Field Guide to Invasive Alien Species in European Forests

The Slovenian Forestry Institute has published a Field Guide to Invasive Alien Species in European Forests which was translated to English and made available as PDF file.

This guide was first prepared in Slovenian, within the framework of the project Awareness Raising, Training and Measures on Invasive alien Species in forests (LIFE ARTEMIS), which is funded by the European Commission in the framework of the LIFE financial mechanism, the Ministry of the Environment and Spatial Planning of the Republic of Slovenia, the City of Ljubljana and the Slovenian Research Agency.

Kus Veenvliet, J., P. Veenvliet, M. de Groot & L. Kutnar (eds.). 2019. A Field Guide to Invasive Alien Species in European Forests. Nova vas: Institute Symbioisis, so. e.; Ljubljana: The Silva Slovenica Publishing Centre, Slovenian Forestry Institute. [PDF 32 MB]

 Page content last updated on 2019-12-12. Last proofread by Caroline Grounds on 2019-12-12.

Asclepias syriaca L.

English Common milkweed Status LU: uncertain. 1st record: LU & ITW <1836.
Lëtzebuergesch Gewéinlech Seideplanz Status Eur.: established. 1st record: unkn. IAS of EU concern (2017).
Français Asclépiade commune RA: ISEIA: B0, Alert List. Harmonia+: 0,19
Deutsch Gewöhnliche Seidenpflanze Wikipedia: Wikipedia - English Wikipedia - Français Wikipedia - Deutsch Wikipedia - Nederlands | Wikispecies: Wikispecies | CABI
Nederlands Gewoon zijdeplant Back to the list of neophytes

Brief description

Asclepias syriaca sl 2

Asclepias syriaca on ballast near Karnabrunn, district Korneuburg, Lower Austria. Photo: Stefan.lefnaer, 30 June 2012. commons.wikimedia.org

Asclepias syriaca L. originates from North America and is cultivated as an ornamental. It was introduced into European countries from France to Bulgaria and Russia (potentially invasive) (EPPO 2019). A. syriaca colonises a variety of communities from woodlands to cleared grasslands and marshlands. It grows in clumps or patches in meadows, fencerows, roadsides, railways, waste places, reduced-tillage fields, and other open habitats (CABI 2010). Larger populations can displace native plant and animal species, especially if the species penetrates into habitats worthy of protection. The large leaves shade the soil and thus prevent the emergence of other species (Anonymous 2014).

In 2014, Switzerland added Asclepias syriaca L. to its List of Invasive Alien Plants (EPPO 2019, Anonymous 2014).

IAS of Union concern

In 2017, Asclepias syriaca L. was added to the list of invasive alien species of Union concern (Anonymous 2017) which implies that member states shall take all necessary steps to prevent it’s unintentional introduction or spread.

Status and distribution in Luxembourg

Asclepias syriaca L. was first mentioned for Luxembourg by Tinant (1836: 134): “This plant, native to Syria and cultivated in our gardens, is found wild on the rocks around Bellevue [Ed.: Petrusse valley in Luxembourg City], where it is quite common […]. It also grows in the park of Clervaux.”1 Klein (1897: 54-55) mentions the Common milkweed to be grown as an ornamental plant.

Lambinon & Verloove (2012: 534) mention this melliferous species as very rare in the territory of the flora; recently in expansion in Belgium.

The databases contain no observations of Asclepias syriaca L. for Luxembourg (MNHNL, iNaturalist & GBIF, 2020).

Risk assessment

ISEIA protocol

B0 (3+2+2+2) = Alert List. First assessed 22 November 2019 by Christian Ries.

Harmonia+ protocol

Overall risk score 0,19 = (Overall Invasion score 0,52 x Overall Impact score 0,37) (Ries et al. 2020).

