The Structure and Phenology of Non-Native Scolytine Beetle Communities in Coffee Plantations on Kauaʻi

Populations and communities are known to respond to abiotic conditions, but the forces determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.


Introduction
Abiotic factors influence seasona communities (e.g., diversity). Specific k species geographical distributions and modeling, developing conservation str pest species [1][2][3][4]. Many ambrosia and pests of natural ecosystems, forestry, a resources to combat them [5][6][7][8][9]. Promin ponderosae Hopkins, which has devast important pines in the Pacific Northwes compactus (Eichhoff), which is a pest of and the coffee berry borer, Hypothenem pest of coffee production [12]. Much scolytines in the context of damage i Jared Bernard 1, * , Curtis P. Ewing 2 and Russell H. Messing 1 1 Kaua www.mdpi.com/journal/insects to abiotic conditions, but the forces etimes overlooked in the process of ulionidae: Scolytinae) are important ctors that influence their success are host to over three dozen invasive agriculture. We analyzed scolytine d site in a tropical wet forest and a r regression analyses show overall s were four species: the tropical nut lus, and Xyleborinus andrewesi. These ies being markedly dissimilar despite d no influence from temperature or an be valuable for the timing of pest scolytine arrivals on Kauaʻi, and for .
distribution; phenology; Scolytinae; ns and thus the structure of their ctions is invaluable to understanding nces for the purposes of ecological ecies, and directing management of dae: Scolytinae) are important global g tremendous financial and human e mountain pine beetle, Dendroctonus s of ecologically and commercially 10]; the black twig borer, Xylosandrus amental plants, and forest trees [11]; ch is the most impactful pantropical oted to elucidating life histories of developing monitoring and control

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control a, HI 96746, USA; messing@hawaii.edu 2 Research Extension Center, University of Hawai www.mdpi.com/journal/insects dj@hawaii.edu; Tel.: +1-808-320-0566 Accepted: 18 September 2018; Published: 20 September 2018 nd communities are known to respond to abiotic conditions, but the forces ution of particular insect pests are sometimes overlooked in the process of hods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important and ecosystems worldwide, yet the factors that influence their success are ecies. The Hawaiian archipelago is host to over three dozen invasive ich occur on Kauaʻi and are pests of agriculture. We analyzed scolytine t two coffee estates: a hand-harvested site in a tropical wet forest and a site in a tropical dry savanna. Our regression analyses show overall s negatively correlated with rainfall, as were four species: the tropical nut scurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These d to the compositions of the communities being markedly dissimilar despite s richness. Multivariate analysis found no influence from temperature or munity dynamics. This information can be valuable for the timing of pest edicting the success of possible new scolytine arrivals on Kauaʻi, and for ecies may spread with climate change. ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; ence seasonal variation of populations and thus the structure of their ity). Specific knowledge of these interactions is invaluable to understanding tributions and fluctuations of abundances for the purposes of ecological nservation strategies for threatened species, and directing management of ambrosia and bark beetles (Curculionidae: Scolytinae) are important global ms, forestry, and agriculture, requiring tremendous financial and human [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ch has devastated millions of hectares of ecologically and commercially cific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus ich is a pest of hundreds of crops, ornamental plants, and forest trees [11]; r, Hypothenemus hampei (Ferrari), which is the most impactful pantropical n [12]. Much attention has been devoted to elucidating life histories of t of damage to host plants, and to developing monitoring and control eived: 19 August 2018; Accepted: 18 September 2018; Published: 20 September 2018 stract: Populations and communities are known to respond to abiotic conditions, but the forces termining the distribution of particular insect pests are sometimes overlooked in the process of veloping control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important sts of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are known for many species. The Hawaiian archipelago is host to over three dozen invasive lytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine munity dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a chanically harvested site in a tropical dry savanna. Our regression analyses show overall lytine abundance was negatively correlated with rainfall, as were four species: the tropical nut rer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These ationships contributed to the compositions of the communities being markedly dissimilar despite ving the same species richness. Multivariate analysis found no influence from temperature or rvest method on community dynamics. This information can be valuable for the timing of pest trol methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for ecasting how these species may spread with climate change.
ywords: community ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; asive species troduction Abiotic factors influence seasonal variation of populations and thus the structure of their munities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding ies geographical distributions and fluctuations of abundances for the purposes of ecological eling, developing conservation strategies for threatened species, and directing management of species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global s of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human urces to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus erosae Hopkins, which has devastated millions of hectares of ecologically and commercially ortant pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus pactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical of coffee production [12]. Much attention has been devoted to elucidating life histories of ytines in the context of damage to host plants, and to developing monitoring and control i Island in September 2010 [16]. It was subsequently recorded in the Ka www.mdpi.com/journal/insects .edu; Tel.: +1-808-320-0566 18 September 2018; Published: 20 September 2018 unities are known to respond to abiotic conditions, but the forces articular insect pests are sometimes overlooked in the process of and ambrosia beetles (Curculionidae: Scolytinae) are important stems worldwide, yet the factors that influence their success are e Hawaiian archipelago is host to over three dozen invasive on Kauaʻi and are pests of agriculture. We analyzed scolytine fee estates: a hand-harvested site in a tropical wet forest and a tropical dry savanna. Our regression analyses show overall ely correlated with rainfall, as were four species: the tropical nut . brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These mpositions of the communities being markedly dissimilar despite . Multivariate analysis found no influence from temperature or ynamics. This information can be valuable for the timing of pest he success of possible new scolytine arrivals on Kauaʻi, and for spread with climate change.
diversity; spatiotemporal distribution; phenology; Scolytinae; sonal variation of populations and thus the structure of their fic knowledge of these interactions is invaluable to understanding and fluctuations of abundances for the purposes of ecological strategies for threatened species, and directing management of and bark beetles (Curculionidae: Scolytinae) are important global ry, and agriculture, requiring tremendous financial and human minent examples include the mountain pine beetle, Dendroctonus vastated millions of hectares of ecologically and commercially west and Rocky Mountains [10]; the black twig borer, Xylosandrus st of hundreds of crops, ornamental plants, and forest trees [11]; enemus hampei (Ferrari), which is the most impactful pantropical uch attention has been devoted to elucidating life histories of ge to host plants, and to developing monitoring and control ulations and communities are known to respond to abiotic conditions, but the forces he distribution of particular insect pests are sometimes overlooked in the process of ntrol methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important , forestry, and ecosystems worldwide, yet the factors that influence their success are many species. The Hawaiian archipelago is host to over three dozen invasive ny of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine ynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a harvested site in a tropical dry savanna. Our regression analyses show overall dance was negatively correlated with rainfall, as were four species: the tropical nut enemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These ontributed to the compositions of the communities being markedly dissimilar despite me species richness. Multivariate analysis found no influence from temperature or d on community dynamics. This information can be valuable for the timing of pest ds, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for w these species may spread with climate change. mmunity ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; es ctors influence seasonal variation of populations and thus the structure of their .g., diversity). Specific knowledge of these interactions is invaluable to understanding hical distributions and fluctuations of abundances for the purposes of ecological loping conservation strategies for threatened species, and directing management of 4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global l ecosystems, forestry, and agriculture, requiring tremendous financial and human bat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus kins, which has devastated millions of hectares of ecologically and commercially in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus hoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical production [12]. Much attention has been devoted to elucidating life histories of e context of damage to host plants, and to developing monitoring and control i Island in May 2011, Dole's Waialua Estate on O www.mdpi.com/journal/insects tember 2018 biotic conditions, but the forces es overlooked in the process of nidae: Scolytinae) are important that influence their success are to over three dozen invasive iculture. We analyzed scolytine e in a tropical wet forest and a ression analyses show overall re four species: the tropical nut nd Xyleborinus andrewesi. These ing markedly dissimilar despite influence from temperature or valuable for the timing of pest tine arrivals on Kauaʻi, and for ibution; phenology; Scolytinae; nd thus the structure of their s is invaluable to understanding for the purposes of ecological s, and directing management of Scolytinae) are important global mendous financial and human untain pine beetle, Dendroctonus ecologically and commercially the black twig borer, Xylosandrus tal plants, and forest trees [11]; the most impactful pantropical to elucidating life histories of loping monitoring and control ahu in December 2014, and on Maui in December 2016. The richness of the non-native scolytine community in the Hawaiian Islands is thus ideal for analyzing responses of different species to various environmental variables.
The largest coffee estate in the United States is the Kaua t: Populations and communities are known to respond to abiotic conditions, but the forces ining the distribution of particular insect pests are sometimes overlooked in the process of ing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important f crops, forestry, and ecosystems worldwide, yet the factors that influence their success are n for many species. The Hawaiian archipelago is host to over three dozen invasive es, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine nity dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a ically harvested site in a tropical dry savanna. Our regression analyses show overall e abundance was negatively correlated with rainfall, as were four species: the tropical nut ypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These ships contributed to the compositions of the communities being markedly dissimilar despite the same species richness. Multivariate analysis found no influence from temperature or method on community dynamics. This information can be valuable for the timing of pest methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for ting how these species may spread with climate change.
rds: community ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; e species uction otic factors influence seasonal variation of populations and thus the structure of their ities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding eographical distributions and fluctuations of abundances for the purposes of ecological g, developing conservation strategies for threatened species, and directing management of ies [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global natural ecosystems, forestry, and agriculture, requiring tremendous financial and human s to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus e Hopkins, which has devastated millions of hectares of ecologically and commercially t pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus s (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical offee production [12]. Much attention has been devoted to elucidating life histories of s in the context of damage to host plants, and to developing monitoring and control l variation of populations and thus the structure of their nowledge of these interactions is invaluable to understanding fluctuations of abundances for the purposes of ecological ategies for threatened species, and directing management of bark beetles (Curculionidae: Scolytinae) are important global and agriculture, requiring tremendous financial and human ent examples include the mountain pine beetle, Dendroctonus tated millions of hectares of ecologically and commercially st and Rocky Mountains [10]; the black twig borer, Xylosandrus f hundreds of crops, ornamental plants, and forest trees [11]; us hampei (Ferrari), which is the most impactful pantropical attention has been devoted to elucidating life histories of to host plants, and to developing monitoring and control i, where H. hampei has not yet been discovered. Nonetheless, X. compactus is also known to be a pest of coffee [11] and is established on Kaua

