Distribution, Hosts and Parasitoids of Monochamus galloprovincialis (Coleoptera: Cerambycidae) in Portugal Mainland
Distribuição, hospedeiros e parasitóides de Monochamus galloprovincialis em Portugal Continental
Distribution des hôtes et parasitoïdes de Monochamus galloprovincialis (Coleoptera: Cerambycidae) au Portugal continental
*Ricardo Petersen-Silva, **Pedro Naves, ***Paulo Godinho, ***Edmundo Sousa and ****Juli Pujade-Villar
*Research grant-holder E-mail: ricardo.petersen@iniav.pt
**Researcher FCT E-mail: pedro.naves@iniav.pt
***Senior Researcher, Instituto Nacional de Investigação Agrária e Veterinária, IP., Unidade Estratégica de Investigação de Sistemas Agrários e Florestais e Sanidade Vegetal. Quinta do Marquês, 2780-159 Oeiras, PORTUGAL. E-mail: paulo.godinho@iniav.pt, edmundo.sousa@iniav.pt
]]> ****Professor, Universitat de Barcelona, Facultat de Biologia, Departament de Biología Animal. Avda. Diagonal 643, 08028-Barcelona, SPAIN. E-mails: rnpetersen@netcabo.pt, jpujade@ub.edu
ABSTRACT
The geographic distribution of the pine sawyer (Monochamus galloprovincialis), vector of the pine wood nematode (Bursaphelenchus xylophilus), is presented for Portugal mainland, according to the UTM 10x10km square grid. Field surveys were conducted in an area of approximately 15% of the maritime pine (Pinus pinaster) forested area. Simultaneously, a literature review was made for the previous references on the insect in Portugal. The insect vector was found in 94% of the surveyed locations, suggesting a widespread distribution. A total of eight forest and ornamental conifers were sampled, and for the first time the pine sawyer was detected in Portugal in two of them, namely Pinus sylvestris and Pinus halepensis. The beetle was not found associated with Pinus pinea, Pinus radiata, Cupressus lusitanica, Larix decidua and Chamaecyparis lawsoniana. The parasitoid guild associated with the pine sawyerin Portugal was reviewed and updated, with new geographic locations for a total of seven species.
Key words: Cerambycidae, Pinus pinaster; Pine Wood Nematode, Cyanopterus flavator, geographic distribution
RESUMO
É apresentada e discutida a distribuição geográfica em Portugal Continental do cerambicídeo Monochamus galloprovincialis, vetor do nemátode da madeira do pinheiro (Bursaphelenchus xylophilus), baseada na grelha UTM 10x10km. Foram realizadas prospeções em aproximadamente 15% da área florestal de pinheiro-bravo (Pinus pinaster). Simultaneamente, foi efetuada uma extensa revisão bibliográfica sobre as referências existentes para este cerambicídeo em Portugal. Foi confirmada a sua presença em 94% dos locais estudados, sugerindo uma distribuição alargada e generalizada. Foram prospetadas um total de oito espécies florestais e ornamentais, tendo-se encontrado pela primeira vez em Portugal a associação do inseto vetor com outros hospedeiros, nomeadamente com Pinus sylvestris e Pinus halepensis. Não se detetou a presença do inseto em Pinus pinea, Pinus radiata, Cupressus lusitanica, Larix decidua e Chamaecyparis lawsoniana. Adicionalmente, foi também revista e atualizada a comunidade de parasitóides associada a este cerambicídeo, num total de sete espécies, para as quais são apresentadas novas localizações geográficas.
Palavras-chave: Cerambycidae, Pinus pinaster, Nemátode da Madeira do Pinheiro, Cyanopterus flavator, distribuição geográfica
]]> RÉSUMÉ
La répartition géographique du scieur du pin (Monochamus galloprovincialis), vecteur du nématode du bois de pin (Bursaphelenchus xylophilus), est présentée au niveau du territoire continental portugais selon le quadrillage UTM 10x10km. Sur le terrain les études ont été menées en 15% environ de la surface forestière de pin maritime (Pinus pinaster). Simultanément a été faite une révision bibliographique des références précédentes de l'insecte au Portugal. Sa présence a été confirmée en 94% des endroits étudiés, ce qui suggère une répartition généralisée. Un total de huit conifères forestières et ornementales ont été échantillonnés, et le scieur de pin a été détecté pour la première fois au Portugal sur deux d'entre eux, a savoir Pinus sylvestris et Pinus halepensis. Le coléoptère n'a pas été trouvé associé à Pinus pinea, Pinus radiata, Cupressus lusitanica, Larix decidua et Chamaecyparis lawsoniana. La guilde des parasitoïdes associés au scieur de pin au Portugal a été revue et mise à jour, avec de nouvelles zones géographiques pour un total de sept espèces.
