribbean pine beetlefrom Midtgaard and Thunes (2002)

Materials and Methods

Specimens A total of 289 adult Dendroctonus spnov were examined from 40 collection sites (Table 1)Specimens reported by Armendariz Toledano et al(2014a) corresponding to Dendroctonus sp nov (ieD frontalis morphotype B) were included in the analy-ses Additionally adult specimens of Dendroctonus spnov were collected by the authors from naturallyinfested Pinus sp or graciously loaned by the followinginstitutions Comision Forestal del Estado de Michoa-can Morelia Michoacan El Colegio de la Frontera SurTapachula Chiapas (ECOSUR) Laboratorio de Varia-cion Biologica y Evolucion Escuela Nacional de Cien-cias Biologicas Instituto Politecnico Nacional MexicoColeccion Cientıfica de Entomologıa Forestal Divisionde Ciencias Forestales Universidad Autonoma Cha-pingo (UACH) Estado de Mexico United States ForestService Southern Research Station Pineville LA

Morphology Sex of specimens was determined bya combination of dimorphic characters the size andabundance of large tubercles on the fronts the stridula-tory apparatus in males and genitalia (Lyon 1958Mendoza-Correa and Zuniga 1991) Species identifica-tion of specimens within the D frontalis complex wasperformed using the key of Lanier et al (1988)

Specimens of Dendroctonus sp nov were distinguishedby means of diagnostic characters reported by Armen-dariz-Toledano et al (2014a) for D frontalis morpho-type B ie the presence of striations on preepisternalarea (PEA) the irregular size of ornamentations of theplates of the eighth tergite of females (AP) and theshape of seminal rod (SSR) (Armendariz et al 2014aFig 1)

External morphology of adults was exhaustivelyexamined and photographed with a NIKON SMZ800dissecting microscope and a Nikon DXM1200F digitalcamera Body measurements were taken with this samemicroscope with a micrometer eye piece (at 20ndash63)For some photos of the holotype and paratype 2(Fig 1) images were recorded at successive focaldepths and automatically combined into a single sharpimage (ie focus stacking) using CombineZM software(Hadley 2008) Morphological details of some speci-mens were photographed with an environmental scan-ning electron microscope ESEM (Evo 40 VP ZeissJena Germany)

The elytra and genitalia from 100 specimens of bothsexes were removed These structures were cleared byincubating them at 50C in 10 KOH for 120ndash180 minfor elytra and for 10ndash15 min for male and female geni-talia After incubation structures were immersed in20 acetic acid solution to neutralize the KOH andsubsequently rinsed with 100 ethanol For both malesand females the elytra were cut into two parts (elytraldisc and declivity) and then mounted on the same slideThe eighth abdominal tergite and spermatheca fromindividual females were mounted in the same slideand the seminal rod and genital capsule of individualmales were separated and mounted in the same slide

All structures were mounted semipermanently inglycerol and observed at 100ndash400 with phase contrastmicroscopy Elements of elytral sculpture (puncturesgranules crenulations) were measured and quantifiedwith a phase contrast microscope with an ocularmicrometer (400) Photographs of male and femalegenitalia were taken with a Nikon DXM1200F digitalcamera mounted in a phase contrast microscopeMorphological details of genitalia were photographedwith an ESEM (Evo 40 VP Zeiss)

From the entire series (nfrac14 289) 98 specimens from26 geographic localities were selected to constitute thetype series (Table 1) In the type series six specimensused by Armendariz-Toledano et al (2014a) in the kar-yological analysis were included The genitalia of theholotype allotype and 18 paratypes were extractedphotographed and preserved in a glycerol-filled micro-vial affixed to the same pin as the respective specimenHolotype was indicated with H1 label allotype wasindicated with A1 label and individual paratypes weredesignated from P1 to P96

Dendroctonus mesoamericanusArmendariz-Toledano amp Sullivan new species

(Figs 1ndash3)

Type Material HOLOTYPE male MexicoChiapas La Trinitaria Parque Nacional Lagunas de










Host





























































Fig

4

Geo

grap

hic

ald

istr

ibu

tion

ofD

endr

octo

nus

mes

oam

eric

anus

sp

nov

T

he

Key

sof

loca

litie

sar

esh

owed

inT

able

1B

el

Bel

ize

ES

E

lS

alva

dor

G

ua

Gu

atem

ala

Hon

H

ond

ura

sM

exM

exic

oN

icN

icar

agu

a









Acknowledgments


References Cited
















































sav020-TF1

sav020-TF2

sav020-TF3









Host





























































Fig

4

Geo

grap

hic

ald

istr

ibu

tion

ofD

endr

octo

nus

mes

oam

eric

anus

sp

nov

T

he

Key

sof

loca

litie

sar

esh

owed

inT

able

1B

el

Bel

ize

ES

E

lS

alva

dor

G

ua

Gu

atem

ala

Hon

H

ond

ura

sM

exM

exic

oN

icN

icar

agu

a









Acknowledgments


References Cited
















































sav020-TF1

sav020-TF2

sav020-TF3
















































Fig

4

Geo

grap

hic

ald

istr

ibu

tion

ofD

endr

octo

nus

mes

oam

eric

anus

sp

nov

T

he

Key

sof

loca

litie

sar

esh

owed

inT

able

1B

el

Bel

ize

ES

E

lS

alva

dor

G

ua

Gu

atem

ala

Hon

H

ond

ura

sM

exM

exic

oN

icN

icar

agu

a









Acknowledgments


References Cited
















































sav020-TF1

sav020-TF2

sav020-TF3








Acknowledgments


References Cited
















































sav020-TF1

sav020-TF2

sav020-TF3












































sav020-TF1

sav020-TF2

sav020-TF3

SYSTEMATICS A New Species of Bark Beetle, … · SYSTEMATICS A New Species of Bark Beetle,...

Documents

Transcript of SYSTEMATICS A New Species of Bark Beetle, … · SYSTEMATICS A New Species of Bark Beetle,...