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control i, as are many other non-native scolytines. Authors have previously addressed the effect of host availability on scolytine communities [6,8,17]; monocultures of crops present an opportunity to factor out aspects of the community structure related to hosts and focus on the abiotic factors. Uncovering the drivers of phenology and structure of invasive scolytine communities on Kaua stract: Populations and communities are known to respond to abiotic conditions, but the forces termining the distribution of particular insect pests are sometimes overlooked in the process of veloping control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important sts of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are known for many species. The Hawaiian archipelago is host to over three dozen invasive lytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine munity dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a chanically harvested site in a tropical dry savanna. Our regression analyses show overall lytine abundance was negatively correlated with rainfall, as were four species: the tropical nut rer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These ationships contributed to the compositions of the communities being markedly dissimilar despite ving the same species richness. Multivariate analysis found no influence from temperature or rvest method on community dynamics. This information can be valuable for the timing of pest trol methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for ecasting how these species may spread with climate change.
ywords: community ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; asive species troduction Abiotic factors influence seasonal variation of populations and thus the structure of their munities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding ies geographical distributions and fluctuations of abundances for the purposes of ecological eling, developing conservation strategies for threatened species, and directing management of species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global s of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human urces to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus erosae Hopkins, which has devastated millions of hectares of ecologically and commercially ortant pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus pactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical of coffee production [12]. Much attention has been devoted to elucidating life histories of ytines in the context of damage to host plants, and to developing monitoring and control i can also inform management strategies and indicate the potential success of H. hampei or other adventive scolytines, and therefore would be highly useful to protecting Kaua www.mdpi.com/journal/insects Abstract: Populations and communities are known to respond to abiotic conditions, but the forces determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control n to respond to abiotic conditions, but the forces pests are sometimes overlooked in the process of beetles (Curculionidae: Scolytinae) are important de, yet the factors that influence their success are hipelago is host to over three dozen invasive are pests of agriculture. We analyzed scolytine nd-harvested site in a tropical wet forest and a savanna. Our regression analyses show overall ith rainfall, as were four species: the tropical nut halus longipilus, and Xyleborinus andrewesi. These e communities being markedly dissimilar despite nalysis found no influence from temperature or formation can be valuable for the timing of pest ossible new scolytine arrivals on Kauaʻi, and for imate change. iotemporal distribution; phenology; Scolytinae; of populations and thus the structure of their f these interactions is invaluable to understanding s of abundances for the purposes of ecological threatened species, and directing management of (Curculionidae: Scolytinae) are important global ture, requiring tremendous financial and human es include the mountain pine beetle, Dendroctonus ns of hectares of ecologically and commercially Mountains [10]; the black twig borer, Xylosandrus of crops, ornamental plants, and forest trees [11]; Ferrari), which is the most impactful pantropical as been devoted to elucidating life histories of nts, and to developing monitoring and control ations and communities are known to respond to abiotic conditions, but the forces distribution of particular insect pests are sometimes overlooked in the process of trol methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important orestry, and ecosystems worldwide, yet the factors that influence their success are any species. The Hawaiian archipelago is host to over three dozen invasive y of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine amics at two coffee estates: a hand-harvested site in a tropical wet forest and a arvested site in a tropical dry savanna. Our regression analyses show overall ance was negatively correlated with rainfall, as were four species: the tropical nut emus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These ntributed to the compositions of the communities being markedly dissimilar despite e species richness. Multivariate analysis found no influence from temperature or on community dynamics. This information can be valuable for the timing of pest s, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for these species may spread with climate change. munity ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; ors influence seasonal variation of populations and thus the structure of their ., diversity). Specific knowledge of these interactions is invaluable to understanding ical distributions and fluctuations of abundances for the purposes of ecological ping conservation strategies for threatened species, and directing management of . Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global ecosystems, forestry, and agriculture, requiring tremendous financial and human bat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ns, which has devastated millions of hectares of ecologically and commercially n the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus off), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; rry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical oduction [12]. Much attention has been devoted to elucidating life histories of context of damage to host plants, and to developing monitoring and control tors influence seasonal variation of populations and thus the structure of their .g., diversity). Specific knowledge of these interactions is invaluable to understanding hical distributions and fluctuations of abundances for the purposes of ecological loping conservation strategies for threatened species, and directing management of 4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global ecosystems, forestry, and agriculture, requiring tremendous financial and human bat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus kins, which has devastated millions of hectares of ecologically and commercially in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus hoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; erry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical roduction [12]. Much attention has been devoted to elucidating life histories of e context of damage to host plants, and to developing monitoring and control communities are known to respond to abiotic conditions, but the forces ion of particular insect pests are sometimes overlooked in the process of ds. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important d ecosystems worldwide, yet the factors that influence their success are ies. The Hawaiian archipelago is host to over three dozen invasive h occur on Kauaʻi and are pests of agriculture. We analyzed scolytine two coffee estates: a hand-harvested site in a tropical wet forest and a ite in a tropical dry savanna. Our regression analyses show overall negatively correlated with rainfall, as were four species: the tropical nut rus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These to the compositions of the communities being markedly dissimilar despite richness. Multivariate analysis found no influence from temperature or unity dynamics. This information can be valuable for the timing of pest icting the success of possible new scolytine arrivals on Kauaʻi, and for cies may spread with climate change. cology; diversity; spatiotemporal distribution; phenology; Scolytinae; nce seasonal variation of populations and thus the structure of their ). Specific knowledge of these interactions is invaluable to understanding butions and fluctuations of abundances for the purposes of ecological ervation strategies for threatened species, and directing management of brosia and bark beetles (Curculionidae: Scolytinae) are important global s, forestry, and agriculture, requiring tremendous financial and human [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus has devastated millions of hectares of ecologically and commercially ic Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus is a pest of hundreds of crops, ornamental plants, and forest trees [11]; Hypothenemus hampei (Ferrari), which is the most impactful pantropical [12]. wn to respond to abiotic conditions, but the forces pests are sometimes overlooked in the process of beetles (Curculionidae: Scolytinae) are important ide, yet the factors that influence their success are rchipelago is host to over three dozen invasive d are pests of agriculture. We analyzed scolytine and-harvested site in a tropical wet forest and a savanna. Our regression analyses show overall ith rainfall, as were four species: the tropical nut phalus longipilus, and Xyleborinus andrewesi. These the communities being markedly dissimilar despite analysis found no influence from temperature or information can be valuable for the timing of pest ossible new scolytine arrivals on Kauaʻi, and for limate change. tiotemporal distribution; phenology; Scolytinae; of populations and thus the structure of their of these interactions is invaluable to understanding ns of abundances for the purposes of ecological threatened species, and directing management of s (Curculionidae: Scolytinae) are important global lture, requiring tremendous financial and human les include the mountain pine beetle, Dendroctonus ons of hectares of ecologically and commercially y Mountains [10]; the black twig borer, Xylosandrus of crops, ornamental plants, and forest trees [11]; (Ferrari), which is the most impactful pantropical has been devoted to elucidating life histories of ants, and to developing monitoring and control

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control a Bay Coffee is hand-harvested, permitting us to additionally question whether different harvesting methods influence the bark beetle community. By collecting samples for two growing seasons at the Kaua Abstract: Populations and communities are known to respond to abiotic conditions, but the forces determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control munities are known to respond to abiotic conditions, but the forces f particular insect pests are sometimes overlooked in the process of Bark and ambrosia beetles (Curculionidae: Scolytinae) are important cosystems worldwide, yet the factors that influence their success are The Hawaiian archipelago is host to over three dozen invasive cur on Kauaʻi and are pests of agriculture. We analyzed scolytine coffee estates: a hand-harvested site in a tropical wet forest and a in a tropical dry savanna. Our regression analyses show overall tively correlated with rainfall, as were four species: the tropical nut ), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These e compositions of the communities being markedly dissimilar despite ess. Multivariate analysis found no influence from temperature or y dynamics. This information can be valuable for the timing of pest g the success of possible new scolytine arrivals on Kauaʻi, and for may spread with climate change.
gy; diversity; spatiotemporal distribution; phenology; Scolytinae; seasonal variation of populations and thus the structure of their ecific knowledge of these interactions is invaluable to understanding ons and fluctuations of abundances for the purposes of ecological tion strategies for threatened species, and directing management of sia and bark beetles (Curculionidae: Scolytinae) are important global restry, and agriculture, requiring tremendous financial and human . Prominent examples include the mountain pine beetle, Dendroctonus s devastated millions of hectares of ecologically and commercially orthwest and Rocky Mountains [10]; the black twig borer, Xylosandrus pest of hundreds of crops, ornamental plants, and forest trees [11]; othenemus hampei (Ferrari), which is the most impactful pantropical . Much attention has been devoted to elucidating life histories of amage to host plants, and to developing monitoring and control a Bay Coffee, this study compared the species composition of scolytine communities at both sites. To understand how abiotic forces influence scolytines at these sites, we analyzed the fluctuating abundances over time for each species and the community as a whole for correlations with rainfall, temperature, and harvest techniques. We furthermore assessed the correlations between scolytine species to explore interspecies determinants of community structure.

