Exotic Odonata in Europe

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Exotic Odonata in Europe

Exotic Odonata in Europe Gerold Laister1, Gerhard Lehmann2 & Andreas Martens3 Hans-Hofmann-Ring 3, 4470 Enns, Austria; Haunfeldstraße 14, 6330 Kufstein, Austria; 3 University of Education Karlsruhe, Bismarckstraße 10, 76133 Karlsruhe, Germany; 1

2

Received 2nd February 2014; revised and accepted 24th March 2014 Abstract. Between 1991 and 2011, more than 1,000 adults of exotic odonate species were recorded from glasshouses of a wholesaler dealer of aquarium plants near Wels, Austria. Twenty-three species could be identified to species level. All species were accidentally introduced as eggs or larvae. The majority are widely distributed and common southeastern Asian species. About 17 taxa are first recorded from Europe bringing the list of exotic Odonata in Europe to 41 taxa. These odonates are mainly introduced via aquarist trade and many species have emerged from home aquaria and glasshouses. Currently, tropical plants for aquarists are mainly imported to Europe from Singapore, Indonesia and Thailand, which suggests that the exotic Odonata originated in those countries. So far, the introduction of exotic Odonata species into Europe is not ecologically relevant because none of the introduced species have become established in the wild. However, this study will improve understanding of the significance of trading connections in establishing exotic species which could become invasive. Key words. Dragonfly, neobiota, check-list, aquaristic trade, incidental introduction, dispersal, invasive species, long-term study

Introduction Invasive species are one of the major threats to biodiversity (CBD 2000). In freshwater habitats, one of the most important factors triggering biological invasions is the pet and aquarium trade (Duggan 2010; Chucholl 2013). Non-indigenous ornamental fish and crayfish, in particular, have been introduced and become established in the wild (Duggan et al. 2006; Chucholl 2013). In addition to traded species, several other species are introduced incidentally (Duggan 2010), including Odonata. Non-indigenous species can be divided into ‘invasive species’ (i.e. alien species, neobiota), which are able to establish new populations in a new area, and those which do not, termed ‘exotic species’. Odonatologica 43(1/2) 2014: 125-135

126

G. Laister, G. Lehmann & A. Martens

In Europe, the knowledge of exotic Odonata is based on small studies (e.g. Valtonen 1985; Brooks 1988) and reports of single observations (e.g. Wasscher & Groutbeek 1998; Martens & Schiel 2009). Until now, 25 exotic odonate taxa are known from Europe (Wasscher & Bos 2000; Bos et al. 2007; Martens & Griese 2009; Seehausen 2012a). The last review was by Kipping (2006). Several species were found as adults in tropical glasshouses or aquarium trading shops; others were reared from larva to adult in home aquaria. There is a more or less constant flow of eggs and larvae of exotic odonates into Europe with trading of aquarium plants. Living Odonata are not traded. So far, in Europe, odonates have not become invasive species. The data presented here are from the first long-time study of exotic odonate species found in a glasshouse. An overview of all published records in Europe so far is also presented. The results demonstrate the principal role of the world-wide aquarium trade in the incidental introduction of exotic Odonata to Europe. Study site and methods The study was conducted at two glasshouses of a wholesale dealer of aquarium plants near Wels, Austria (48.08°N, 14.00°E). Both glasshouses together had a total size of ca 650 m². Aquatic plants were cultivated in long rows of 1.2 m wide tanks on the ground as well as in 0.3 m wide containers hanging from the roof above these containers. Both glasshouses had vents for thermal regulation. The climate was humid and tropical throughout the year. Outside, additional small containers of 1 m² were used for cultivation in the summer. Between 1991 and 2011 the first author visited the glasshouses more than 80 times. All odonate imagines seen were collected. The imagines were taken with a small insect net (24 cm diameter). Additionally, in the first few years of the study the small water containers outside were checked in summer. The imagines were stored dry. They were identified to species level by the second author. Native species which entered the glasshouses via vents were also recorded. Exuviae were also collected, but so far have not been identified. Odonatologica 43(1/2) 2014: 125-135

