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Improved Identification and New Records of Dendroctonus Bark

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Improved Identification and New Records of Dendroctonus Bark Article Improved Identification and New Records of Dendroctonus Bark Beetles Attacking Pinus contorta in the Subalpine Forest of the Southern Rocky Mountains Javier E. Mercado Rocky Mountain Research Station, USDA/Forest Service, Fort Collins, CO 80526, USA; [email protected] Received: 16 May 2020; Accepted: 3 June 2020; Published: 9 June 2020 Abstract: Research Highlights: Atypical and poorly understood attacks by Dendroctonus rufipennis (Kirby) to Pinus contorta Doug. ex Loudon were detected in the southern Rocky Mountains (SRM). The phenomenon is confirmed across all examined area. Its reproduction is described for the first time as well as the first attacks of D. adjunctus Blandf. in that host. Improved detection and diagnostics of D. rufipennis will allow a simpler, and efficient identification of the species. It will improve the detection capacity by pest detection specialists and entomologists, which will increase our understanding of the phenomena within and beyond the known range. Background and Objectives: In addition to D. ponderosae Hopk. other Dendroctonus species, sometimes together, attacked P. contorta that grew intermixed with Picea engelmannii in the SRM’ subalpine forest. The identification of these beetles was difficult. The goal was to improve the detection and identification of the species from similar Dendroctonus spp. attacking that host and to uncover biological facts about the phenomena. Materials and Methods: Dendroctonus attacking P. contorta were collected along the entire SRM, their attack signs and behavior were recorded. These characteristics were revised from those in the literature and new characters were introduced and tested. Results: The identification of Dendroctonus bark beetles attacking P. contorta in the SRM was improved using revised and new characters including attack signs, attack behavior, and adult beetle characters. An improved identification key couplet is presented to effectively distinguish D. murrayanae from D. rufipennis. Conclusions: Simplified insect identifications that are both accessible to users with different levels of expertise and are based on insect characters, their attack pattern, and signs, like the present, improve detection of insects of interest. Efficient insect detections allow a better understanding of the capabilities they have and the impact they cause to the woodland ecosystems we study, protect, and manage around the globe. Keywords: non-host attack; post-epidemic; facilitation; endemic population strategies 1. Introduction In the southern Rocky Mountains (SRM) of Colorado and southern Wyoming, Pinus contorta Douglas (lodgepole pine) grows from 2400 to 3200 m, intermixing with Picea engelmannii Parry ex Engelm (Engelmann spruce) in the subalpine zone, at elevations above 2800 m [1]. In this temperate mountain forest, native Dendroctonus bark beetles kill trees altering forest structure and species composition. Two Dendroctonus bark beetles attack P. contorta natively, D. ponderosae Hopk. (mountain pine beetle) and D. murrayanae Hopk. (lodgepole pine beetle). The magnitude of the impact on their host is different. For instance, during the most recent epidemic, D. ponderosae killed approximately 60% of the mature P. contorta within the SRMs [2,3], whereas D. murrayanae was reported affecting only small patches of trees [4]. The disparity of the beetles’ impact reflects their different biologies. While D. ponderosae attacks all Pinus species in the SRM, D. murrayanae only attacks P.contorta. Moreover, Forests 2020, 11, 656; doi:10.3390/f11060656 www.mdpi.com/journal/forests Forests 2020, 11, 656 2 of 11 Forests 2020, 11, x FOR PEER REVIEW 2 of 11 D.different ponderosae biologies.occurs While throughout D. ponderosae the SRMs, attacks while allD. Pinus murrayanae species onlyin the occurs SRM, above D. murrayanae latitude 39.4only◦ , asattacks it appears P. contorta to be. limitedMoreover, to cooler,D. ponderosae northern occurs regions throughout [5]. In addition,the SRMs,P. while engelmannii D. murrayanaehas a native only Dendroctonusoccurs aboveenemy, latitudeDendroctonus 39.4°, as it appears rufipennis to(Kirby) be limited (North to cooler, American northern spruce regions beetle), [5]. with In D.addition, ponderosae P. attackingengelmanniiPicea hasspp. a native on rareDendroctonus occasions. enemy, With Dendroctonus reddish elytra rufipennis that contrasts (Kirby) with(North a darkAmerican brown spruce head andbeetle), prothorax, with D. adult ponderosaeD. rufipennis attackingand PiceaD. murrayanaespp. on rareclosely occasions. resemble With reddish each other elytra in color,that contrasts but also inwith size, a shape,dark brown and other head externaland