The Kaua
Non-Native n Coffee Plantations pond to abiotic conditions, but the forces e sometimes overlooked in the process of (Curculionidae: Scolytinae) are important he factors that influence their success are o is host to over three dozen invasive ts of agriculture. We analyzed scolytine vested site in a tropical wet forest and a . Our regression analyses show overall fall, as were four species: the tropical nut ngipilus, and Xyleborinus andrewesi. These unities being markedly dissimilar despite found no influence from temperature or ion can be valuable for the timing of pest ew scolytine arrivals on Kauaʻi, and for ange. Abstract: Populations and communities are known to respond to abiotic conditions, but the forces determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human i between Hanapepe River and the Kalaheo Gulch (UTM: 4Q 442039 E, 2421806 N). The visitor center is situated at the center of the estate along Highway 540 at~87.2 m elevation. Because the weather stations around Numila have incomplete data for both the sampling period and the 1981-2010 climate normals, the station from the adjacent district of Kōloa (GHCND: US1HIKI0020) was used. According to the Köppen-Geiger climate classification, the area is a tropical dry savanna (As) [19] with an average annual precipitation of 552.7 mm and air temperature of 19.8-28.3 • C, averaging 24.1 • C [20]. The soil at this location is a fine isohyperthermic silty clay loam with a basic igneous parent material ca. 5 million years old [21]. The estate has been used since 1996 for the production of unshaded coffee (Coffea arabica L.) cultivars, having previously been used since the mid-nineteenth century for intensive www.mdpi.com/journal/insects community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control i Coffee include typica, blue mountain, and, most predominantly, catuai, a Brazilian hybrid cultivar that has varietals of either yellow or red fruit. The Kaua www.mdpi.com/journal/insects awaiian archipelago is host to over three dozen invasive Kauaʻi and are pests of agriculture. We analyzed scolytine estates: a hand-harvested site in a tropical wet forest and a ropical dry savanna. Our regression analyses show overall correlated with rainfall, as were four species: the tropical nut runneus, Cryphalus longipilus, and Xyleborinus andrewesi. These ositions of the communities being markedly dissimilar despite ultivariate analysis found no influence from temperature or amics. This information can be valuable for the timing of pest success of possible new scolytine arrivals on Kauaʻi, and for pread with climate change. iversity; spatiotemporal distribution; phenology; Scolytinae; al variation of populations and thus the structure of their knowledge of these interactions is invaluable to understanding d fluctuations of abundances for the purposes of ecological trategies for threatened species, and directing management of d bark beetles (Curculionidae: Scolytinae) are important global , and agriculture, requiring tremendous financial and human inent examples include the mountain pine beetle, Dendroctonus stated millions of hectares of ecologically and commercially est and Rocky Mountains [10]; the black twig borer, Xylosandrus of hundreds of crops, ornamental plants, and forest trees [11]; mus hampei (Ferrari), which is the most impactful pantropical h attention has been devoted to elucidating life histories of to host plants, and to developing monitoring and control i Coffee Company utilizes mechanical harvesters and processing in an industrial mill, with its annual harvest starting in late September or October and lasting until late December or January, similar to the seasonality in Mesoamerica. This method of harvest strips the berries from the trees, leaving only minute numbers behind. Less accessible terrain on the estate, such as hilltops and gulches, is dominated by haole koa (Leucaena leucocephala [Lam.] de Wit).
Contrarily, Moloa www.mdpi.com/journal/insects wn to respond to abiotic conditions, but the forces pests are sometimes overlooked in the process of beetles (Curculionidae: Scolytinae) are important ide, yet the factors that influence their success are chipelago is host to over three dozen invasive are pests of agriculture. We analyzed scolytine and-harvested site in a tropical wet forest and a savanna. Our regression analyses show overall ith rainfall, as were four species: the tropical nut phalus longipilus, and Xyleborinus andrewesi. These he communities being markedly dissimilar despite analysis found no influence from temperature or nformation can be valuable for the timing of pest ossible new scolytine arrivals on Kauaʻi, and for limate change. tiotemporal distribution; phenology; Scolytinae; of populations and thus the structure of their f these interactions is invaluable to understanding ns of abundances for the purposes of ecological threatened species, and directing management of s (Curculionidae: Scolytinae) are important global lture, requiring tremendous financial and human les include the mountain pine beetle, Dendroctonus ns of hectares of ecologically and commercially y Mountains [10]; the black twig borer, Xylosandrus of crops, ornamental plants, and forest trees [11]; (Ferrari), which is the most impactful pantropical has been devoted to elucidating life histories of nts, and to developing monitoring and control a Bay Coffee uses hand-harvesting more or less year-round. This estate is on the northeast windward shore of Kaua www.mdpi.com/journal/insects epted: 18 September 2018; Published: 20 September 2018 ommunities are known to respond to abiotic conditions, but the forces n of particular insect pests are sometimes overlooked in the process of s. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important ecosystems worldwide, yet the factors that influence their success are s. The Hawaiian archipelago is host to over three dozen invasive occur on Kauaʻi and are pests of agriculture. We analyzed scolytine o coffee estates: a hand-harvested site in a tropical wet forest and a e in a tropical dry savanna. Our regression analyses show overall egatively correlated with rainfall, as were four species: the tropical nut us), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These the compositions of the communities being markedly dissimilar despite chness. Multivariate analysis found no influence from temperature or nity dynamics. This information can be valuable for the timing of pest ting the success of possible new scolytine arrivals on Kauaʻi, and for es may spread with climate change.
ology; diversity; spatiotemporal distribution; phenology; Scolytinae; e seasonal variation of populations and thus the structure of their . Specific knowledge of these interactions is invaluable to understanding utions and fluctuations of abundances for the purposes of ecological rvation strategies for threatened species, and directing management of rosia and bark beetles (Curculionidae: Scolytinae) are important global forestry, and agriculture, requiring tremendous financial and human 9]. Prominent examples include the mountain pine beetle, Dendroctonus has devastated millions of hectares of ecologically and commercially Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus s a pest of hundreds of crops, ornamental plants, and forest trees [11]; ypothenemus hampei (Ferrari), which is the most impactful pantropical 2]. Much attention has been devoted to elucidating life histories of damage to host plants, and to developing monitoring and control Abstract: Populations and communities are known to respond to abiotic conditions, but the forces determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control a (UTM: 4Q 466094 E, 2453687 N). The Anahola weather station (GHCND: US1HIKI0021) provided data for the sampling period, while the 1981-2010 climate normals were obtained from the nearby town of Kīlauea. The area receives an average annual precipitation of 1742.9 mm and a temperature of 19.7-26.8 • C, with an average of 23.3 • C [20], giving it a Köppen-Geiger classification of tropical rainforest (Af ) [19]. Like Numila, this site has very fine isohyperthermic silty clay soil with a basic igneous dust substrate [21]. Moloa ities are known to respond to abiotic conditions, but the forces icular insect pests are sometimes overlooked in the process of nd ambrosia beetles (Curculionidae: Scolytinae) are important ems worldwide, yet the factors that influence their success are Hawaiian archipelago is host to over three dozen invasive n Kauaʻi and are pests of agriculture. We analyzed scolytine estates: a hand-harvested site in a tropical wet forest and a ropical dry savanna. Our regression analyses show overall correlated with rainfall, as were four species: the tropical nut runneus, Cryphalus longipilus, and Xyleborinus andrewesi. These positions of the communities being markedly dissimilar despite ultivariate analysis found no influence from temperature or amics. This information can be valuable for the timing of pest success of possible new scolytine arrivals on Kauaʻi, and for pread with climate change.
iversity; spatiotemporal distribution; phenology; Scolytinae; al variation of populations and thus the structure of their knowledge of these interactions is invaluable to understanding d fluctuations of abundances for the purposes of ecological trategies for threatened species, and directing management of d bark beetles (Curculionidae: Scolytinae) are important global , and agriculture, requiring tremendous financial and human inent examples include the mountain pine beetle, Dendroctonus stated millions of hectares of ecologically and commercially est and Rocky Mountains [10]; the black twig borer, Xylosandrus of hundreds of crops, ornamental plants, and forest trees [11]; emus hampei (Ferrari), which is the most impactful pantropical h attention has been devoted to elucidating life histories of to host plants, and to developing monitoring and control a Bay Coffee estate is 6.1 ha that includes a mix of shaded and unshaded arabica coffee production, all of which is the typica varietal, interspersed with two rows each of cacao (Theobroma cacao L.), Latundan banana (Musa acuminata × balbisiana silk cultivar), as well as citrus trees. This site also includes hedgerows of vinegartree (Lophostemon confertus [R.Br.] Wilson & Waterh.) and elephant-ear tree (Enterolobium cyclocarpum [Jacq.] Griseb.). Areas around the estate are dominated by haole koa. re known to respond to abiotic conditions, but the forces insect pests are sometimes overlooked in the process of brosia beetles (Curculionidae: Scolytinae) are important orldwide, yet the factors that influence their success are iian archipelago is host to over three dozen invasive aʻi and are pests of agriculture. We analyzed scolytine es: a hand-harvested site in a tropical wet forest and a l dry savanna. Our regression analyses show overall lated with rainfall, as were four species: the tropical nut s, Cryphalus longipilus, and Xyleborinus andrewesi. These ons of the communities being markedly dissimilar despite ariate analysis found no influence from temperature or . This information can be valuable for the timing of pest ss of possible new scolytine arrivals on Kauaʻi, and for with climate change. ty; spatiotemporal distribution; phenology; Scolytinae; riation of populations and thus the structure of their ledge of these interactions is invaluable to understanding ctuations of abundances for the purposes of ecological ies for threatened species, and directing management of beetles (Curculionidae: Scolytinae) are important global agriculture, requiring tremendous financial and human examples include the mountain pine beetle, Dendroctonus millions of hectares of ecologically and commercially d Rocky Mountains [10]; the black twig borer, Xylosandrus dreds of crops, ornamental plants, and forest trees [11]; ampei (Ferrari), which is the most impactful pantropical ntion has been devoted to elucidating life histories of ost plants, and to developing monitoring and control a, we sampled only during the latter period. We excluded both Junes from statistical analysis, however, because we sampled during only part of the month. We collected scolytines in Brocap ® traps (Cirad, Montpellier, France). To maintain the efficacy of the traps, every three months we replaced each trap's 3:1 methanol-ethanol lure (Scentry Biologicals, Inc., Billings, MT, USA) and 2.5 × 2.5 cm Vaportape TM II insecticidal strip (2,2-dichlorovinyldimethylphosphate; Aberdeen Road Company, Emigsville, PA, USA). Despite the recommendation by Scentry Biologicals, Inc. to replace the lures every three weeks, we found them to be effective for at least three months, with no impact on collection (data not shown). Although intended for surveying for H. hampei [22], this trapping system proved to be a non-specific means for collecting not only nearly two dozen species of scolytines, but also many other pests of agriculture, including several species each of nitidulids, bostrichids, anthribids, cerambycids, and coccinellids, as well as many hemipterans, hymenopterans (including sphecids, ants, and numerous parasitoids), blattellids, dipterans, lepidopterans, and other insects (data not shown). We hence used only these traps, thereby standardizing our collection methods to avoid accounting for variable trapping methods in our data. We placed the traps to encapsulate as much of each property as possible. At the Kaua communities are known to respond to abiotic conditions, but the forces ion of particular insect pests are sometimes overlooked in the process of ds. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important d ecosystems worldwide, yet the factors that influence their success are ies. The Hawaiian archipelago is host to over three dozen invasive h occur on Kauaʻi and are pests of agriculture. We analyzed scolytine two coffee estates: a hand-harvested site in a tropical wet forest and a ite in a tropical dry savanna. Our regression analyses show overall negatively correlated with rainfall, as were four species: the tropical nut rus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These to the compositions of the communities being markedly dissimilar despite richness. Multivariate analysis found no influence from temperature or unity dynamics. This information can be valuable for the timing of pest icting the success of possible new scolytine arrivals on Kauaʻi, and for cies may spread with climate change.

