Biodiversity Data Journal :
Taxonomic Paper
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Corresponding author: Levente-Péter Kolcsár (kolcsar.peter@gmail.com)
Academic editor: Torsten Dikow
Received: 07 Apr 2021 | Accepted: 14 Jun 2021 | Published: 21 Jul 2021
© 2021 Levente-Péter Kolcsár, Pjotr Oosterbroek, Dmitry Gavryushin, Kjell Magne Olsen, Nikolai Paramonov, Valentin Pilipenko, Jaroslav Starý, Alexei Polevoi, Vladimir Lantsov, Eulalia Eiroa, Michael Andersson, Jukka Salmela, Clovis Quindroit, Micha d'Oliveira, E. Geoffrey Hancock, Jorge Mederos, Pete Boardman, Esko Viitanen, Kozo Watanabe
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Kolcsár L-P, Oosterbroek P, Gavryushin DI, Olsen KM, Paramonov NM, Pilipenko VE, Starý J, Polevoi A, Lantsov VI, Eiroa E, Andersson M, Salmela J, Quindroit C, d'Oliveira MC, Hancock EG, Mederos J, Boardman P, Viitanen E, Watanabe K (2021) Contribution to the knowledge of Limoniidae (Diptera: Tipuloidea): first records of 244 species from various European countries. Biodiversity Data Journal 9: e67085. https://doi.org/10.3897/BDJ.9.e67085
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Limoniidae is one of the most species-rich Dipteran families, with 661 reported species in Europe. Despite the fact that the European limoniid crane fly fauna has been studied ever since Carolus Linnaeus, it is still poorly known.
In this study, we summarise the taxonomic and faunistic studies of European Limoniidae, which described new species and reported first country records, between 2010 and 2020. We also report occurrence data of 244 Limoniidae species which represent the first country records or conformational records from various European countries, as we report ten species from Albania, one from Austria, thirty-seven from Belarus, five from Belgium, two from Bulgaria, two from Estonia, six from Finland, seven from France, fourteen from Greece, sixteen from Hungary, two from Iceland, six from Italy, ten from Latvia, one from Malta, nine from Montenegro, two from The Netherlands, ten from North Macedonia, forty-two from Norway, one from Poland, five from Portugal, twenty from Romania, thirty-eight from Serbia, six from Slovenia, five from Spain and seven species from Sweden for the first time. From the European territory of Russia, we report twenty-eight species from Central European Russia, seventy-two from East European Russia, fifteen from North European Russia, one from Northwest European Russia and seven from North Caucasus for the first time. Confirmatory records and corrigenda are also included.
biodiversity, Catalogue of the Craneflies of the World (CCW), collection data, database, Global Biodiversity Information Facility (GBIF), fauna, terminalia photos
Crane flies (superfamily Tipuloidea) are one of the most species-rich dipteran groups, with more than 15600 recognised taxa (
Since the description of the first limoniid crane fly species by C. Linnaeus in 1758, the number of described/reported species from Europe has continuously increased (Fig.
Species | Publication | Distribution |
Austrolimnophila (Austrolimnophila) cretica Starý, 2014 |
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Greece (Crete) |
Austrolimnophila (Austrolimnophila) vikhrevi Gavryushin, 2015 |
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Azerbajian, Georgia and Russia: North Caucasus |
Baeoura rotherayi Hancock, 2020 |
|
Spain (Jaen) |
Chionea dolomitana Vanin, 2010 |
|
Italy (Belluno, Treviso, Udine) |
Chionea olympiae Vanin, 2010 |
|
Italy (Biella) |
Dicranomyia (Idiopyga) boreobaltica Salmela, 2014 |
|
Finland (northern Baltic area) |
Dicranophragma relictum Mederos, 2020 |
|
Spain (Barcelona) |
Ellipteroides (Ramagonomyia) mendli Hancock and Starý, 2019 |
|
Portugal (Faro) and Spain (south) |
Erioptera (Erioptera) octobris Gavryushin, 2011 |
|
Russia: North Caucasus |
Geranomyia eugeniana Lantsov, 2015 |
|
Azerbajian, Georgia and Russia: North Caucasus |
Geranomyia fuscior Starý, 2012 |
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Albania (Durazzo), Portugal (Faro) and Libya (Tripolis) |
Gonempeda mariya Pilipenko, 2017 |
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Russia: North Caucasus |
Gonomyia (Gonomyia) securiformis Starý, 2011 |
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Bulgaria, Czech Rep., Italy (Piemonte) and Switzerland |
Gonomyia (Gonomyia) undiformis Starý, 2011 |
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Slovakia |
Idiocera (Idiocera) cretopunctata Starý, 2020 |
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Greece (Crete) |
Idiocera (Idiocera) falcistylus Starý, 2020 |
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Spain (Almeria) |
Limonia enormis Starý, 2017 |
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Greece (Crete) |
Limonia hartveldae Starý, 2017 |
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Portugal (Guarda) |
Lipsothrix galiciensis Hancock and Hewitt, 2015 |
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Spain (Lugo) |
Molophilus (Molophilus) balcanicus Kolcsár, 2015 |
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Bulgaria (Stara Planina Mts) |
Molophilus (Molophilus) brevifurcatus Starý, 2011 |
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Czech Rep. and Slovakia |
Molophilus (Molophilus) calabricus Starý, 2011 |
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Italy (Calabria) |
Molophilus (Molophilus) carbonis Starý, 2011 |
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Portugal (Faro) |