0,52Invasion
0,37Impact
0,19Risk

Worldwide distribution

Bibliography

  • Anonymous, 2014. Asclepias syriaca in: Info Flora – Das nationale Daten- und Informationszentrum der Schweizer Flora. URL: https://www.infoflora.ch/de/assets/content/documents/neophyten/inva_ascl_syr_d.pdf
  • Anonymous, 2017. Commission implementing regulation (EU) 2017/1263 of 12 July 2017 updating the list of invasive alien species of Union concern established by Implementing Regulation (EU) 2016/1141 pursuant to Regulation (EU) No 1143/2014 of the European Parliament and of the Council. Official Journal of the European Union L 182: 37-39 (13.7.2017).
  • CABI, 2010. Asclepias syriaca L. [original text by Claire Teeling]. In: Invasive Species Compendium. Wallingford, UK: CAB International. URL: www.cabi.org/isc [accessed 2019-11-22]
  • EPPO, 2019. EPPO Global Database: Asclepias syriaca L. URL: https://gd.eppo.int [accessed 2019-11-22]
  • Klein, E. J., 1897. Die Flora der Heimat (sowie die hauptsächlichsten bei uns kultivierten fremden Pflanzenarten biologisch betrachtet. Eine Anleitung zur selbständigen Beobachtung der Lebens- und Anpassungserscheinungen in der Pflanzenwelt). 552 S., Buchdruckerei Justin Schroell, Diekirch.
  • Lambinon J. & F. Verloove, 2012. Nouvelle flore de la Belgique, du grand-duché de Luxembourg, du Nord de la France et des régions voisines. Sixième édition. Avec la collaboration de L. Delvosalle, B. Toussaint, D. Geerinck, I. Hoste, F. Van Rossum, B. Cornier, R. Schumacker, A. Vanderpoorten et H. Vannerom. Jardin botanique national de Belgique, Meise. CXXXIX + 1195 pp. ISBN : 9789072619884.
  • MNHNL, iNaturalist & GBIF, 2020. Asclepias syriaca in MNHNL-mdata, online portal combining species observation from Recorder-Lux, iNaturalist and GBIF. National Museum of Natural History, Luxembourg. URL: https://mdata.mnhn.lu [Accessed 2020-02-28]
  • Ries, C. & Y. Krippel, 2021. First records of 56 invasive alien vascular plants in Luxembourg. Bulletin de la Société des naturalistes luxembourgeois 123: 115-127. [PDF 241 KB]
  • Ries, C., Y. Krippel & M. Pfeiffenschneider, 2020. Risk assessment after the Harmonia+ protocol of invasive alien vascular plant species in Luxembourg. Bull. Soc. Nat. luxemb. 122: 197-205. [PDF 132 KB]
  • Tinant, F. A., 1836. Flore luxembourgeoise, ou, Description des plantes phanérogames, recueillies et observées dans le grand-duché de Luxembourg, classées d’après le système sexuel de Linnée. 512 p. Luxembourg, J. P. Kuborn.

 Page content last updated on 2021-07-14. Last proofread by Caroline Grounds on 2019-11-27.

  1. Original text: “Cette plante originaire de Syrie et cultivée dans nos jardins, se trouve sauvage sur les rochers aux environs de Bellevue, où elle est assez fréquente […]. Elle croît aussi dans le parc de Clervaux.”[]

First record of Solanum rostratum in Luxembourg

Solanum rostratum Dunal was first observed in the wild in Luxembourg on 21 August 2019 by Jörg Zoldan and Annette Steinbach-Zoldan on the border of a maize field in the municipality of Kayl (Southern part of Luxembourg), during a field study on behalf of SICONA.

Description

Solanum rostratum Dunal (spiny or prickly nightshade, Mexican thistle) is a species of nightshade that is native to the US and northern and central Mexico. It is an annual, self-compatible herb that forms a tumbleweed and has abundant spines on the stems and leaves. In its native range S. rostratum is pollinated by medium- to large-sized bees including bumblebees. The seeds are released when the berries dry and dehisce (split apart) while still attached to the plant (Wikipedia contributors 2019).

This species represents one of the later scientific interests of famed biologist Charles Darwin, who just over a week prior to his death had ordered seeds from a colleague in America, so as to investigate their heteranthery, a topic he was interested in (Wikipedia contributors 2019).

Solanum rostratum is the ancestral host plant of the Colorado potato beetle, Leptinotarsa decemlineata, but this pest adopted the potato, Solanum tuberosum as a new (and more succulent) host, a fact first reported in eastern Nebraska in 1859. It then expanded its range rapidly eastward on potato crops in the next two decades (Wikipedia contributors 2019).