Experimental Design
cology; diversity; spatiotemporal distribution; phenology; Scolytinae; nce seasonal variation of populations and thus the structure of their ). Specific knowledge of these interactions is invaluable to understanding butions and fluctuations of abundances for the purposes of ecological ervation strategies for threatened species, and directing management of brosia and bark beetles (Curculionidae: Scolytinae) are important global s, forestry, and agriculture, requiring tremendous financial and human [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus has devastated millions of hectares of ecologically and commercially ic Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus i Coffee Company at Numila, we placed 12 traps: seven in the fields at the center of the property, two flanking the hopper at the mill (to encounter specimens transported from distant parts of the property), and three at the corners of the estate. We positioned two traps at opposite ends of the Moloa seasonal variation of populations and thus the structure of their ecific knowledge of these interactions is invaluable to understanding ons and fluctuations of abundances for the purposes of ecological tion strategies for threatened species, and directing management of sia and bark beetles (Curculionidae: Scolytinae) are important global restry, and agriculture, requiring tremendous financial and human a Bay Coffee property, which is proportionally adequate given that this estate is 0.5% the size of the estate at Numila. To ameliorate potential edge effects, traps near the perimeters of each estate were a minimum of 6 m from the limits of coffee production, beyond which was another 3-10 m of buffer to the edge of the property. The predominant haole koa surrounding each estate would further help to mitigate differences between sites.

Analysis
After identifying specimens following Wood [23,24], Beaver and Maddison [25], and Samuelson [14], we preserved them in 95% EtOH and verified vouchers prior to accessioning at the University of differed between the sites, we calculated the similarity between the communities using relative abundances with the Bray-Curtis dissimilarity index. The Shannon-Wiener index enabled us to assess the diversity because it can be related to species richness to determine the similarity of species' abundances or species evenness. As this provides a snapshot of the community structure, we plotted these metrics over time to evaluate the communities' structure in the context of phenology. We assessed log-transformed data (x' = log 10 [x + 1]) for normality using the Shapiro-Wilk test and for skewness using Pearson's coefficient before conducting regression analyses with R 3.5.0 [26] to discern correlation of the communities' phenology to abiotic factors, both within and among sites. When both beetle and environmental data were normally distributed, we used parametric linear regression, and for the others we used Kendall's τ b correlation. Multiple analysis of variance (MANOVA) tests enabled us to further assess any potential interactive effects of the variables. Finally, to analyze additional influences of the structure of the communities from beetle-beetle interactions, we constructed a scale-free correlation network using the R package "igraph" [27] with log-transformed abundance data. With the aid of the R package "Matrix" [28], we filtered the correlation network for significant p-values. We projected the networks using the Fruchterman-Reingold algorithm because it employs natural repellent forces to display distances between nodes [29].

Results
This study identified 20 species of scolytines, with 4841 specimens collected and preserved (raw data deposited on Figshare.com). Three of these species are new records for Kaua