Molophilus (Molophilus) creticola Starý, 2011 |
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Greece (Crete) |
Molophilus (Molophilus) cypricola Starý, 2011 |
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Cyprus |
Molophilus (Molophilus) ibericus Starý, 2011 |
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Spain and Morocco (High Atlas) |
Molophilus (Molophilus) rohaceki Starý and Oboňa, 2019 |
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Greece (Taygetos Mts) |
Molophilus (Molophilus) suboccultus Starý, 2011 |
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Portugal (Faro) |
Molophilus (Molophilus) zonzensis Boardman and Starý, 2020 |
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France (Corsica) |
Prionolabis pjotri Hancock, 2015 |
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Portugal and Spain (Lugo) |
Thaumastoptera (Thaumastoptera) spathifera Starý and Obaňa, 2018 |
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Italy (Sicily) |
In the last decade, many new faunistic studies have been published on European Limoniidae, of which we list the ones that report at least one species for the first time from a country (Table
Summarised list of publications reporting new country records from Europe during the period 2010–2020, including those that describe new species. The total number includes questionable and unconfirmed species.
Country | Publications with new country record(s) during the period 2010-2020 | No. of species reported as new during the period 2010-2020 | No. of species reported as new in this article | Total no. of species including the new records |
Albania |
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1 | 10 | 71 |
Andorra |
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12 | 0 | 28 |
Austria |
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8 | 1 | 277 |
Belarus |
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14 | 37 | 63 |
Belgium |
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1 | 5 | 165 |
Bosnia and Herzegovina |
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2 | 0 | 71 |
Bulgaria |
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7 | 2 | 222 |
Croatia |
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14 | 0 | 87 |
Cyprus |
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1 | 0 | 20 |
Czech Republic (Czechia) |
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6 | 0 | 300 |
Denmark |
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11 | 0 | 171 |
Estonia | 0 | 2 | 82 | |
Finland |
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8 | 6 | 204 |
France |
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18 | 7 | 279 |
Germany |
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1 | 0 | 292 |
Greece |
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14 | 14 | 116 |
Hungary |
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30 | 16 | 177 |
Iceland | 0 | 2 | 13 | |
Ireland | 0 | 0 | 131 | |
Italy |
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9 | 6 | 280 |
Latvia | 0 | 11 | 113 | |
Liechtenstein | 0 | 0 | 0 | |
Lithuania |
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1 | 0 | 202 |
Luxembourg |
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4 | 0 | 35 |
Malta |
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10 | 1 | 12 |
Moldova | 0 | 0 | 18 | |
Monaco | 0 | 0 | 0 | |
Montenegro |
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2 | 9 | 70 |
Netherlands |
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7 | 2 | 153 |
North Macedonia |
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1 | 10 | 93 |
Norway |
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40 | 42 | 216 |
Poland |
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13 | 1 | 232 |
Portugal |
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55 | 5 | 85 |
Romania |
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9 | 20 | 285 |
Russia: Central European Russia (RUC) |
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62 | 28 | 141 |
Russia: East European Russia (RUE) |
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11 | 72 | 103 |
Russia: North European Russia (RUN) |
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5 | 15 | 149 |
Russia: Northwest European Russia (RUW) | 0 | 1 | 141 | |
Russia: South European Russia (RUS) |
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7 | 0 | 65 |
Russia: North Caucasus (NC) |
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15 | 7 | 154 |
San Marino | 0 | 0 | 0 | |
Serbia |
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3 | 38 | 109 |
Slovakia |
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5 | 0 | 320 |
Slovenia | 0 | 6 | 113 | |
Spain |
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43 | 5 | 181 |
Sweden |
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16 | 7 | 223 |
Switzerland |
|
2 | 0 | 298 |
Ukraine |
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2 | 0 | 243 |
United Kingdom |
|
5 | 0 |
228 |
Turkey (European part) |
|
65 | 0 |
67 |
The aim of this paper is to publish the first records of limonid crane flies from various European countries. A collection of previously unpublished scientific records from CCW uploaded between 2010 and 2020 is also included, which represent the first country record(s) of the species in question. Confirmatory records and some corrigenda are also included. In total, we present 859 records belonging to 244 species.