Solanum rostratum as an invasive alien weed

Solanum rostratum is a taprooted annual plant known to be a noxious weed in parts of the USA and Canada, and is included in the Global Compendium of Weeds as an ‘agricultural weed’, ‘casual alien’, ‘environmental weed’, ‘naturalised’, ‘noxious weed’, and ‘weed’ across tropical and temperate regions of the world. It is a fast-growing, vigorous weed native to Mexico and the United States and now widely introduced including into Asia, Oceania and Europe (Austria, Bulgaria, Denmark, Germany, Hungary, Moldova, Russia, Slovakia, UK, Ukraine). The species invades ecosystems by forming dense colonies, and a single plant can produce hundreds of seeds which are dispersed by both biotic and abiotic vectors, as well as self-propelled by its dehiscent fruit (Datiles & Acevedo-Rodríguez 2014).

Bibliography

 Page content last updated on 2019-11-11. Last proofread by Caroline Grounds on 2019-11-11.

Publication of a guide for the identification and management of invasive alien plant species on construction sites

The Luxembourg Nature and Forestry Administration has published a guide on the identification and management of invasive alien plant species on construction sites. The 88-page guide is written in French and can be useful to other concerned parties such as biological stations, municipalities and other state administrations that are active in other activities such as nature sites, the development and maintenance of green spaces and along roads.

The guide can be ordered or downloaded from the following web page: https://environnement.public.lu/fr/publications/conserv_nature/plantes_exotiques_envahissantes.html

Citation: Pfeiffenschneider M. et al., 2019. Guide d’identification et de gestion d’espèces de plantes exotiques envahissantes sur les chantiers. 88 pp. Administration de la nature et des forêts (éd.).

 Page content last updated on 2019-11-11. Last proofread by Caroline Grounds on 2019-11-11.

Lysichiton americanus Hultén & H. St. John

English American skunk-cabbage, swamp lantern Status LU: casual. 1st record: 2022
Lëtzebuergesch Amerikanesch Heckepëppchen Status Eur.: established. 1st record: UK 1901.1 IAS of EU concern (2016).
Français Lysichiton américain RA: ISEIA: B1, Watch List. Harmonia+: 0,14
Deutsch Amerikanischer Stinktierkohl Wikipedia: Wikipedia - English - American skunk-cabbage Wikipedia - Français - Lysichiton américain Wikipedia - Deutsch - Amerikanischer Stinktierkohl Wikipedia - Nederlands - Moeraslantaarn | Wikispecies: Wikispecies - Lysichiton americanus | CABI
Nederlands Moeraslantaarn Back to the list of neophytes

Brief description

Lysichiton americanus 38261Lysichiton americanus Hultén & H. St. John grows in the transition zone of terrestrial, semi-aquatic and aquatic habitats like swamps, fens, wet meadows, marshy and alluvial woodlands, along streams, riverbanks, lakesides and ponds. It has no specific site condition requirements except the presence of saturated organic soils. It is often found in protected semi-natural habitats. Lysichiton reproduces almost exclusively by seeds, which may be dispersed downstream along waterways. However, spread by natural means is not frequent and rather limited. L. americanus has become established locally in swamp forests and associated wetlands in the EPPO region (resulting most of the time from plantation in the site). After some years, its huge leaves build a dense layer excluding light from native species which are usually not adapted to extreme darkness. It can displace and cause local extinction of rare species of mosses and vascular plants (Carex echinata, Viola palustris, and orchids). (Branquart et al. 2010).

IAS of Union concern

In 2016, Lysichiton americanus Hultén & H. St. John was added to the list of invasive alien species of Union concern (Anonymous 2016) which implies that member states shall take all necessary steps to prevent it’s unintentional introduction or spread.

Status and distribution in Luxembourg

Records of somethingelse in Luxembourg. Data source: Recorder-Lux, iNaturalist & GBIF, 2024-11-23.

Lysichiton americanus Hultén & H. St. John was first documented by Christiane Wolff via iNaturalist on 2022-05-31 and was confirmed by Maximilien Oly on 2022-06-02 in the Mamer valley close to Kopstal. Three individuals have been differenciated next to a tributary of the Mamer. The small population was eradicated by the local forestry service. The origin of the population has not been identified.

L. americanus in the Mamer valley on 2022-06-02 – plant in the background (EFOR-ERSA, M. Oly)

L. americanus next to a tributary of the Mamer (EFOR-ERSA, M. Oly)

Risk assessment

ISEIA protocol

B1 (2+3+3+2) = Watch List. First assessed 2019-01-24 by Manou Pfeiffenschneider and Christian Ries.