Mots-clés: Cerambycidae, Pinus pinaster, Nématode du Bois de Pin, Cyanopterus flavator, répartition géographique
Introduction
One of the most important sanitary problems affecting the maritime pine (Pinus pinaster Aiton) is the pine wilt disease (PWD). The causal agent is the pine wood nematode (PWN) Bursaphelenchus xylophilus (Steiner and Buhrer) Nickle (Nematoda: Tylenchida: Aphelenchoididae), which was first reported in Portugal by MOTA et al. in 1999. To be dispersed the PWN requires a vector insect, being the most effective the beetles of the genus Monochamus Dejan (Coleoptera: Cerambycidae) (KOBAYASHI et al., 1984; LINIT 1988; KISHI 1995; NAVES et al., 2007). In Portugal, the sole vector of the PWN is the pine sawyer Monochamus galloprovincialis (Olivier) (SOUSA et al., 2001). This beetle was firstly reported in the end of the XIXth century by OLIVEIRA (1894), who collected specimens from the Leiria region which he erroneously identified as Monochamus sutor Linnaeus. The species remains absent in subsequent catalogues and collection during the XXth century (CORRÊA DE BARROS, 1916; NEVES, 1950), until it was collected by NEVES et al. (1978) in burned P. pinaster trees near Arganil, central Portugal.
The pine sawyer is widely distributed in Europe (except in the United Kingdom, Ireland and Cyprus), being common in the Mediterranean area, namely in Italy, France, Spain, and Portugal (FRANCARDI and PENNACCHIO, 1996; VIVES, 2000; BRUSTEL et al., 2002; KOUTROUMPA, 2007). Outside of Europe, M. galloprovincialis can also be found in North Africa, the Caucasus, Russia, China and Mongolia (HELLRIGL, 1971).
Due to the increasing importance of this species as vector of the PWN, several recent studies have detailed various aspects of its biology and ecology (e.g. KOUTROUMPA et al., 2008; NAVES et al., 2008; AKBULUT, 2009), the application of chemical substances to control it (SOUSA et al., 2013) and its natural enemies (NAVES et al., 2005; PETERSEN-SILVA et al., 2012). Despite the recently-acquired knowledge, the PWN has continued to spread and is now causing mortality on pine forests in central and northern Portugal (SOUSA et al., 2011), (Figure 1). Even though the human transport of infested wood is responsible for long-distance dispersal of the disease, the beetle's flight activity and dispersal is also important when considering the spreading of PWD at a local scale. A detailed knowledge on the distribution and hosts of the vector beetle is important to understand the establishment and incidence of wilt disease in the terrain, and is lacking for Portugal. In this paper we present the first mapping of the distribution and hosts of M. galloprovincialis in continental Portugal, and review the parasitoid guild with new geographic records. A detailed bibliographic review on M. galloprovincialis distribution in Portugal was conducted.
]]> Materials and methods
Geographic distribution and hosts of Monochamus galloprovincialis
Surveys in the terrain were conducted by the INIAV (Instituto Nacional de Investigação Agrária e Veterinária, Oeiras, Portugal) entomological team. The surveys were mostly directed to areas where P. pinaster is the dominant tree species (ICNF, 2010), recording the locations in the UTM 10x10 km coordinates system (Universal Transverse Mercator coordinate system) and on the NUTS II system (Nomenclature of Territorial Units for Statistics). Surveys braced approximately 15% of the P. pinaster forested area. The surveyed areas were chosen based on the current distribution of the zones affected by the pine wilt disease (Figure 1, Annex 1), and on the proximity with Spain (buffer zone). The distribution of M. galloprovincialis was assessed by the presence of adult insects or larval instars inside decaying or dead trees. The characteristic round emergence holes made by the adults also allowed identifying past presence of the pine sawyer, but wood dissection was necessary to distinguish from similar holes made by carpenter bees of the genus Xylocopa Latreille or by wood wasps of the genus Sirex Linnaeus.
Although the majority of the survey was carried on P. pinaster, other conifers were also sporadically sampled, namely Pinus halepensis Mill., Pinus pinea L., Pinus sylvestris L., Pinus radiata Don, Cupressus lusitanica Mill., Larix decidua Mill. and Chamaecyparis lawsoniana (A. Murraybis) Parl.
Parasitoid guild associated with Monochamus galloprovincialis
The parasitoid guild distribution was studied from 2002 to 2003 and from 2011 to 2014. Locations where the surveys were conducted were randomly chosen among the locations surveyed for M. galloprovincialis distribution (Table 1, Annex 2). Field methods were based in the felling of trees with M. galloprovincialis presence and the creation of trap trees in the field. Collected wood was transported and reared at the INIAV laboratory, as described in NAVES el al. (2005) and PETERSEN-SILVA et al. (2012). The identification of parasitoid specimens was confirmed by Dr. Sergey Belokobylskij (Polish Academy of Sciences, Warszawa, Poland) and Dr. Dmitri Kasparyan (St. Petersburg, Russia).
Distribution data
Maps were created with the ESRI® ArcMapTM 10.0 program, and the presence/absence of M. galloprovincialis and the parasitoid guild was recorded in the UTM square grid (10 x 10 km) for continental Portugal. New records are signaled in the tables.