In this study, we focus on the territory of Europe (Fig.
Abbreviations used for European territories of Russia:
RUN – North European Russia
RUW – Northwest European Russia
RUC – Central European Russia
RUE – East European Russia
RUS – South European Russia
NC – North Caucasus
Studied material has been collected by recognised manual field methods, such as sweep or hand-netting or trapped by the use of Malaise traps, SLAM-traps, light traps, trunk emergence traps, Lindgren funnel traps, pitfall traps, pan traps and window traps. Some records are based on photos or field observations. The preserved material is pinned or stored in ethanol and deposited in various public or private collections.
About 99% of the material is identified by the authors (excluding K. Watanabe) and the identifier(s) are responsible for the corresponding records. Collection data also avaible on the GBIF.org database (
Photographs of wing, body and terminalia of some rare species were taken by the first author, by using a Zeiss Stemi 508 stereomicroscope, equipped with a Canon Kiss M digital camera or with Optik microscope, equipped with a LM Digital SLR Adapter and Canon 650D camera.
BioFokus – BioFokus' office (https://biofokus.no), Oslo, Norway.
CKLP – Private Collection of L.-P. Kolcsár.
FRIP – Forest Research Institute, Petrozavodsk, Russia.
HMUG – Hunterian Museum, University of Glasgow, Glasgow, United Kindom.
HNHM – Hungarian National History Museum, Budapest, Hungary.
IEMT – Tembotov Institute of Ecology of Mountain Territories of Russian Academy of Sciences, Nalchik, Russia.
LMM – Regional Museum of Lapland, Rovaniemi, Finland.
MCNB – Natural Sciences Museum of Barcelona, Barcelona, Spain.
MNHN – National Museum of Natural History, Paris, France.
MZLU – Museum of Biology of Lund, Lund, Sweden.
NBCN – Naturalis Biodiversity Center in Leiden, The Netherlands.
NHMO – Natural History Museum at the University of Oslo, Oslo, Norway.
NHRS – Entomological Collections, Swedish Museum of Natural History, Stockholm, Sweden.
OCIC – Observatoire Conservatoire des Insectes de Corse, Corte, France.
PCCQ – Private Collection of C. Quindroit, Angers, France.
PCJS – Private Collection of J. Starý, Olomouc, Czechia.
PCML – Private Collection of M. Lindström, Sweden.
PCKMO – Private Collection of K. M. Olsen, Norway.
PCMCO – Private Collection of M. C. d'Oliveira, Haarlem, The Netherlands.
USC – Entomological Collection, Zoology Department, University of Santiago de Compostela, Spain.
VPMC – Private Collection of V. E. Pilipenko, Moscow, Russia.
ZIN – Zoological Institute, Russian Academy of Sciences, St Petersburg, Russia.
ZMMU – Zoological Museum of Moscow State University, Moscow, Russia.
ZMUB – Zoological Museum, University of Bergen, Bergen, Norway.
ZMUT – Zoological Museum of the University of Turku, Turku, Finland.
Several species listed below have already been reported from different countries (geopolitical units) in the CCW, based on unpublished information and referred to as in litt in the CCW. Here, we publish the collection data for these records as well. If a record was credited to another person other than one of the authors of this paper, we highlight this, otherwise we simply refer to it as the first country record(s). General information and distribution of each species is available on the CCW and can be accessed by the links provided below.