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control unities are known to respond to abiotic conditions, but the forces articular insect pests are sometimes overlooked in the process of k and ambrosia beetles (Curculionidae: Scolytinae) are important ystems worldwide, yet the factors that influence their success are e Hawaiian archipelago is host to over three dozen invasive r on Kauaʻi and are pests of agriculture. We analyzed scolytine ffee estates: a hand-harvested site in a tropical wet forest and a a tropical dry savanna. Our regression analyses show overall ely correlated with rainfall, as were four species: the tropical nut . brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These ompositions of the communities being markedly dissimilar despite s. Multivariate analysis found no influence from temperature or ynamics. This information can be valuable for the timing of pest the success of possible new scolytine arrivals on Kauaʻi, and for y spread with climate change.
; diversity; spatiotemporal distribution; phenology; Scolytinae; sonal variation of populations and thus the structure of their ific knowledge of these interactions is invaluable to understanding s and fluctuations of abundances for the purposes of ecological n strategies for threatened species, and directing management of and bark beetles (Curculionidae: Scolytinae) are important global try, and agriculture, requiring tremendous financial and human ominent examples include the mountain pine beetle, Dendroctonus evastated millions of hectares of ecologically and commercially hwest and Rocky Mountains [10]; the black twig borer, Xylosandrus est of hundreds of crops, ornamental plants, and forest trees [11]; henemus hampei (Ferrari), which is the most impactful pantropical uch attention has been devoted to elucidating life histories of age to host plants, and to developing monitoring and control ive species duction biotic factors influence seasonal variation of populations and thus the structure of their nities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding geographical distributions and fluctuations of abundances for the purposes of ecological ing, developing conservation strategies for threatened species, and directing management of ecies [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global f natural ecosystems, forestry, and agriculture, requiring tremendous financial and human ces to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus sae Hopkins, which has devastated millions of hectares of ecologically and commercially ant pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus tus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; e coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical f coffee production [12]. Much attention has been devoted to elucidating life histories of nes in the context of damage to host plants, and to developing monitoring and control  (Figures 1 and 2). We calculated the Shannon-Wiener diversity index (H') using monthly totals to describe changes to the community structure over time, and superimposed over this is the monthly species evenness (E), based on species richness, for both Moloa Abstract: Populations and communities are known to respond to abiotic conditions, but the forces determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control a and Numila (Figure 3).  ulations and communities are known to respond to abiotic conditions, but the forces the distribution of particular insect pests are sometimes overlooked in the process of ontrol methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important s, forestry, and ecosystems worldwide, yet the factors that influence their success are r many species. The Hawaiian archipelago is host to over three dozen invasive any of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine ynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a harvested site in a tropical dry savanna. Our regression analyses show overall ndance was negatively correlated with rainfall, as were four species: the tropical nut henemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These contributed to the compositions of the communities being markedly dissimilar despite ame species richness. Multivariate analysis found no influence from temperature or od on community dynamics. This information can be valuable for the timing of pest ods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for ow these species may spread with climate change. ommunity ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; ies n actors influence seasonal variation of populations and thus the structure of their e.g., diversity). Specific knowledge of these interactions is invaluable to understanding phical distributions and fluctuations of abundances for the purposes of ecological eloping conservation strategies for threatened species, and directing management of -4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global al ecosystems, forestry, and agriculture, requiring tremendous financial and human mbat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus pkins, which has devastated millions of hectares of ecologically and commercially s in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus hhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical production [12]. Much attention has been devoted to elucidating life histories of Abstract: Populations and communities are known to respond to abiotic conditions, but the forces determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of  Abstract: Populations and communities are known to respond to abiotic conditions, but the forces determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of ct: Populations and communities are known to respond to abiotic conditions, but the forces ining the distribution of particular insect pests are sometimes overlooked in the process of ping control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important f crops, forestry, and ecosystems worldwide, yet the factors that influence their success are wn for many species. The Hawaiian archipelago is host to over three dozen invasive nes, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine nity dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a nically harvested site in a tropical dry savanna. Our regression analyses show overall ne abundance was negatively correlated with rainfall, as were four species: the tropical nut Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These nships contributed to the compositions of the communities being markedly dissimilar despite the same species richness. Multivariate analysis found no influence from temperature or t method on community dynamics. This information can be valuable for the timing of pest l methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for ting how these species may spread with climate change. rds: community ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; e species uction iotic factors influence seasonal variation of populations and thus the structure of their ities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding geographical distributions and fluctuations of abundances for the purposes of ecological  Relative humidity data were lacking during the sampling period at both sites, and so were excluded from the analysis aside from the inference via rainfall (rain being 100% RH). Therefore, the abiotic factors analyzed in this study included temperature, precipitation, and harvest techniques. Temperature was not significantly different between the two sites (F(1,11.09) = 3.98, p = 0.07); as temperature remained relatively stable year-round, we found no correlation between precipitation and temperature ((F(1,22) = 0.34, p = 0.57). Linear regressions also found no statistical trends between temperature and abundance of scolytines, either for the overall communities (F(1,22) = 1.85, p = 0.19) or for individual species (listed in Table S1). We also note that fluctuation in scolytine abundance did not align with dates on which we replaced the MeOH-EtOH lures (τb = −0.12, p = 0.50). Figure 4 shows total monthly abundances of scolytines at both Numila and Moloaʻa overlaid against monthly rainfall. Pooled data from Moloaʻa and Numila had normal distribution (p = 0.12 for scolytine abundances, p = 0.99 for precipitation from nearby weather stations; n = 23 months). pulations and communities are known to respond to abiotic conditions, but the forces the distribution of particular insect pests are sometimes overlooked in the process of control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important s, forestry, and ecosystems worldwide, yet the factors that influence their success are r many species. The Hawaiian archipelago is host to over three dozen invasive any of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a y harvested site in a tropical dry savanna. Our regression analyses show overall undance was negatively correlated with rainfall, as were four species: the tropical nut thenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These s contributed to the compositions of the communities being markedly dissimilar despite ame species richness. Multivariate analysis found no influence from temperature or hod on community dynamics. This information can be valuable for the timing of pest ods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for ow these species may spread with climate change. community ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; cies n factors influence seasonal variation of populations and thus the structure of their (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding aphical distributions and fluctuations of abundances for the purposes of ecological veloping conservation strategies for threatened species, and directing management of -4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global ral ecosystems, forestry, and agriculture, requiring tremendous financial and human ombat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus pkins, which has devastated millions of hectares of ecologically and commercially a Bay Coffee.
Relative humidity data were lacking during the sampling period at both sites, and so were excluded from the analysis aside from the inference via rainfall (rain being 100% RH). Therefore, the abiotic factors analyzed in this study included temperature, precipitation, and harvest techniques. Temperature was not significantly different between the two sites (F (1,11.09) = 3.98, p = 0.07); as temperature remained relatively stable year-round, we found no correlation between precipitation and temperature ((F (1,22) = 0.34, p = 0.57). Linear regressions also found no statistical trends between temperature and abundance of scolytines, either for the overall communities (F (1,22) = 1.85, p = 0.19) or for individual species (listed in Table S1). We also note that fluctuation in scolytine abundance did not align with dates on which we replaced the MeOH-EtOH lures (τ b = −0.12, p = 0.50).

