Overwintering Hosts for the Exotic Leafroller Parasitoid, Colpoclypeus florus: Implications for Habitat Manipulation to Augment Biological Control of Leafrollers in Pome Fruits

Thirty sites of managed and native habitats were surveyed for leafrollers (Lepidoptera: Tortricidae) in the apple producing region of central Washington State and northern Oregon from September through November 1997–2000 to discover species that supported overwintering by the parasitoid Colpoclypeus...

Ausführliche Beschreibung

Gespeichert in:
Bibliographische Detailangaben
Veröffentlicht in:Journal of insect science (Tucson, Ariz.) Ariz.), 2010, Vol.10 (75), p.1-13
Hauptverfasser: Pfannenstiel, R. S., Unruh, T. R., Brunner, J. F.
Format: Artikel
Sprache:eng
Schlagworte:
Online-Zugang:Volltext
Tags: Tag hinzufügen
Keine Tags, Fügen Sie den ersten Tag hinzu!
container_end_page 13
container_issue 75
container_start_page 1
container_title Journal of insect science (Tucson, Ariz.)
container_volume 10
creator Pfannenstiel, R. S.
Unruh, T. R.
Brunner, J. F.
description Thirty sites of managed and native habitats were surveyed for leafrollers (Lepidoptera: Tortricidae) in the apple producing region of central Washington State and northern Oregon from September through November 1997–2000 to discover species that supported overwintering by the parasitoid Colpoclypeus florus (Walker) (Hymenoptera: Eulophidae). C. florus, a species introduced from Europe, requires medium to large host larvae late in autumn on which to overwinter, and few leafroller species display this biology. Over the four years, five potential C. florus hosts were collected, including: Ancylis comptana (Froelich), Xenotemna pallorana (Robinson), and Syndemis sp. (Tortricidae), Filatima sp. (Gelechiidae), and Caloptilia burgessiellia (Zeller) (Gracillariidae). Of these, A. comptana, Syndemis sp., and Filatima sp. have been confirmed as overwintering hosts for C. florus. During the four years, the Syndemis sp. was rare and observed at only one location feeding on redosier dogwood, Cornus sericea L. (Cornales: Cornaceae) although, at this location, many of the larvae collected were parasitized by C. florus. Filatima sp. was common in the Yakima valley feeding on balsam poplar, Populus balsamifera L. ssp. trichocarpa (Torr. & Gray ex Hook) Brayshaw (Malpighiales: Salicaceae) but was rarely parasitized. A. comptana, however, was collected at many locations in central Washington and was frequently found as an overwintering host for C. florus. A. comptana was found feeding on two Rosaceae: Wood's rose, Rosa woodsii Lindl., and strawberry, Fragaria ananassa Duchesne (Rosales: Rosaceae). Based on the number of host larvae collected, A. comptana appears to be the primary overwintering host for C. florus in Washington. Introduction of A. comptana populations to near-orchard habitats may facilitate biological control of leafrollers that are orchard pests.
doi_str_mv 10.1673/031.010.7501
format Article
fullrecord <record><control><sourceid>gale_pubme</sourceid><recordid>TN_cdi_pubmedcentral_primary_oai_pubmedcentral_nih_gov_3383417</recordid><sourceformat>XML</sourceformat><sourcesystem>PC</sourcesystem><galeid>A248733900</galeid><oup_id>10.1673/031.010.7501</oup_id><sourcerecordid>A248733900</sourcerecordid><originalsourceid>FETCH-LOGICAL-b587t-a06e147f226d459b406f7d977943f492ef35378ff11556ca4053c1dfe743933d3</originalsourceid><addsrcrecordid>eNqFkktvEzEUhUcIREthxxp5h0BN8GvGmS6QQtSSSEGteKwtZ8aeGnl8B9tT2n_FT8QlpSQbKi_8-s65fpyieEnwlFSCvcOMTHGeiBKTR8UhKVk1oZzTxzvjg-JZjN8xppjP6qfFAcVZSWp2WPw6v9Lhp_VJB-s7tISYIjIQULrU6PQakm3QWisTwDkd0IUKKtoEtj1GC3ADNO5m0GOWOAhjPEGrfnC2UcmC3_os1cYmldAn5e0wuj87KAGaj12vfUIfLDjossRlQ59yHQRmp2RE1qML6DU6C6NN8XnxxCgX9Yu7_qj4dnb6dbGcrM8_rhbz9WRTzkSaKFxpwoWhtGp5WW84roxoayFqzgyvqTasZGJmDCFlWTWK45I1pDVacFYz1rKj4v3Wdxg3vW6bfNagnByC7VW4kaCs3N_x9lJ2cCUZmzFORDZ4fWcQ4MeoY5K9jY12TnkNY5R1yatcnNCHSZI5XNHqQVLk76WVKOtMTrdkp5yW1hvIh2xya3VvG_Da2Lw-p3wmGKsxzoI3e4LMJH2dOjXGKFdfPu-zx1u2CRBj0Ob-WQiWt6GUOZQyh1LehjLjr3af8h7-m8J_14JxeMjq7ZbcWMiX-D_8Gwn39xY</addsrcrecordid><sourcetype>Open