First records from Norway and Russia: RUE.
First record from Portugal.
First records from Russia: RUE.
First records from Norway.
First records from Belarus and Greece (from mainland).
The species was first recorded from Finland in
First records from Russia: RUC. The species was considered doubtful for Norway, see discussion in
First records from Belarus, Latvia, Russia: RUC, RUE and Serbia.
First record from Slovenia.
The species was reported from Belgium by J. Soors (in litt. 2020) in the CCW; here, we publish the collection data of that record.
First records from Albania and Montenegro. The species reported before from mainland Italy and here, we publish the first records from Sardinia and Sicily.
First record from Belarus.
First records from Russia: RUE and Serbia.
First record from Slovenia.
First record from Hungary.
First records from Norway.
First record from Russia: RUE.
First records from Belarus and Russia: RUE.
First record from Sweden.
First record from Romania.
First record from Russia: RUE.
First records from Russia: RUC and Serbia.
First records from Hungary.
First record from Serbia.
Records of Dactylolabis sexmaculata from the European part of Turkey reported by
First records from Greece (from Corfu and mainland).
First records from France (from mainland), Norway, Spain (from mainland) and Sweden.
First records from Romania.
First record from Norway.
First record from Romania.
First record from Serbia.
First record from Serbia.
First record from Russia: RUE.
First record from Serbia.
First records from Italy (from mainland) and Russia: RUE.
First records from Russia: RUE.
First record from Belarus.
First record from Russia: RUE.
First records from Russia: RUC.
First records from Belarus and Russia: RUE.
First records from Albania, Montenegro, North Macedonia, Portugal, Russia: RUC, RUE, RUN and Serbia.
First records from Bulgaria, Romania and Russia: RUC.
First record from Albania.
First records from Finland and Russia: RUE.
First record from Belarus.
First record from Serbia.
First records from Norway.
First records from Norway.
First record from Spain (from mainland).
First record from Norway.
First record from Austria.
First records from Greece (from mainland).
Presence of the species in Norway mentioned in
First records from Belarus and Russia: RUE, RUN.
First records from Albania, Belarus, Iceland and Russia: RUE.
First records from Albania, North Macedonia, Russia: RUE and Serbia.
First record from Norway.
First record from Russia: RUE.
First records from Italy (from Sicily).
First records from North Macedonia, Norway, Russia: RUE, Serbia and Sweden.
Presence of the species in Norway mentioned in
First record from Russia: RUE.
First record from the mainland Spain, previously reported from Canary Is.
First records from Sweden.
First records from Montenegro.
First records from Russia: RUC, RUE.
First records from Latvia and Russia: RUE, RUN.
First records from Belarus and Russia: RUE.
First records from Belarus, Hungary, Latvia, Poland, Russia: RUE and Serbia. We confirm the presence of the species in Russia: RUN.
First record from Russia: RUE.
First records from Hungary and Russia: RUE.
Records of Dicranoptycha paralivescens from Europan part of Turkey reported by
First records from Latvia and Russia: RUE.
First records from Norway. We confirm the presence of the species in Hungary.
First records from Romania and Russia: RUE.
The species removed from the Swedish checklist by
First records from Norway.
First records from Greece (from mainland) and Montenegro.
First record from Estonia.
First record from Slovenia.
Presence of the species in Russia: NC mentioned in
First records from North Macedonia.
First records from Bulgaria, Greece (from mainland) and Hungary.
First record from Serbia.
First record from Portugal.
Figs
First record from Romania.
First records from Hungary, Russia: RUC and Serbia.
First record from Hungary.
First records from Hungary and Serbia.
First record from Greece (from mainland).
First records from Russia: RUN. The species previously reported from mainland Italy and here, we report first time from Sardinia and Sicily.
First record from Serbia.
First records from Belarus and Russia: RUE.
First records from Norway.
First records from Russia: RUC.
First records from Russia: RUE.
First records from Russia: RUC, RUE.
First records from Norway.
First record from Russia: RUE.
First record from Italy (from mainland).
First records from Belarus and Russia: RUE.
First records from Greece (from mainland).
First records from Norway and Russia: RUN.
First records from Russia: RUC, RUN.