Figure 4 shows total monthly abundances of scolytines at both Numila and Moloa
Insects 2018, 9, x; doi: www.mdpi.com/journal/insects community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control a overlaid against monthly rainfall. Pooled data from Moloa www.mdpi.com/journal/insects which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine s at two coffee estates: a hand-harvested site in a tropical wet forest and a ted site in a tropical dry savanna. Our regression analyses show overall was negatively correlated with rainfall, as were four species: the tropical nut obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These uted to the compositions of the communities being markedly dissimilar despite cies richness. Multivariate analysis found no influence from temperature or ommunity dynamics. This information can be valuable for the timing of pest predicting the success of possible new scolytine arrivals on Kauaʻi, and for e species may spread with climate change. ity ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; nfluence seasonal variation of populations and thus the structure of their ersity). Specific knowledge of these interactions is invaluable to understanding distributions and fluctuations of abundances for the purposes of ecological conservation strategies for threatened species, and directing management of ny ambrosia and bark beetles (Curculionidae: Scolytinae) are important global stems, forestry, and agriculture, requiring tremendous financial and human em [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus hich has devastated millions of hectares of ecologically and commercially Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus hich is a pest of hundreds of crops, ornamental plants, and forest trees [11]; orer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical tion [12]. Much attention has been devoted to elucidating life histories of text of damage to host plants, and to developing monitoring and control a and Numila had normal distribution (p = 0.12 for scolytine abundances, p = 0.99 for precipitation from nearby weather stations; n = 23 months). Pearson's coefficient of skewness (Sk 2 ) also showed that sample distribution was not significantly skewed in either data set (Sk 2 = −0.27 for scolytines, Sk 2 = 0.38 for rainfall), both being less than twice the standard error of the skewness, and their variances were equal (F = 1.20, critical value = 2.04). A linear regression revealed a negative correlation between total scolytine abundance and precipitation (F (1,22) = 12.21, p = 0.00) with Pearson's correlation coefficient r = −0.62 and R 2 = 0.38 ( Figure 5). When analyzing the sites separately, a significant negative correlation with rainfall also emerged independently at Numila (F (1,15) = 4.98, p = 0.04, R 2 = 0.28), although the trend did not arise at the wetter Moloa www.mdpi.com/journal/insects ed: 20 September 2018 ond to abiotic conditions, but the forces sometimes overlooked in the process of urculionidae: Scolytinae) are important e factors that influence their success are is host to over three dozen invasive s of agriculture. We analyzed scolytine sted site in a tropical wet forest and a Our regression analyses show overall ll, as were four species: the tropical nut gipilus, and Xyleborinus andrewesi. These nities being markedly dissimilar despite ound no influence from temperature or n can be valuable for the timing of pest w scolytine arrivals on Kauaʻi, and for nge. al distribution; phenology; Scolytinae; lations and thus the structure of their teractions is invaluable to understanding ndances for the purposes of ecological d species, and directing management of onidae: Scolytinae) are important global iring tremendous financial and human e the mountain pine beetle, Dendroctonus tares of ecologically and commercially ins [10]; the black twig borer, Xylosandrus ornamental plants, and forest trees [11]; hich is the most impactful pantropical devoted to elucidating life histories of to developing monitoring and control a site (F (1,8) = 0.00, p = 0.97). However, a MANOVA found no significant combined effect of precipitation and temperature on scolytine abundance.         www.mdpi.com/journal/insects mmunity dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a chanically harvested site in a tropical dry savanna. Our regression analyses show overall lytine abundance was negatively correlated with rainfall, as were four species: the tropical nut rer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These ationships contributed to the compositions of the communities being markedly dissimilar despite ving the same species richness. Multivariate analysis found no influence from temperature or rvest method on community dynamics. This information can be valuable for the timing of pest ntrol methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for ecasting how these species may spread with climate change.
ywords: community ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; asive species troduction Abiotic factors influence seasonal variation of populations and thus the structure of their munities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding cies geographical distributions and fluctuations of abundances for the purposes of ecological eling, developing conservation strategies for threatened species, and directing management of t species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global ts of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human urces to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus derosae Hopkins, which has devastated millions of hectares of ecologically and commercially ortant pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus pactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical t of coffee production [12]. Much attention has been devoted to elucidating life histories of ytines in the context of damage to host plants, and to developing monitoring and control a Bay Coffee with the one at Kaua www.mdpi.com/journal/insects iculture. We analyzed scolytine e in a tropical wet forest and a gression analyses show overall ere four species: the tropical nut and Xyleborinus andrewesi. These eing markedly dissimilar despite influence from temperature or e valuable for the timing of pest tine arrivals on Kauaʻi, and for ibution; phenology; Scolytinae; and thus the structure of their s is invaluable to understanding s for the purposes of ecological s, and directing management of Scolytinae) are important global emendous financial and human untain pine beetle, Dendroctonus f ecologically and commercially the black twig borer, Xylosandrus ntal plants, and forest trees [11]; the most impactful pantropical to elucidating life histories of loping monitoring and control i Coffee Company, we assessed only the 2016-2017 growing season when both sites were surveyed (n = 7 months). Relative monthly abundances (i.e., density) were used to eliminate the bias of the sampling area. Although Pearson's coefficient of skewness indicated that the data were not significantly skewed (Sk 2 = 1.40 for Moloa www.mdpi.com/journal/insects Bark and ambrosia beetles (Curculionidae: Scolytinae) are important cosystems worldwide, yet the factors that influence their success are The Hawaiian archipelago is host to over three dozen invasive ccur on Kauaʻi and are pests of agriculture. We analyzed scolytine coffee estates: a hand-harvested site in a tropical wet forest and a in a tropical dry savanna. Our regression analyses show overall atively correlated with rainfall, as were four species: the tropical nut ), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These e compositions of the communities being markedly dissimilar despite ness. Multivariate analysis found no influence from temperature or ty dynamics. This information can be valuable for the timing of pest ng the success of possible new scolytine arrivals on Kauaʻi, and for may spread with climate change.
ogy; diversity; spatiotemporal distribution; phenology; Scolytinae; seasonal variation of populations and thus the structure of their pecific knowledge of these interactions is invaluable to understanding ions and fluctuations of abundances for the purposes of ecological ation strategies for threatened species, and directing management of sia and bark beetles (Curculionidae: Scolytinae) are important global restry, and agriculture, requiring tremendous financial and human . Prominent examples include the mountain pine beetle, Dendroctonus s devastated millions of hectares of ecologically and commercially orthwest and Rocky Mountains [10]; the black twig borer, Xylosandrus a pest of hundreds of crops, ornamental plants, and forest trees [11]; pothenemus hampei (Ferrari), which is the most impactful pantropical . Much attention has been devoted to elucidating life histories of amage to host plants, and to developing monitoring and control a, Sk 2 = 1.31 for Numila), the Shapiro-Wilk test found the log-transformed data to have a non-normal distribution (p = 0.05 for Moloa Insects 2018, 9, x; doi: www.mdpi.com/journal/insects determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control a, p = 0.00 for Numila) and unequal variances (F = 0.94, critical value = 0.23). The non-parametric Mann-Whitney U test rejected significant difference of total monthly scolytine abundances between the two sites (U = 23, c.v. = 8). In contrasting monthly species richness (adjusted per trap), we found no difference between Numila and Moloa www.mdpi.com/journal/insects : 20 September 2018 d to abiotic conditions, but the forces metimes overlooked in the process of rculionidae: Scolytinae) are important factors that influence their success are host to over three dozen invasive f agriculture. We analyzed scolytine ed site in a tropical wet forest and a ur regression analyses show overall as were four species: the tropical nut ilus, and Xyleborinus andrewesi. These ties being markedly dissimilar despite nd no influence from temperature or can be valuable for the timing of pest scolytine arrivals on Kauaʻi, and for e. distribution; phenology; Scolytinae; ions and thus the structure of their actions is invaluable to understanding ances for the purposes of ecological pecies, and directing management of idae: Scolytinae) are important global ing tremendous financial and human he mountain pine beetle, Dendroctonus res of ecologically and commercially [10]; the black twig borer, Xylosandrus namental plants, and forest trees [11]; ich is the most impactful pantropical voted to elucidating life histories of developing monitoring and control a (U = 18, c.v. = 8). As a test of the relationship of monthly scolytine abundances to harvest at Numila, we calculated the area of remaining coffee fields adjacent to traps from the total area of the Kaua Abstract: Populations and communities are known to respond to abiotic conditions, but the forces determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control i Coffee Company according to their harvest dates. Regression analyses of scolytine abundance against harvested area over two harvest seasons showed no trend, either for the community overall (F (1,9) = 0.26, p = 0.63) or for individual species. We also found no interactive effects of harvest with temperature and/or precipitation on scolytine abundance (F (1,9) = 0.22, p = 0.81).
Additional Mann-Whitney tests allowed us to detect differences between sites for individual species' densities, with values adjusted per trap. We found significant differences for the following species: Xylosandrus compactus (U = 0, c.v. = 8; Figure 7a = 7 months). Relative monthly abundances (i.e., density) were used to eliminate the bias of the sampling area. Although Pearson's coefficient of skewness indicated that the data were not significantly skewed (Sk2 = 1.40 for Moloaʻa, Sk2 = 1.31 for Numila), the Shapiro-Wilk test found the log-transformed data to have a non-normal distribution (p = 0.05 for Moloaʻa, p = 0.00 for Numila) and unequal variances (F = 0.94, critical value = 0.23). The non-parametric Mann-Whitney U test rejected significant difference of total monthly scolytine abundances between the two sites (U = 23, c.v. = 8). In contrasting monthly species richness (adjusted per trap), we found no difference between Numila and Moloaʻa (U = 18, c.v. = 8).
As a test of the relationship of monthly scolytine abundances to harvest at Numila, we calculated the area of remaining coffee fields adjacent to traps from the total area of the Kauaʻi Coffee Company according to their harvest dates. Regression analyses of scolytine abundance against harvested area over two harvest seasons showed no trend, either for the community overall (F(1,9) = 0.26, p = 0.63) or for individual species. We also found no interactive effects of harvest with temperature and/or precipitation on scolytine abundance (F(1,9) = 0.22, p = 0.81).
Additional Mann-Whitney tests allowed us to detect differences between sites for individual species' densities, with values adjusted per trap. We found significant differences for the following species: Xylosandrus compactus (U = 0, c.v. = 8; Figure 7a  To address any influence of beetle-beetle interactions on the communities, we constructed scalefree correlation networks. Figure 8 shows only relationships wherein R 2 > 0.35, and displays significant correlations (p < 0.05) alongside weak trends (0.1 > p > 0.05) to give a more complete depiction of the community structure. There were no significant negative correlations at either site. At Numila (Figure 8a  ulations and communities are known to respond to abiotic conditions, but the forces the distribution of particular insect pests are sometimes overlooked in the process of ntrol methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important , forestry, and ecosystems worldwide, yet the factors that influence their success are many species. The Hawaiian archipelago is host to over three dozen invasive any of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine ynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a harvested site in a tropical dry savanna. Our regression analyses show overall ndance was negatively correlated with rainfall, as were four species: the tropical nut enemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These contributed to the compositions of the communities being markedly dissimilar despite me species richness. Multivariate analysis found no influence from temperature or od on community dynamics. This information can be valuable for the timing of pest ds, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for w these species may spread with climate change. ommunity ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; ies ctors influence seasonal variation of populations and thus the structure of their .g., diversity). Specific knowledge of these interactions is invaluable to understanding To address any influence of beetle-beetle interactions on the communities, we constructed scale-free correlation networks. Figure 8 shows only relationships wherein R 2 > 0.35, and displays significant correlations (p < 0.05) alongside weak trends (0.1 > p > 0.05) to give a more complete depiction of the community structure. There were no significant negative correlations at either site. At Numila (Figure 8a) www.mdpi.com/journal/insects f crops, forestry, and ecosystems worldwide, yet the factors that influence their success are wn for many species. The Hawaiian archipelago is host to over three dozen invasive nes, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine unity dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a nically harvested site in a tropical dry savanna. Our regression analyses show overall ne abundance was negatively correlated with rainfall, as were four species: the tropical nut (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These nships contributed to the compositions of the communities being markedly dissimilar despite the same species richness. Multivariate analysis found no influence from temperature or t method on community dynamics. This information can be valuable for the timing of pest l methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for sting how these species may spread with climate change. rds: community ecology; diversity; spatiotemporal distribution; phenology; Scolytinae; e species duction iotic factors influence seasonal variation of populations and thus the structure of their nities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding geographical distributions and fluctuations of abundances for the purposes of ecological g, developing conservation strategies for threatened species, and directing management of cies [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global natural ecosystems, forestry, and agriculture, requiring tremendous financial and human es to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ae Hopkins, which has devastated millions of hectares of ecologically and commercially nt pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus us (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical coffee production [12]. Much attention has been devoted to elucidating life histories of es in the context of damage to host plants, and to developing monitoring and control a (Figure 8b), we found significant correlations between C. longipilus and H. eruditus (R 2 = 0.93), and between E. fornicatus and Ptilopodius pacificus Schedl (R 2 = 0.97). The strong correlations present at Numila were weak at Moloa Insects 2018, 9, x; doi: www.mdpi.com/journal/insects determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control a, and those in Moloa www.mdpi.com/journal/insects to abiotic conditions, but the forces etimes overlooked in the process of ulionidae: Scolytinae) are important ctors that influence their success are host to over three dozen invasive agriculture. We analyzed scolytine site in a tropical wet forest and a regression analyses show overall s were four species: the tropical nut us, and Xyleborinus andrewesi. These es being markedly dissimilar despite no influence from temperature or n be valuable for the timing of pest colytine arrivals on Kauaʻi, and for .
istribution; phenology; Scolytinae; ns and thus the structure of their tions is invaluable to understanding nces for the purposes of ecological ecies, and directing management of ae: Scolytinae) are important global g tremendous financial and human mountain pine beetle, Dendroctonus s of ecologically and commercially 10]; the black twig borer, Xylosandrus mental plants, and forest trees [11]; h is the most impactful pantropical oted to elucidating life histories of eveloping monitoring and control a were not present in Numila.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of