]]>Results
Geographic distribution and hosts of Monochamus galloprovincialis
A total of 96 UTM squares with dominant forest cover of Pinus pinaster were surveyed over the years (Table 1, Figure 2, and Annex 1). The pine sawyer M. galloprovincialis was found in 90 of them (93.8%), being widely distributed in the Portuguese continental territory. This cerambycidae showed an obvious preference for the maritime pine, although it was also found associated with other hosts such as P. sylvestris in Viana do Castelo (UTM NG21), and Pinus halepensis in Cascais (UTM MC68). All of these are new host records for Portugal. The pine sawyer was not detected in other surveyed conifers, such as Pinus pinea, Cupressus lusitanica, Larix decidua and Chamaecyparis lawsoniana.
Parasitoid guild associated with Monochamus galloprovincialis
The surveys lead to the collection of 188 parasitoid specimens found associated with M. galloprovincialis larvae. Specimens belong to the families Ichneumonidae and Braconidae and include seven species, namely: Atanycolus denigrator (L.); Atanycolus ivanowi (Kokujev); Cyanopterus flavator (Fabricius); Coeloides sordidator Ratzeburg; Doryctes striatellus (Nees); Iphiaulax impostor (Scopoli) and Xorides depressus (Holmgren). Their distribution, percentage of species and the UTM grid reference can be found in Table 1, Figure 3, and Annex 2,of which most are new distribution localities for Portugal. Cyanopterus flavator is the more frequently detected species, being found in seven of the 15 locations analyzed.
]]> Conclusions/ Discussion
The pine sawyer was found to be widely distributed on most of the Portuguese territory where pine forests are abundant, not being conditioned by edapho-climatic variations of the different localities, which was to be expected considering its vast distribution range (HELLRIGL, 1971). The absence of this widespread and conspicuous species from the previous catalogues of OLIVEIRA (1894), CORRÊA DE BARROS (1916), NEVES (1950) and SERRANO (1983) is somewhat intriguing, although these authors were not specifically looking for this beetle, unlike the surveys conducted by the INIAV team. It is also possible that the pine sawyer may have benefited from the great campaigns of forestation with P. pinaster that occurred in Portugal during the XXth century, experiencing a distribution expansion due to the higher availability of hosts. Insect populations may have also benefited from the sanitary problems and forest fires which occurred after this forest expansion, as the dead and scorcher pines tend to attract Monochamus beetles for breeding (BYERS, 1989a; b; SATO and MAETO, 2006; BONIFÁCIO et al., 2012).
Although maritime pine is clearly the preferred host in Portugal, the association of M. galloprovincialis with other hosts such as P. sylvestris and P. halepensisis is not surprising, as these are very frequent hosts all over the Mediterranean basin (HELLRIGL, 1971; EVANS et al., 1996; FRANCARDI and PENNACCHIO, 1996).Other unusual hosts include Pinus strobus (VINCENT, 2007), Pinus uncinata (BEHALOVÁ, 2006) and even Picea abies (L.) H. Karst.and Picea orientalis (L.) Peterm. (YUKSEL, 1998), although all of these species are inexistent or rare in Portugal.
Regarding the parasitoids associated with the insect vector, a fairly diverse guild was detected over the years, being dominated by Cyanopterus flavator, the most abundant species and with the widest distribution. Nevertheless and as already mentioned by NAVES et al. (2005) and PETERSEN-SILVA et al. (2012), these are all generalist species which attack a vast array of Cerambycidae and even other coleopteran, and therefore, their use as biological control agents seems ineffective and unlikely. Future samplings should focus on other natural enemies, such as bacteria or fungi, which have been found to affect other Monochamus species elsewhere, and have not yet been studied in detail in Portugal.
This is a first approach to elaborate a detailed mapping on the distribution range and hosts of M. galloprovincialis in Portugal, and may constitute an important tool for future assessments of pine wilt disease distribution and dispersal, and for the implementation of management and control strategies of this complex forest sanitary problem.
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Submitted for publication in April 2014
Accepted in May 2014
Acknowledgments
We would like to thank the colleagues of Instituto Nacional de Investigação Agrária e Veterinária, Mr. Adérito Matos, Dr. Luis Bonifácio and Mr. Victor Gonçalves for assistance in the field surveys and M. galloprovincialis laboratory handling, and Drª Lurdes Inácio for her comments and valuable text revision. We are also grateful to Dr. Sergey Belokobylskij of the Polish Academy of Sciences, Warszawa (Poland) and Dr. Dmitri Kasparyan (St. Petersburg, Russia) for confirming the identification of some parasitoid specimens. We would also like to thank Dr. José Manuel Rodrigues for the map presenting the current area affected by the pine wilt disease. This work was partially funded by the project REPHRAME Development of improved methods for detection, control and eradication of pine wood nematode in support of European Union Plant Health policy.
]]> Annexes
Annex 1
Distribution of Monochamus galloprovincialis in continental Portugal (UTM grid of 10 x 10km) and hosts
(a) – New host for M. galloprovincialis in Portugal; 1- Ribeiro, 1992; 2- Neves, et al., 1978; 3- Oliveira, 1894; Unknown – M. galloprovincialis adult specimens collected outside the host.
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