Access Repository</sourcetype><iscdi>true</iscdi><recordtype>article</recordtype><pqid>748926759</pqid></control><display><type>article</type><title>Overwintering Hosts for the Exotic Leafroller Parasitoid, Colpoclypeus florus: Implications for Habitat Manipulation to Augment Biological Control of Leafrollers in Pome Fruits</title><source>Oxford Journals Open Access Collection</source><source>MEDLINE</source><source>DOAJ Directory of Open Access Journals</source><source>BioOne Open Access Titles</source><source>Elektronische Zeitschriftenbibliothek - Frei zugängliche E-Journals</source><source>PubMed Central</source><creator>Pfannenstiel, R. S. ; Unruh, T. R. ; Brunner, J. F.</creator><creatorcontrib>Pfannenstiel, R. S. ; Unruh, T. R. ; Brunner, J. F.</creatorcontrib><description>Thirty sites of managed and native habitats were surveyed for leafrollers (Lepidoptera: Tortricidae) in the apple producing region of central Washington State and northern Oregon from September through November 1997–2000 to discover species that supported overwintering by the parasitoid Colpoclypeus florus (Walker) (Hymenoptera: Eulophidae). C. florus, a species introduced from Europe, requires medium to large host larvae late in autumn on which to overwinter, and few leafroller species display this biology. Over the four years, five potential C. florus hosts were collected, including: Ancylis comptana (Froelich), Xenotemna pallorana (Robinson), and Syndemis sp. (Tortricidae), Filatima sp. (Gelechiidae), and Caloptilia burgessiellia (Zeller) (Gracillariidae). Of these, A. comptana, Syndemis sp., and Filatima sp. have been confirmed as overwintering hosts for C. florus. During the four years, the Syndemis sp. was rare and observed at only one location feeding on redosier dogwood, Cornus sericea L. (Cornales: Cornaceae) although, at this location, many of the larvae collected were parasitized by C. florus. Filatima sp. was common in the Yakima valley feeding on balsam poplar, Populus balsamifera L. ssp. trichocarpa (Torr. &amp; Gray ex Hook) Brayshaw (Malpighiales: Salicaceae) but was rarely parasitized. A. comptana, however, was collected at many locations in central Washington and was frequently found as an overwintering host for C. florus. A. comptana was found feeding on two Rosaceae: Wood's rose, Rosa woodsii Lindl., and strawberry, Fragaria ananassa Duchesne (Rosales: Rosaceae). Based on the number of host larvae collected, A. comptana appears to be the primary overwintering host for C. florus in Washington. Introduction of A. comptana populations to near-orchard habitats may facilitate biological control of leafrollers that are orchard pests.