Discussion
We found a significant negative correlation between precipitation and total scolytine community abundance ( Figure 5). Although this phenomenon initially appears to be a lag of scolytine success following coffee productivity, which itself would lag after a period of heavier rain [1], some data contradict this speculation. Increases in scolytine abundance in Numila in February and August of 2016, for example, did not follow rainy months, and increases that did follow rainy periods tended to be a few months later (Figure 4). Another explanation for this trend is that some scolytines could prefer drier weather. Spatial distribution modeling in Kenya by Jaramillo et al. [12] indicated that Hypothenemus hampei is more successful in dry climates than in wet ones.
Our data show that other scolytine species are also more abundant in such conditions; supporting the pattern of the overall community, we also found significant inverse correlations with precipitation for four species: H. obscurus, H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi ( Figure 6). Moreover, the scale-free interspecies correlation networks found no significant positive correlations between these four species (Figure 8), which demonstrates that they each have independent relationships with rainfall and that other disparate factors influence their numbers. This indicates that the correlations shown in the interspecies networks provide additive effects to the overall community structure, alongside the importance of rainfall, rather than reflect an indirect effect of species responding to the same forces.
In parsing out the independent variables in a manner similar to Park and Reid [30], we surprisingly found no correlation between beetle abundances and either temperature or harvest times, despite harvest techniques being one of the most obvious differences between the sites. This may hint that scolytines in these communities have little to do with coffee fruit, but we occasionally reared H. obscurus, Xylosandrus compactus, Xyleborus affinis Eichhoff, and Coccotrypes carpophagus (Hornung) from coffee fruit collected at these estates.
Despite the correlation between overall scolytine density and precipitation, this study could not find a correlation between diversity and precipitation. Nevertheless, diversity and evenness of the community at Numila tended to decline as the total abundance of scolytines increased (Figures 3 and 4). In February 2016 and January 2017, total scolytine abundance peaked when the community was less diverse. This is appears to be owing to the proportional dominance of H. obscurus (Figures 1 and 2), although we found no negative correlations between it and other species (Figure 8a). Species evenness was conversely highest in the summer months when overall scolytine abundance was relatively low for the year. ties are known to respond to abiotic conditions, but the forces icular insect pests are sometimes overlooked in the process of nd ambrosia beetles (Curculionidae: Scolytinae) are important ms worldwide, yet the factors that influence their success are awaiian archipelago is host to over three dozen invasive Kauaʻi and are pests of agriculture. We analyzed scolytine estates: a hand-harvested site in a tropical wet forest and a ropical dry savanna. Our regression analyses show overall correlated with rainfall, as were four species: the tropical nut unneus, Cryphalus longipilus, and Xyleborinus andrewesi. These ositions of the communities being markedly dissimilar despite ultivariate analysis found no influence from temperature or mics. This information can be valuable for the timing of pest success of possible new scolytine arrivals on Kauaʻi, and for read with climate change. versity; spatiotemporal distribution; phenology; Scolytinae; al variation of populations and thus the structure of their knowledge of these interactions is invaluable to understanding d fluctuations of abundances for the purposes of ecological rategies for threatened species, and directing management of bark beetles (Curculionidae: Scolytinae) are important global and agriculture, requiring tremendous financial and human inent examples include the mountain pine beetle, Dendroctonus stated millions of hectares of ecologically and commercially st and Rocky Mountains [10]; the black twig borer, Xylosandrus f hundreds of crops, ornamental plants, and forest trees [11]; mus hampei (Ferrari), which is the most impactful pantropical attention has been devoted to elucidating life histories of to host plants, and to developing monitoring and control a depict a rise in diversity alongside a rise in overall abundance in January 2017, peak evenness for the community was in October 2016 when the abundance was lowest.
The Abstract: Populations and communities are known to respond to abiotic conditions, but the forces determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human to abiotic conditions, but the forces etimes overlooked in the process of ulionidae: Scolytinae) are important tors that influence their success are ost to over three dozen invasive agriculture. We analyzed scolytine site in a tropical wet forest and a regression analyses show overall s were four species: the tropical nut s, and Xyleborinus andrewesi. These s being markedly dissimilar despite no influence from temperature or n be valuable for the timing of pest colytine arrivals on Kauaʻi, and for istribution; phenology; Scolytinae; ns and thus the structure of their tions is invaluable to understanding nces for the purposes of ecological ecies, and directing management of a ( Figure 1). We furthermore discovered the following four species to have significantly higher relative abundance at Moloa  (Figure 7). None of the latter four species displayed a relationship with the abiotic variables measured in this study (e.g., precipitation), so their distribution must be influenced by other factors. Their success at Moloa are known to respond to abiotic conditions, but the forces r insect pests are sometimes overlooked in the process of mbrosia beetles (Curculionidae: Scolytinae) are important worldwide, yet the factors that influence their success are aiian archipelago is host to over three dozen invasive uaʻi and are pests of agriculture. We analyzed scolytine tes: a hand-harvested site in a tropical wet forest and a cal dry savanna. Our regression analyses show overall elated with rainfall, as were four species: the tropical nut eus, Cryphalus longipilus, and Xyleborinus andrewesi. These ions of the communities being markedly dissimilar despite ivariate analysis found no influence from temperature or s. This information can be valuable for the timing of pest ess of possible new scolytine arrivals on Kauaʻi, and for d with climate change. ity; spatiotemporal distribution; phenology; Scolytinae; a could be attributed to the few non-coffee crops available there; for instance, X. compactus and X. crassiusculus are able to use cacao as a host [11,31]. Pérez-De La Cruz et al. [32] list cacao as the host for several species, including H. eruditus and Xyleborus affinis, the latter of which we found in Numila but not Moloa a. Moreover, these authors do not list cacao as a host for