prothorax, characters. adult TheD. rufipennis two Dendroctonus and D. murrayanaeare also di closelyfficult resemble to distinguish each behaviorallyother in color, since but theyalso in have size, similar shape, eggand galleries,other external egg placingcharacters. patterns, The two and Dendroctonus brood aggregation are also patternsdifficult [ to6]. distinguish Therefore, behaviorally the distinction since between they have the similar beetle species egg galleries, is usually egg basedplacing on patterns, the infested and hostbrood [5, 7aggregation]. Dendroctonus patterns rufipennis [6]. Therefore,populations the distinction have been between irruptive the during beetle thespecies last is 20 usually years, killingbased largeon the numbers infested of host suitable [5,7].P. Dendroctonus engelmannii rufipennisin the SRM. populations have been irruptive during the last 20 years,In thekilling SRM, large subalpine numbers forest of suitable coincides P. withengelmannii elevations in the at whichSRM. D. ponderosae activity can regularly becomeIn attenuatedthe SRM, subalpine by low temperatures forest coincides [8]. Consequently,with elevationsD. at ponderosae which D. attacks ponderosae to P. activity contorta canare seldomregularly documented become attenuated above 3000 by m oflow elevation temperatures [8–10]. [8]. However, Consequently, during the D. last ponderosae epidemic, attacksD. ponderosae to P. attackscontorta were are seldom observed documented above 3000 above m providing 3000 m of the elevation opportunity [8–10]. to However, study their during activity the inlast areas epidemic, where theyD. ponderosae occur irregularly. attacks were While observed studying above this 3000 in them providing Roosevelt the National opportunity Forest to in study northern their activity SRM in Colorado,in areas where red-elytra theyDendroctonus occur irregularly.beetles While resembling studyingD. murrayanae this in the and RooseveltD. rufipennis Nationalwere Forest attracted in tonorthern baited (Exobrevicomin-Myrcene-Transverbenol, SRM in Colorado, red‐elytra Dendroctonus Synergy beetles Semiochemicals) resembling funnelD. murrayanae traps (12-funnel and D. Lindgren)rufipennis monitoring were attractedD. ponderosae to baited (Exobrevicominactivity. Due to‐ theirMyrcene resemblance‐Transverbenol, and the Synergy response Semiochemicals) to D. ponderosae lures,funnel that traps included (12‐funnel one Lindgren)P. contorta monitoringtree component, D. ponderosae these beetles activity. were Due di toffi theircult toresemblance identify to and species. the Attacksresponse by to similar D. ponderosae bark beetles lures, werethat included also documented one P. contorta in southern tree component, parts of thethese SRM beetles in Colorado, were wheredifficultD. to murrayanae identify tohas species. not been Attacks documented. by similar bark On allbeetles occasions, were also these documented bark beetles in southern were found parts in mixedof the standsSRM in of Colorado,P. contorta wherewith D.P. murrayanae engelmannii has. Thus, not been the question documented. of whether On all occasions, it was D. murrayanaethese bark orbeetlesD. rufipennis were foundthe species in mixed responsible stands of forP. contorta some of with the attacksP. engelmannii to P. contorta. Thus, inthe the question region’s of subalpine whether forestsit was becameD. murrayanae a topic or of D. relevance. rufipennis the species responsible for some of the attacks to P. contorta in the region’sDendroctonus subalpine rufipennis forestshas became been a topic reported of relevance. attacking P. contorta in the SRMs before, from Dendroctonus rufipennis has been reported attacking P. contorta in the SRMs before, from 1944– 1944–1949 [11] and in 1957 [12] (Figure1); however, a taxonomic authority [ 5,7] contradicted those 1949 [11] and in 1957 [12] (Figure 1); however, a taxonomic authority [5,7] contradicted those determinations. The phenomena of D. rufipennis attacks to P. contorta in the SRM may be poorly determinations. The phenomena of D. rufipennis attacks to P. contorta in the SRM may be poorly understood due to the difficulty of separating it from D. murrayanae attacking that host in subalpine understood due to the difficulty of separating it from D. murrayanae attacking that host in subalpine forests. In this study, Dendroctonus beetles attacking P. contorta in SRM’ subalpine forests of Wyoming forests. In this study, Dendroctonus beetles attacking P. contorta in SRM’ subalpine forests of Wyoming and Colorado were examined. The objectives of this study were to (1) improve the identification and Colorado were examined. The objectives of this study were to (1) improve the identification of of adult Dendroctonus and the detection of their attack signs and patterns, (2) confirm the identity of Dendroctonus spp. with red-elytra attacking P. contorta in subalpine
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