</description><identifier>ISSN: 1536-2442</identifier><identifier>EISSN: 1536-2442</identifier><identifier>DOI: 10.1673/031.010.7501</identifier><identifier>PMID: 20673193</identifier><language>eng</language><publisher>Oxford, UK: University of Wisconsin Library</publisher><subject>alternate host ; Ancylis comptana ; Animals ; Bioclimatology ; Biological control ; Caloptilia ; Cornaceae ; Cornales ; Cornus ; diapause ; Ecosystem ; Environmental aspects ; Eulophidae ; Fragaria ; Fragaria ananassa ; Fruit - parasitology ; Gelechiidae ; Gracillariidae ; Host-Parasite Interactions ; Hymenoptera ; Lepidoptera ; Malus ; Moths - parasitology ; Parasitoids ; Pest Control, Biological ; Physiological aspects ; Populus balsamifera ; Rosa woodsii ; Rosaceae ; Rosales ; Salicaceae ; Seasons ; Tortricidae ; Tortricids ; Wasps - physiology ; Winter</subject><ispartof>Journal of insect science (Tucson, Ariz.), 2010, Vol.10 (75), p.1-13</ispartof><rights>This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits unrestricted use, provided that the paper is properly attributed.</rights><rights>This is an open access paper. We use the Creative Commons Attribution 3.0 license that permits unrestricted use, provided that the paper is properly attributed. 2010</rights><rights>COPYRIGHT 2010 Oxford University Press</rights><rights>2010 2010</rights><lds50>peer_reviewed</lds50><oa>free_for_read</oa><woscitedreferencessubscribed>false</woscitedreferencessubscribed><citedby>FETCH-LOGICAL-b587t-a06e147f226d459b406f7d977943f492ef35378ff11556ca4053c1dfe743933d3</citedby><cites>FETCH-LOGICAL-b587t-a06e147f226d459b406f7d977943f492ef35378ff11556ca4053c1dfe743933d3</cites></display><links><openurl>$$Topenurl_article</openurl><openurlfulltext>$$Topenurlfull_article</openurlfulltext><thumbnail>$$Tsyndetics_thumb_exl</thumbnail><linktopdf>$$Uhttps://bioone.org/doi/pdf/10.1673/031.010.7501$$EPDF$$P50$$Gbioone$$H</linktopdf><linktohtml>$$Uhttps://www.ncbi.nlm.nih.gov/pmc/articles/PMC3383417/$$EHTML$$P50$$Gpubmedcentral$$Hfree_for_read</linktohtml><link.rule.ids>109,230,314,723,776,780,860,881,4010,27900,27901,27902,52694,53766,53768</link.rule.ids><backlink>$$Uhttps://www.ncbi.nlm.nih.gov/pubmed/20673193$$D View this record in MEDLINE/PubMed$$Hfree_for_read</backlink></links><search><creatorcontrib>Pfannenstiel, R. S.</creatorcontrib><creatorcontrib>Unruh, T. R.</creatorcontrib><creatorcontrib>Brunner, J. F.</creatorcontrib><title>Overwintering Hosts for the Exotic Leafroller Parasitoid, Colpoclypeus florus: Implications for Habitat Manipulation to Augment Biological Control of Leafrollers in Pome Fruits</title><title>Journal of insect science (Tucson, Ariz.)</title><addtitle>J Insect Sci</addtitle><description>Thirty sites of managed and native habitats were surveyed for leafrollers (Lepidoptera: Tortricidae) in the apple producing region of central Washington State and northern Oregon from September through November 1997–2000 to discover species that supported overwintering by the parasitoid Colpoclypeus florus (Walker) (Hymenoptera: Eulophidae). C. florus, a species introduced from Europe, requires medium to large host larvae late in autumn on which to overwinter, and few leafroller species display this biology. Over the four years, five potential C. florus hosts were collected, including: Ancylis comptana (Froelich), Xenotemna pallorana (Robinson), and Syndemis sp. (Tortricidae), Filatima sp. (Gelechiidae), and Caloptilia burgessiellia (Zeller) (Gracillariidae). Of these, A. comptana, Syndemis sp., and Filatima sp. have been confirmed as overwintering hosts for C. florus. During the four years, the Syndemis sp. was rare and observed at only one location feeding on redosier dogwood, Cornus sericea L. (Cornales: Cornaceae) although, at this location, many of the larvae collected were parasitized by C. florus. Filatima sp. was common in the Yakima valley feeding on balsam poplar, Populus balsamifera L. ssp. trichocarpa (Torr. &amp; Gray ex Hook) Brayshaw (Malpighiales: Salicaceae) but was rarely parasitized. A. comptana, however, was collected at many locations in central Washington and was frequently found as an overwintering host for C. florus. A. comptana was found feeding on two Rosaceae: Wood's rose, Rosa woodsii Lindl., and strawberry, Fragaria ananassa Duchesne (Rosales: Rosaceae). Based on the number of host larvae collected, A. comptana appears to be the primary overwintering host for C. florus in Washington. Introduction of A. comptana populations to near-orchard habitats may facilitate biological control of leafrollers that are orchard pests.