Harmonia+ protocol

Overall risk score 0,14 = (Overall Invasion score 0,34 x Overall Impact score 0,40) (Ries et al. 2020).

0,34Invasion
0,40Impact
0,14Risk

Worldwide distribution

Bibliography

  • Anonymous, 2016. Commission implementing regulation (EU) 2016/1141 of 13 July 2016 adopting a list of invasive alien species of Union concern pursuant to Regulation (EU) No 1143/2014 of the European Parliament and of the Council. Official Journal of the European Union L 189: 4-5.
  • Branquart, E., I. Stiers, S. Vanderhoeven, W. Van Landuyt, F. Van Rossum & F. Verloove, 2010. Harmonia database: Lysichiton americanus Hultén & H. St. John. Harmonia version 1.2, Belgian Forum on Invasive Species. URL: http://ias.biodiversity.be [accessed on 2019-11-21]
  • CABI, 2021. Lysichiton americanus. In: Invasive Species Compendium. Wallingford, UK: CAB International. URL: www.cabi.org/isc [accessed 2021-03-04]
  • MNHNL, iNaturalist & GBIF, 2020. Lysichiton americanus in MNHNL-mdata, online portal combining species observation from Recorder-Lux, iNaturalist and GBIF. National Museum of Natural History, Luxembourg. URL: https://mdata.mnhn.lu [Accessed 2020-03-02]
  • Oly, M., 2022. Kartierung invasiver gebietsfremder Wasserpflanzen im Großherzotum Luxemburg, Untersuchungen zur Verbreitung von invasiven gebietsfremden Wasserpflanzen in einer beispielhaften Auswahl an Stillgewässern, Musée national d’histoire naturelle, Luxembourg. 117 pp. 
  • Ries, C., Y. Krippel & M. Pfeiffenschneider, 2020. Risk assessment after the Harmonia+ protocol of invasive alien vascular plant species in Luxembourg. Bull. Soc. Nat. luxemb. 122: 197-205. [PDF 132 KB]

 Page content last updated on 2023-08-17. Last proofread by Caroline Grounds on 2019-11-19.

  1. Cf. CABI 2021.[]

Ludwigia peploides (Kunth.) P.H. Raven

English Creeping water primrose Status LU: absent. 1st record: n/a.
Lëtzebuergesch Krauchend Waasserprimmel Status Eur.: established. 1st record: FR 1830s. 1 IAS of EU concern (2016).
Français Jussie rampante RA: ISEIA: A0, Alert List. Harmonia+: 0,44
Deutsch Kriechendes Heusenkraut Wikipedia: Wikipedia - English - Creeping water primrose Wikipedia - Français - Jussie rampante Wikipedia - Nederlands - Kleine Waterteunisbloem | Wikispecies: Wikispecies - Ludwigia_peploides | CABI
Nederlands Kleine waterteunisbloem Back to the list of neophytes

Brief descritpion

Ludwigia peploides 3Ludwigia peploides (Kunth.) P.H. Raven is an amphibious plant living in ponds, lakes, ditches, channels and slow-running rivers as well as in humid meadows. It shows a high tolerance to different water levels. Its growth is favoured by water eutrophication but the plant is able to develop in oligotrophic environments. Both clonal and sexual reproductions contribute to plant spread across watersheds. Water primroses are highly detrimental to the environment in western Europe. They quickly develop and make very thick monospecific floating carpets at the surface of water bodies. They alter the physico-chemical quality of water (reduction of light and dissolved oxygen) and possess an allelopathic activity that influences the water quality throughout the year and reduces the germination and survival rates of other plant species. They outcompete most native water plants and create an anoxic environment detrimental to many plant and animal species. In addition, they modify water flow and cause wetland drying (Branquart et al. 2013).

IAS of Union concern

In 2016, Ludwigia peploides (Kunth.) P.H. Raven was added to the list of invasive alien species of Union concern (Anonymous 2016) which implies that member states shall take all necessary steps to prevent it’s unintentional introduction or spread.

Status and distribution in Luxembourg

Ludwigia peploides (Kunth.) P.H. Raven has not yet been observed in Luxembourg (MNHNL, iNaturalist & GBIF 2020).

Risk assessment

ISEIA protocol

A0 (3+3+2+3) = Alert List. First assessed 24 January 2019 by Manou Pfeiffenschneider and Christian Ries.