Introduction
Abiotic factors influence seasonal variation of populations and thus the structure of their communities (e.g., diversity). Specific knowledge of these interactions is invaluable to understanding species geographical distributions and fluctuations of abundances for the purposes of ecological modeling, developing conservation strategies for threatened species, and directing management of pest species [1][2][3][4]. Many ambrosia and bark beetles (Curculionidae: Scolytinae) are important global pests of natural ecosystems, forestry, and agriculture, requiring tremendous financial and human resources to combat them [5][6][7][8][9]. Prominent examples include the mountain pine beetle, Dendroctonus ponderosae Hopkins, which has devastated millions of hectares of ecologically and commercially important pines in the Pacific Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus compactus (Eichhoff), which is a pest of hundreds of crops, ornamental plants, and forest trees [11]; and the coffee berry borer, Hypothenemus hampei (Ferrari), which is the most impactful pantropical pest of coffee production [12]. Much attention has been devoted to elucidating life histories of scolytines in the context of damage to host plants, and to developing monitoring and control i. Both were first detected in Numila in July 2016, and progressed to become, respectively, the third and fifth most numerous species of the 2016-2017 harvest season ( Figure 2 o respond to abiotic conditions, but the forces ts are sometimes overlooked in the process of tles (Curculionidae: Scolytinae) are important yet the factors that influence their success are elago is host to over three dozen invasive pests of agriculture. We analyzed scolytine -harvested site in a tropical wet forest and a nna. Our regression analyses show overall rainfall, as were four species: the tropical nut s longipilus, and Xyleborinus andrewesi. These mmunities being markedly dissimilar despite ysis found no influence from temperature or mation can be valuable for the timing of pest ble new scolytine arrivals on Kauaʻi, and for te change. mporal distribution; phenology; Scolytinae; populations and thus the structure of their ese interactions is invaluable to understanding f abundances for the purposes of ecological atened species, and directing management of urculionidae: Scolytinae) are important global , requiring tremendous financial and human nclude the mountain pine beetle, Dendroctonus of hectares of ecologically and commercially untains [10]; the black twig borer, Xylosandrus rops, ornamental plants, and forest trees [11]; ari), which is the most impactful pantropical been devoted to elucidating life histories of and to developing monitoring and control a Bay Coffee at the same time in our data, but unfortunately the shorter sampling period there precluded determining whether this cryphaline was present a priori. If H. brunneus was extant in Numila during the first year of sampling, it would have been at exceedingly low densities. This biodiversity study also extended the known distributions for several other species [15,24,35- communities are known to respond to abiotic conditions, but the forces n of particular insect pests are sometimes overlooked in the process of s. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important ecosystems worldwide, yet the factors that influence their success are es. The Hawaiian archipelago is host to over three dozen invasive occur on Kauaʻi and are pests of agriculture. We analyzed scolytine o coffee estates: a hand-harvested site in a tropical wet forest and a te in a tropical dry savanna. Our regression analyses show overall egatively correlated with rainfall, as were four species: the tropical nut us), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These the compositions of the communities being markedly dissimilar despite chness. Multivariate analysis found no influence from temperature or nity dynamics. This information can be valuable for the timing of pest cting the success of possible new scolytine arrivals on Kauaʻi, and for es may spread with climate change. ology; diversity; spatiotemporal distribution; phenology; Scolytinae; ce seasonal variation of populations and thus the structure of their . Specific knowledge of these interactions is invaluable to understanding utions and fluctuations of abundances for the purposes of ecological rvation strategies for threatened species, and directing management of rosia and bark beetles (Curculionidae: Scolytinae) are important global forestry, and agriculture, requiring tremendous financial and human 9]. Prominent examples include the mountain pine beetle, Dendroctonus has devastated millions of hectares of ecologically and commercially Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus is a pest of hundreds of crops, ornamental plants, and forest trees [11]; ypothenemus hampei (Ferrari), which is the most impactful pantropical 2]. Much attention has been devoted to elucidating life histories of damage to host plants, and to developing monitoring and control i, detailed alongside vouchers in S1.

Conclusions
On Kaua ond to abiotic conditions, but the forces sometimes overlooked in the process of urculionidae: Scolytinae) are important e factors that influence their success are is host to over three dozen invasive s of agriculture. We analyzed scolytine ested site in a tropical wet forest and a Our regression analyses show overall all, as were four species: the tropical nut gipilus, and Xyleborinus andrewesi. These nities being markedly dissimilar despite ound no influence from temperature or n can be valuable for the timing of pest ew scolytine arrivals on Kauaʻi, and for nge. ral distribution; phenology; Scolytinae; lations and thus the structure of their teractions is invaluable to understanding ndances for the purposes of ecological d species, and directing management of ionidae: Scolytinae) are important global iring tremendous financial and human e the mountain pine beetle, Dendroctonus ctares of ecologically and commercially ins [10]; the black twig borer, Xylosandrus ornamental plants, and forest trees [11]; which is the most impactful pantropical devoted to elucidating life histories of to developing monitoring and control i we found a unique opportunity to observe two sites that are similar in many dimensions (e.g., crops, soil, temperature, surrounding vegetation, scolytine density, scolytine richness) but differed in key factors (e.g., harvest method, precipitation, marginal crops). Our findings demonstrate that precipitation regime and species composition determine scolytine community dynamics; certain species have an inverse relationship with rainfall (such as those in Figure 6), which will dominate communities in drier environments/periods, but will be displaced by other species (such as those in Figure 7) in wetter environments/periods. Communities with such members will experience an overall increase in density in drier periods/environments (Figures 4 and 5). Marginal crops may be an additional factor in determining the community composition. Knowledge of these relationships could prove useful to management strategies, because pest management can be targeted in response to scolytine numbers, such as when they are at low densities, depending on the control method. These trends can also help predict the establishment of future invasive species; scolytines such as H. hampei that have an inverse relationship with wet climates [12] would be more successful in drier regions like Numila and less so in wetter ones like Moloa nown to respond to abiotic conditions, but the forces ct pests are sometimes overlooked in the process of sia beetles (Curculionidae: Scolytinae) are important wide, yet the factors that influence their success are archipelago is host to over three dozen invasive nd are pests of agriculture. We analyzed scolytine hand-harvested site in a tropical wet forest and a ry savanna. Our regression analyses show overall d with rainfall, as were four species: the tropical nut ryphalus longipilus, and Xyleborinus andrewesi. These f the communities being markedly dissimilar despite te analysis found no influence from temperature or is information can be valuable for the timing of pest f possible new scolytine arrivals on Kauaʻi, and for h climate change. Abstract: Populations and communities are known to respond to abiotic conditions, but the forces determining the distribution of particular insect pests are sometimes overlooked in the process of developing control methods. Bark and ambrosia beetles (Curculionidae: Scolytinae) are important pests of crops, forestry, and ecosystems worldwide, yet the factors that influence their success are unknown for many species. The Hawaiian archipelago is host to over three dozen invasive scolytines, many of which occur on Kauaʻi and are pests of agriculture. We analyzed scolytine community dynamics at two coffee estates: a hand-harvested site in a tropical wet forest and a mechanically harvested site in a tropical dry savanna. Our regression analyses show overall scolytine abundance was negatively correlated with rainfall, as were four species: the tropical nut borer (Hypothenemus obscurus), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These relationships contributed to the compositions of the communities being markedly dissimilar despite having the same species richness. Multivariate analysis found no influence from temperature or harvest method on community dynamics. This information can be valuable for the timing of pest control methods, for predicting the success of possible new scolytine arrivals on Kauaʻi, and for forecasting how these species may spread with climate change.

Introduction
i as climate change progresses [12,39]. Scolytines that have low or nonexistent densities in wetter climates may be able to expand into such areas as the climate changes.
Supplementary Materials: The following are available online at http://www.mdpi.com/2075-4450/9/4/123/s1. S1: Bark beetle vouchers with notes for new Kaua www.mdpi.com/journal/insects occur on Kauaʻi and are pests of agriculture. We analyzed scolytine o coffee estates: a hand-harvested site in a tropical wet forest and a te in a tropical dry savanna. Our regression analyses show overall egatively correlated with rainfall, as were four species: the tropical nut us), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These the compositions of the communities being markedly dissimilar despite chness. Multivariate analysis found no influence from temperature or nity dynamics. This information can be valuable for the timing of pest cting the success of possible new scolytine arrivals on Kauaʻi, and for es may spread with climate change. ology; diversity; spatiotemporal distribution; phenology; Scolytinae; e seasonal variation of populations and thus the structure of their . Specific knowledge of these interactions is invaluable to understanding utions and fluctuations of abundances for the purposes of ecological rvation strategies for threatened species, and directing management of rosia and bark beetles (Curculionidae: Scolytinae) are important global forestry, and agriculture, requiring tremendous financial and human 9]. Prominent examples include the mountain pine beetle, Dendroctonus has devastated millions of hectares of ecologically and commercially Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus is a pest of hundreds of crops, ornamental plants, and forest trees [11]; ypothenemus hampei (Ferrari), which is the most impactful pantropical 2]. Much attention has been devoted to elucidating life histories of damage to host plants, and to developing monitoring and control i adventive and naturalization records. Table S1: No statistical relationship between log-transformed monthly scolytine abundances and log-transformed monthly averages of temperature in bark beetle communities on Kaua www.mdpi.com/journal/insects s. The Hawaiian archipelago is host to over three dozen invasive occur on Kauaʻi and are pests of agriculture. We analyzed scolytine o coffee estates: a hand-harvested site in a tropical wet forest and a e in a tropical dry savanna. Our regression analyses show overall gatively correlated with rainfall, as were four species: the tropical nut us), H. brunneus, Cryphalus longipilus, and Xyleborinus andrewesi. These the compositions of the communities being markedly dissimilar despite chness. Multivariate analysis found no influence from temperature or nity dynamics. This information can be valuable for the timing of pest ting the success of possible new scolytine arrivals on Kauaʻi, and for es may spread with climate change.
ology; diversity; spatiotemporal distribution; phenology; Scolytinae; e seasonal variation of populations and thus the structure of their Specific knowledge of these interactions is invaluable to understanding tions and fluctuations of abundances for the purposes of ecological vation strategies for threatened species, and directing management of rosia and bark beetles (Curculionidae: Scolytinae) are important global forestry, and agriculture, requiring tremendous financial and human 9]. Prominent examples include the mountain pine beetle, Dendroctonus as devastated millions of hectares of ecologically and commercially Northwest and Rocky Mountains [10]; the black twig borer, Xylosandrus s a pest of hundreds of crops, ornamental plants, and forest trees [11]; ypothenemus hampei (Ferrari), which is the most impactful pantropical 2]. Much attention has been devoted to elucidating life histories of damage to host plants, and to developing monitoring and control