</description><subject>alternate host</subject><subject>Ancylis comptana</subject><subject>Animals</subject><subject>Bioclimatology</subject><subject>Biological control</subject><subject>Caloptilia</subject><subject>Cornaceae</subject><subject>Cornales</subject><subject>Cornus</subject><subject>diapause</subject><subject>Ecosystem</subject><subject>Environmental aspects</subject><subject>Eulophidae</subject><subject>Fragaria</subject><subject>Fragaria ananassa</subject><subject>Fruit - parasitology</subject><subject>Gelechiidae</subject><subject>Gracillariidae</subject><subject>Host-Parasite Interactions</subject><subject>Hymenoptera</subject><subject>Lepidoptera</subject><subject>Malus</subject><subject>Moths - parasitology</subject><subject>Parasitoids</subject><subject>Pest Control, Biological</subject><subject>Physiological aspects</subject><subject>Populus balsamifera</subject><subject>Rosa woodsii</subject><subject>Rosaceae</subject><subject>Rosales</subject><subject>Salicaceae</subject><subject>Seasons</subject><subject>Tortricidae</subject><subject>Tortricids</subject><subject>Wasps - physiology</subject><subject>Winter</subject><issn>1536-2442</issn><issn>1536-2442</issn><fulltext>true</fulltext><rsrctype>article</rsrctype><creationdate>2010</creationdate><recordtype>article</recordtype><sourceid>TOX</sourceid><sourceid>EIF</sourceid><recordid>eNqFkktvEzEUhUcIREthxxp5h0BN8GvGmS6QQtSSSEGteKwtZ8aeGnl8B9tT2n_FT8QlpSQbKi_8-s65fpyieEnwlFSCvcOMTHGeiBKTR8UhKVk1oZzTxzvjg-JZjN8xppjP6qfFAcVZSWp2WPw6v9Lhp_VJB-s7tISYIjIQULrU6PQakm3QWisTwDkd0IUKKtoEtj1GC3ADNO5m0GOWOAhjPEGrfnC2UcmC3_os1cYmldAn5e0wuj87KAGaj12vfUIfLDjossRlQ59yHQRmp2RE1qML6DU6C6NN8XnxxCgX9Yu7_qj4dnb6dbGcrM8_rhbz9WRTzkSaKFxpwoWhtGp5WW84roxoayFqzgyvqTasZGJmDCFlWTWK45I1pDVacFYz1rKj4v3Wdxg3vW6bfNagnByC7VW4kaCs3N_x9lJ2cCUZmzFORDZ4fWcQ4MeoY5K9jY12TnkNY5R1yatcnNCHSZI5XNHqQVLk76WVKOtMTrdkp5yW1hvIh2xya3VvG_Da2Lw-p3wmGKsxzoI3e4LMJH2dOjXGKFdfPu-zx1u2CRBj0Ob-WQiWt6GUOZQyh1LehjLjr3af8h7-m8J_14JxeMjq7ZbcWMiX-D_8Gwn39xY</recordid><startdate>2010</startdate><enddate>2010</enddate><creator>Pfannenstiel, R. S.</creator><creator>Unruh, T. R.</creator><creator>Brunner, J. F.</creator><general>University of Wisconsin Library</general><general>Oxford University Press</general><scope>TOX</scope><scope>CGR</scope><scope>CUY</scope><scope>CVF</scope><scope>ECM</scope><scope>EIF</scope><scope>NPM</scope><scope>AAYXX</scope><scope>CITATION</scope><scope>ISR</scope><scope>7X8</scope><scope>7QG</scope><scope>7SN</scope><scope>7SS</scope><scope>C1K</scope><scope>7T7</scope><scope>8FD</scope><scope>FR3</scope><scope>P64</scope><scope>5PM</scope></search><sort><creationdate>2010</creationdate><title>Overwintering Hosts for the Exotic Leafroller Parasitoid, Colpoclypeus florus: Implications for Habitat Manipulation to Augment Biological Control of Leafrollers in Pome Fruits</title><author>Pfannenstiel, R. S. ; Unruh, T. R. ; Brunner, J. F.