Harmonia+ protocol

Overall risk score 0,44 = (Overall Invasion score 0,59 x Overall Impact score 0,75) (Ries et al. 2020).

0,59Invasion
0,75Impact
0,44Risk

Worldwide distribution

Bibliography

  • Anonymous, 2016. Commission implementing regulation (EU) 2016/1141 of 13 July 2016 adopting a list of invasive alien species of Union concern pursuant to Regulation (EU) No 1143/2014 of the European Parliament and of the Council. Official Journal of the European Union L 189: 4-5.
  • Branquart, E., S. Vanderhoeven, W. Van Landuyt, F. Van Rossum & F. Verloove, 2013. Harmonia database: Ludwigia peploides (Kunth.) P.H. Raven. Harmonia version 1.2, Belgian Forum on Invasive Species. URL: http://ias.biodiversity.be [accessed on 2019-11-21]
  • CABI, 2021. Ludwigia peploides. In: Invasive Species Compendium. Wallingford, UK: CAB International. URL: www.cabi.org/isc [accessed 2021-03-04]
  • MNHNL, iNaturalist & GBIF, 2020. Ludwigia peploides in MNHNL-mdata, online portal combining species observation from Recorder-Lux, iNaturalist and GBIF. National Museum of Natural History, Luxembourg. URL: https://mdata.mnhn.lu [Accessed 2020-03-02]
  • Ries, C., Y. Krippel & M. Pfeiffenschneider, 2020. Risk assessment after the Harmonia+ protocol of invasive alien vascular plant species in Luxembourg. Bull. Soc. Nat. luxemb. 122: 197-205. [PDF 132 KB]

 Page content last updated on 2021-03-04. Last proofread by Caroline Grounds on 2019-11-19.

Ludwigia grandiflora (Michx.) Greuter & Burdet

English Water primrose Status LU: absent. 1st record: n/a.
Lëtzebuergesch Groussbléieg Waasserprimmel Status Eur.: established. 1st record: FR 1830s.1 IAS of EU concern (2016).
Français Jussie à grandes fleurs, Grande J. RA: ISEIA: A0, Alert List. Harmonia+: 0,28
Deutsch Großblütiges Heusenkraut Wikipedia: Wikipedia - Français - Jussie à grandes fleurs Wikipedia - Deutsch - Großblütige Heusenkraut Wikipedia - Nederlands - Waterteunisbloem | Wikispecies: Wikispecies - Ludwigia_grandiflora | CABI
Nederlands Waterteunisbloem Back to the list of neophytes

Brief description

Ludwigia grandiflora kz3Ludwigia grandiflora (Michx.) Greuter & Burdet is an amphibious species living in ponds, lakes, ditches, channels and slow-running rivers as well as in humid meadows. It shows a high tolerance to different water levels. Its growth is favoured by water eutrophication but the plant is able to develop in oligotrophic environments. Invasive spread across watersheds is almost exclusively clonal and brought about by the dispersal of vegetative propagules by waterflow. Water primroses are highly detrimental to the environment in western Europe. They quickly develop and make very thick monospecific floating carpets at the surface of water bodies. They alter the physico-chemical quality of water (reduction of light and dissolved oxygen) and possess an allelopathic activity that influences the water quality throughout the year and reduces the germination and survival rates of other plant species. They outcompete most native water plants and create an anoxic environment detrimental to many plant and animal species. In addition, they modify water flow and cause wetland drying (Branquart et al. 2013).

IAS of Union concern

In 2016, Ludwigia grandiflora (Michx.) Greuter & Burdet was added to the list of invasive alien species of Union concern (Anonymous 2016) which implies that member states shall take all necessary steps to prevent it’s unintentional introduction or spread.

Status and distribution in Luxembourg

Ludwigia grandiflora (Michx.) Greuter & Burdet has not yet been observed in Luxembourg (MNHNL, iNaturalist & GBIF 2020).

Risk assessment

ISEIA protocol

A0 (3+3+2+3) = Alert List. First assessed 24 January 2019 by Manou Pfeiffenschneider and Christian Ries.

Harmonia+ protocol

Overall risk score 0,28 = (Overall Invasion score 0,46 x Overall Impact score 0,62) (Ries et al. 2020).