First records from Norway and Russia: RUC.
First records from Norway and Sweden.
First records from Norway and Russia: RUC. The species was previously reported from mainland Italy and here, we report for the first time from Sicily.
First records from Russia: RUC.
First records from Russia: RUN.
First records from Russia: RUE.
First records from Greece (from mainland).
First record from mainland Spain, previously reported from Canary Islands.
First record from Serbia.
First records from Belarus, Montenegro and North Macedonia.
First records from Latvia and Russia: RUE, RUN.
First records from Belarus and Romania.
First records from Montenegro and Russia: NC.
First records from Portugal.
First record from Serbia.
First record from Serbia.
First records from Albania and Serbia.
First records from Albania, Greece (from mainland), Montenegro and Romania.
First records from Belarus and Russia: RUE. The species is recorded from Belgium in the CCW by J. Menten and K. Peeters (in litt. 2012); here we publish the collection data for those records.
Presence of the species in Russia: NC mentioned in
First records from Russia: RUE and Serbia.
The species is mentioned from Belgium in the CCW by K. Peeters (in litt. 2012); here, we publish the collection data for that record.
First record from Serbia.
First records from Norway and Russia: RUN. The species is mentioned from Slovenia in the CCW by D. Janevic (in litt. 2017); here, we publish the collection data for that record.
First record from France (from mainland).
First record from Serbia.
First record from Finland.
First records from France (from mainland).
First records from Belarus and Russia: RUE.
Fig.
First records from Norway and Romania. The species is reported from Belgium in the CCW by K. Verhoeyen (in litt. 2012); we publish the collection data for that record.
First records from Russia: RUE.
First records from Belarus.
First records from Norway and Russia: RUE.
First record from Finland.
First records from Norway and Russia: RUW.
First records from Norway.
First records from Portugal and Russia: RUE, NC.
First record from Romania. This species was denoted uncertain for Norway in
First records from Belarus.
First records from Belarus.
First records from Belarus and Malta.
First record from Estonia.
First records from Belarus, Latvia and Russia: RUC, RUE.
First records from Russia: RUC, RUE.
First records from Belarus and Russia: RUE.
First records from Russia: RUC, RUE, RUN.
First records from Hungary and Russia: RUC.
First records from Hungary and Italy (from mainland). The species is reported from Belgium in CCW by K. Peeters (in litt. 2015, 2016); we here publish the collection data for those records.
First record from Serbia.
Species previously reported from mainland France; here, we publish the first record from Corsica.
First records from Albania, Belarus and Spain (from mainland). We confirm the presence of the species in Latvia.
First records from Albania and Russia: RUE.
First records from Romania and Russia: RUN.
First record from Belarus.
First record from Romania.
First records from Russia: RUE.
First records from Belarus.
First records from Russia: RUC, RUE.
First records from Belarus, Hungary and Romania.
First record from France (from mainland).
First records from Russia: RUE.
First records from Russia: RUE.
First records from Latvia, North Macedonia, Russia: RUE, RUN and Serbia.
First records from Hungary and Serbia.
First records from Russia: RUE and Serbia.
First records from Italy (from mainland).
First record from Slovenia.
First records from Norway.
First records from Russia: RUE.
First record from Serbia.
First records from Belarus and Russia: RUE. We confirm the presence of the species in Greece (from mainland).
First records from Norway. Species previously reported form Balearic Islands and here, we report the first records from mainland Spain.
First records from Belarus and Russia: NC.
First records from Norway and Russia: RUC.
First records from Serbia.
First records from Norway.
First record from Hungary.
First records from Russia: NC.
First records from Russia: RUC.
First records from Russia: RUC.
First records from Norway and The Netherlands.
First records from Russia: RUC.
First records from Russia: RUE.
First records from Albania, Greece (from Samothraki) and North Macedonia. Presence of the species in Latvia is confirmed.
First record from Slovenia.
The species was deleted from the Norwegian species list by
First records from Norway.
First records from Norway and Russia: RUE.
First record from France (from mainland).
First records from Russia: RUC, RUE.
First record from Russia: RUE.
The species reported from Finland by
We confirm the presence of the species in Russia: RUE.
First records from Romania.
The species was deleted from the Finnish species list by
First records from Belarus.
First records from Belarus and Russia: RUE.
First records from Russia: RUE.