</author></sort><facets><frbrtype>5</frbrtype><frbrgroupid>cdi_FETCH-LOGICAL-b587t-a06e147f226d459b406f7d977943f492ef35378ff11556ca4053c1dfe743933d3</frbrgroupid><rsrctype>articles</rsrctype><prefilter>articles</prefilter><language>eng</language><creationdate>2010</creationdate><topic>alternate host</topic><topic>Ancylis comptana</topic><topic>Animals</topic><topic>Bioclimatology</topic><topic>Biological control</topic><topic>Caloptilia</topic><topic>Cornaceae</topic><topic>Cornales</topic><topic>Cornus</topic><topic>diapause</topic><topic>Ecosystem</topic><topic>Environmental aspects</topic><topic>Eulophidae</topic><topic>Fragaria</topic><topic>Fragaria ananassa</topic><topic>Fruit - parasitology</topic><topic>Gelechiidae</topic><topic>Gracillariidae</topic><topic>Host-Parasite Interactions</topic><topic>Hymenoptera</topic><topic>Lepidoptera</topic><topic>Malus</topic><topic>Moths - parasitology</topic><topic>Parasitoids</topic><topic>Pest Control, Biological</topic><topic>Physiological aspects</topic><topic>Populus balsamifera</topic><topic>Rosa woodsii</topic><topic>Rosaceae</topic><topic>Rosales</topic><topic>Salicaceae</topic><topic>Seasons</topic><topic>Tortricidae</topic><topic>Tortricids</topic><topic>Wasps - physiology</topic><topic>Winter</topic><toplevel>peer_reviewed</toplevel><toplevel>online_resources</toplevel><creatorcontrib>Pfannenstiel, R. S.</creatorcontrib><creatorcontrib>Unruh, T. R.</creatorcontrib><creatorcontrib>Brunner, J. F.</creatorcontrib><collection>Oxford Journals Open Access Collection</collection><collection>Medline</collection><collection>MEDLINE</collection><collection>MEDLINE (Ovid)</collection><collection>MEDLINE</collection><collection>MEDLINE</collection><collection>PubMed</collection><collection>CrossRef</collection><collection>Gale In Context: Science</collection><collection>MEDLINE - Academic</collection><collection>Animal Behavior Abstracts</collection><collection>Ecology Abstracts</collection><collection>Entomology Abstracts (Full archive)</collection><collection>Environmental Sciences and Pollution Management</collection><collection>Industrial and Applied Microbiology Abstracts (Microbiology A)</collection><collection>Technology Research Database</collection><collection>Engineering Research Database</collection><collection>Biotechnology and BioEngineering Abstracts</collection><collection>PubMed Central (Full Participant titles)</collection><jtitle>Journal of insect science (Tucson, Ariz.)</jtitle></facets><delivery><delcategory>Remote Search Resource</delcategory><fulltext>fulltext</fulltext></delivery><addata><au>Pfannenstiel, R. S.</au><au>Unruh, T. R.</au><au>Brunner, J. F.</au><format>journal</format><genre>article</genre><ristype>JOUR</ristype><atitle>Overwintering Hosts for the Exotic Leafroller Parasitoid, Colpoclypeus florus: Implications for Habitat Manipulation to Augment Biological Control of Leafrollers in Pome Fruits</atitle><jtitle>Journal of insect science (Tucson, Ariz.)</jtitle><addtitle>J Insect Sci</addtitle><date>2010</date><risdate>2010</risdate><volume>10</volume><issue>75</issue><spage>1</spage><epage>13</epage><pages>1-13</pages><issn>1536-2442</issn><eissn>1536-2442</eissn><abstract>Thirty sites of managed and native habitats were surveyed for leafrollers (Lepidoptera: Tortricidae) in the apple producing region of central Washington State and northern Oregon from September through November 1997–2000 to discover species that supported overwintering by the parasitoid Colpoclypeus florus (Walker) (Hymenoptera: Eulophidae). C. florus, a species introduced from Europe, requires medium to large host larvae late in autumn on which to overwinter, and few leafroller species display this biology. Over the four years, five potential C. florus hosts were collected, including: Ancylis comptana (Froelich), Xenotemna pallorana (Robinson), and Syndemis sp. (Tortricidae), Filatima sp. (Gelechiidae), and Caloptilia burgessiellia (Zeller) (Gracillariidae). Of these, A. comptana, Syndemis sp., and Filatima sp. have been confirmed as overwintering hosts for C. florus. During the four years, the Syndemis sp. was rare and observed at only one location feeding on redosier dogwood, Cornus sericea L. (Cornales: Cornaceae) although, at this location, many of the larvae collected were parasitized by C. florus. Filatima sp. was common in the Yakima valley feeding on balsam poplar, Populus balsamifera L. ssp. trichocarpa (Torr. &amp; Gray ex Hook) Brayshaw (Malpighiales: Salicaceae) but was rarely parasitized. A. comptana, however, was collected at many locations in central Washington and was frequently found as an overwintering host for C. florus. A. comptana was found feeding on two Rosaceae: Wood's rose, Rosa woodsii Lindl., and strawberry, Fragaria ananassa Duchesne (Rosales: Rosaceae). Based on the number of host larvae collected, A. comptana appears to be the primary overwintering host for C. florus in Washington. Introduction of A. comptana populations to near-orchard habitats may facilitate biological control of leafrollers that are orchard pests.</abstract><cop>Oxford, UK</cop><pub>University of Wisconsin Library</pub><pmid>20673193</pmid><doi>10.1673/031.010.7501</doi><tpages>13</tpages><oa>free_for_read</oa></addata></record>
fulltext fulltext
identifier ISSN: 1536-2442
ispartof Journal of insect science (Tucson, Ariz.), 2010, Vol.10 (75), p.1-13
issn 1536-2442
1536-2442
language eng
recordid cdi_pubmedcentral_primary_oai_pubmedcentral_nih_gov_3383417
source Oxford Journals Open Access Collection; MEDLINE; DOAJ Directory of Open Access Journals; BioOne Open Access Titles; Elektronische Zeitschriftenbibliothek - Frei zugängliche E-Journals; PubMed Central
subjects alternate host
Ancylis comptana
Animals
Bioclimatology
Biological control
Caloptilia
Cornaceae
Cornales
Cornus
diapause
Ecosystem
Environmental aspects
Eulophidae
Fragaria
Fragaria ananassa
Fruit - parasitology
Gelechiidae
Gracillariidae
Host-Parasite Interactions
Hymenoptera
Lepidoptera
Malus
Moths - parasitology
Parasitoids
Pest Control, Biological
Physiological aspects
Populus balsamifera
Rosa woodsii
Rosaceae
Rosales
Salicaceae
Seasons
Tortricidae
Tortricids
Wasps - physiology
Winter
title Overwintering Hosts for the Exotic Leafroller Parasitoid, Colpoclypeus florus: Implications for Habitat Manipulation to Augment Biological Control of Leafrollers in Pome Fruits
url https://sfx.bib-bvb.de/sfx_tum?ctx_ver=Z39.88-2004&ctx_enc=info:ofi/enc:UTF-8&ctx_tim=2025-02-14T00%3A55%3A04IST&url_ver=Z39.88-2004&url_ctx_fmt=infofi/fmt:kev:mtx:ctx&rfr_id=info:sid/primo.exlibrisgroup.com:primo3-Article-gale_pubme&rft_val_fmt=info:ofi/fmt:kev:mtx:journal&rft.genre=article&rft.atitle=Overwintering%20Hosts%20for%20the%20Exotic%20Leafroller%20Parasitoid,%20Colpoclypeus%20florus:%20Implications%20for%20Habitat%20Manipulation%20to%20Augment%20Biological%20Control%20of%20Leafrollers%20in%20Pome%20Fruits&rft.jtitle=Journal%20of%20insect%20science%20(Tucson,%20Ariz.)&rft.au=Pfannenstiel,%20R.%20S.&rft.date=2010&rft.volume=10&rft.issue=75&rft.spage=1&rft.epage=13&rft.pages=1-13&rft.issn=1536-2442&rft.eissn=1536-2442&rft_id=info:doi/10.1673/031.010.7501&rft_dat=%3Cgale_pubme%3EA248733900%3C/gale_pubme%3E%3Curl%3E%3C/url%3E&disable_directlink=true&sfx.directlink=off&sfx.report_link=0&rft_id=info:oai/&rft_pqid=748926759&rft_id=info:pmid/20673193&rft_galeid=A248733900&rft_oup_id=10.1673/031.010.7501&rfr_iscdi=true