0,46Invasion
0,62Impact
0,28Risk

Worldwide distribution

Bibliography

  • Anonymous, 2016. Commission implementing regulation (EU) 2016/1141 of 13 July 2016 adopting a list of invasive alien species of Union concern pursuant to Regulation (EU) No 1143/2014 of the European Parliament and of the Council. Official Journal of the European Union L 189: 4-5.
  • Branquart, E., I. Stiers, S. Vanderhoeven, W. Van Landuyt, F. Van Rossum & F. Verloove, 2013. Harmonia database: Ludwigia grandiflora (Michx.) Greuter & Burdet. Harmonia version 1.2, Belgian Forum on Invasive Species. URL: http://ias.biodiversity.be [accessed on 2019-11-21]
  • CABI, 2021. Ludwigia grandiflora. In: Invasive Species Compendium. Wallingford, UK: CAB International. URL: www.cabi.org/isc [accessed 2021-03-04]
  • MNHNL, iNaturalist & GBIF, 2020. Ludwigia grandiflorain MNHNL-mdata, online portal combining species observation from Recorder-Lux, iNaturalist and GBIF. National Museum of Natural History, Luxembourg. URL: https://mdata.mnhn.lu [Accessed 2020-03-02]
  • Ries, C., Y. Krippel & M. Pfeiffenschneider, 2020. Risk assessment after the Harmonia+ protocol of invasive alien vascular plant species in Luxembourg. Bull. Soc. Nat. luxemb. 122: 197-205. [PDF 132 KB]

 Page content last updated on 2021-03-04. Last proofread by Caroline Grounds on 2019-11-19.

  1. Cf. CABI 2021.[]

Wandering cribellate spider Zoropsis spinimana discovered in Echternach

On November 15th 2018 Jos A. Massard and Gaby Geimer discovered a female of the wandering cribellate spider Zoropsis spinimana on the window of a pastry shop in Echternach (Massard & Geimer 2018). This spider is a new species for Luxembourg and is currently spreading throughout Europe. The detailed up-to-date distribution of the species can be found on the Luxembourgian Wikipedia (Wikipedia-Bearbeiter 2018).

The home of Zoropsis spinimana is the western Mediterranean region up to the southern edge of the Alps and Dalmatia, as well as North Africa, where it occurs in the open in light forests, under stones and bark, or in and around buildings (Wikipedia contributors 2018).

References

 Page content last updated on 2019-11-11. Last proofread by Caroline Grounds on 2019-11-11.

Increasing understanding of alien species through citizen science

In July 2018 a group of countries launched a European CO-operation in Science and Technology (COST) Action to address multidisciplinary research questions in relation to developing and implementing Citizen Science (CS), advancing scientific understanding of Alien Species (AS) dynamics while informing decision-making, specifically implementation of technical requirements of relevant legislation such as the EU Regulation 1143/2014 on IAS. It will also support the EU biodiversity goals of embedding science within society. The Action will explore and document approaches to establishing a European-wide CS AS network.

All details in the following publication:

Roy H, Groom Q, Adriaens T, Agnello G, Antic M, Archambeau A, Bacher S, Bonn A, Brown P, Brundu G, López B, Cleary M, Cogălniceanu D, de Groot M, De Sousa T, Deidun A, Essl F, Fišer Pečnikar Ž, Gazda A, Gervasini E, Glavendekic M, Gigot G, Jelaska S, Jeschke J, Kaminski D, Karachle P, Komives T, Lapin K, Lucy F, Marchante E, Marisavljevic D, Marja R, Martín Torrijos L, Martinou A, Matosevic D, Mifsud C, Motiejūnaitė J, Ojaveer H, Pasalic N, Pekárik L, Per E, Pergl J, Pesic V, Pocock M, Reino L, Ries C, Rozylowicz L, Schade S, Sigurdsson S, Steinitz O, Stern N, Teofilovski A, Thorsson J, Tomov R, Tricarico E, Trichkova T, Tsiamis K, van Valkenburg J, Vella N, Verbrugge L, Vétek G, Villaverde C, Witzell J, Zenetos A, Cardoso A (2018) Increasing understanding of alien species through citizen science (Alien-CSI). Research Ideas and Outcomes 4: e31412. https://doi.org/10.3897/rio.4.e31412

 Page content last updated on 2019-11-11. Last proofread by Caroline Grounds on 2019-11-11.