First records from Greece (from mainland) and Hungary.
First records from Romania.
First records from Hungary and Russia: RUC, RUE.
First record from Spain (from mainland).
First record from Russia: RUC.
Figs
First records from Romania and Serbia.
First records from Russia: RUE and Serbia.
First records from Belarus and Russia: RUE.
First records from Russia: RUN.
First records from Russia: RUE.
First records from Russia: RUE.
Figs
First record from North Macedonia.
First records from Belarus, Romania and Russia: RUE, RUN.
First records from Belarus and Greece (from mainland).
First records from Belarus and Greece (from mainland).
First records from Russia: RUE.
First records from France (from mainland), Latvia and Norway.
First record from Russia: RUC.
First records from Montenegro and Serbia.
First records from Latvia and Norway.
First records from Latvia and Spain (from mainland).
First records from Hungary, Montenegro and Serbia.
First record from Romania.
First record from Serbia.
First records from Finland and Norway.
First records from Norway.
First records from Romania.
Here, we confirm the presence of the species in Serbia.
First records from Greece (from mainland). First record from the mainland of Italy, previously known from Sicily.
First records from Belarus and Russia: RUE.
First record from Italy (from mainland).
First records from North Macedonia and Russia: RUE.
First record from Serbia.
Records of Rhypholophus varius (Meigen and Wiedemann, 1818) from Hungary reported by
First records from Russia: RUE.
First record from Serbia.
First records from Norway and Russia: RUE.
First records from Belarus.
First records from Norway.
First records from Russia: RUC.
First record from Iceland.
Presence of the species in Russia: NC mentioned in
First records from France (from Corsica), Norway and Serbia. The species was first reported from Sweden in CCW, based on records uploaded to
First records from North Macedonia and Romania.
The species was first reported from The Netherlands in the CCW by van Maanen (in litt. 2012). Here, we publish the collection data for that record.
The first author would like to thank Dr. Edina Török (Vácrátót, Hungary) for her enthusiastic help during the collection trips and thank Dr. Lujza Keresztes for allowing the use of facilities and equipment at Babes-Bolyai University (Cluj-Napoca, Romania). The authors are very grateful to collectors for providing valuable specimens and records, with which they contributed to the knowledge of the distribution of European Limoniidae. We also would like to thank Pasquale Ciliberti (NBCN, Leiden) for providing the collection data for some specimens deposited in the NBCN collection. We would like to thank Dr. Daichi Kato (Tokamachi, Japan) for reviewing the manuscript.
Funding Statement
L.-P. Kolcsár was supported by the Japan Society for the Promotion of Science (JSPS) Postdoctoral Fellowships for Research in Japan Short-term Program (Grant Number: PE18038) and Standard Program (Grant Number: P21094). This research was financially supported by the JSPS Grant-in-Aid for Scientific Research (Grant Numbers: 19K21996, 19H02276). The study of N.M. Paramonov was performed in the framework of the state research project АААА-А17-117030310205-9 and the Presidium RAS program no.41 “Biodiversity of natural systems and biological sources of Russia”. J. Mederos thanks the constant support offered by the MCNB curators Glòria Masó-Ros and Berta Caballero-López, as well as the Barcelona Zoo Foundation. The work of A. Polevoi was carried out under state order implemented by Karelian Research Centre RAS (Forest Research Institute). The study of J. Starý was financially supported by the Ministry of Culture of the Czech Republic by institutional financing of long-term conceptual development of the research institution (the Silesian Museum, MK000100595). The work of V.I. Lantsov was carried out with the constant support of the Directorate of the Tembotov Institute of Ecology of Mountain Territories of Russian Academy of Sciences and the staff of the Dagestan State Nature Reserve, to whom he expresses his sincere gratitude, as well with the financial support from the Russian Fund of Fundamental Research (Grant No. 20-54-56030 Iran_t).
Six main subdivisions of the European territory of Russia have been established, as North, Northwest, Central, East, South European Russia and North Caucasus. We list the territories (respublika, oblast, kray) included in these subdivisions, following the geopolitical boundaries of the Catalogue of the Craneflies of the World. We suggest a modification as Permskaya Oblast and Komi-Permyatskiy Avtonomnyy Okrug were merged in 2005 and it is now called Permskiy Kray. The Russian name of the subdivisions and territories are included.