127

Exotic Odonata in Europe

No. of visits

total

2011

2010

2009

2000

1999

1998

1997

1996

1995

1994

  2           1   2   1   2   1 4   4 1       2  

                      2           2   11   3           1           5     1 4                        

                    3 1     2       10   1           8 21     1 1   16       5 14   1             3      

              24               4       11   2         1 2       3   8       2                        

      1       210       2   2 1 27       1 1   4 14       34 5 14 9 46       2 1 10 19 208       10       1       4   3   105       1       9       262       2   4 2 61       2       7       5       3       14       2

296 470 20

1 28

1 49 37

1 56

7 33 35 1034

2

1 13

2

1

Aciagrion sp.   1 Agriocnemis femina 165 43 A. cf. nana   2 A. pygmaea     Argiocnemis rubescens   1 Ceriagrion cerinorubellum   3 C. chaoi   34 Ischnura senegalensis   11 Pseudagrion austrialasiae     P. microcephalum 2 142 Ictinogomphus rapax   4 Epophthalmia vittigera     Brachydiplax chalybea   4 Crocothemis servilia   67 Diplacodes trivialis   1 Hydrobasileus croceus   3 Neurothemis fluctuans 128 101 Orthetrum chrysis   2 O. sabina   26 O. testaceum     Rhyothemis p. phyllis   7 Tholymis tillarga   5 Tramea transmarina   3 Trithemis aurora 1 8 Urothemis signata   2 total

1993

1992

Taxon

1991

Table 1. Exotic Odonata taxa and number of imagines recorded in glasshouses of a plant trader near Wels, Austria. Species recorded in one year only are given bold.

10 27

9

4

3

7

3

2

84

Results Between 1991 and 2011, more than 1,030 imagines of exotic Odonata were collected. Most Zygoptera and nearly all Anisoptera were taken freshly emerged or immature. Some fully mature individuals of Neurothemis fluc­ Odonatologica 43(1/2) 2014: 125-135

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G. Laister, G. Lehmann & A. Martens

tuans were netted outside. Twenty-three taxa were identified to species level and two to genus level (Tab. 1). Some species, such as N. fluctuans, Agrio­ cnemis femina, Pseudagrion microcephalum and Crocothemis servilia, were collected in large numbers. Some species occurred regularly in low numbers, others were recorded only in one year sometimes as a single individual. Discussion This study significantly extends the list of exotic Odonata known in Europe, to 41 taxa (Tab.  2). The high number of records from glasshouses shows that large numbers of exotic odonates derive from eggs or larvae in or on traded aquatic plants. Other forms of introduction, especially the import of imagines in banana boxes or on ships (Kipping 2006; Corbet et al. 1960; Tab.  2) are comparatively rare. The majority of species are of south-eastern Asian origin (Fig. 1) from where aquatic plants for home aquaria were mainly imported from countries, such as Singapore, Indonesia and Thailand (Gueudre & Tison 2007). Annual differences in the species composition and number of individuals (Tab. 1) suggest that there are large changes in the composition of imported material within a short time.

Figure 1. Origin of exotic Odonata in Europe. Odonatologica 43(1/2) 2014: 125-135

Exotic Odonata in Europe

129

Table 2. Check-list of exotic Odonata taxa in Europe. Taxon

Distribution Remarks

References

Aciagrion sp.

Asia, Africa

Austria, glasshouse. First record for Europe

This study

Agriocnemis femina (Brauer, 1868)

Asia

The Netherlands, aquarium, Austria, glasshouses

Wasscher & Goutbeek 1998; this study

A. cf. nana Laidlaw, 1914

Asia

Austria, glasshouse. First records for Europe

This study

A. pygmaea (Rambur, 1842)

Asia

Germany, aquarium trader, Austria, glasshouses

Martens & Griese 2009; this study

Argia fumipennis (Burmeister, 1839)

North America

UK, glasshouse

Brooks 1988; this study

Argiocnemis rubescens Asia Selys, 1877

Austria, glasshouse. First records for Europe

This study

Ceriagrion cerinorubellum (Brauer, 1865)

Asia

UK, glasshouse; Germany, vicinity of glasshouse; Austria, glasshouse

Brooks 1988; Kipping 2006; this study

C. chaoi Schmidt, 1964

Asia

Austria, glasshouse. First records for Europe

This study

Enallagma signatum (Hagen, 1861)

North America

UK, glasshouse

Brooks 1988

Ischnura posita (Hagen, 1861)

North America

UK, glasshouse

Brooks 1988

I. ramburii (Selys, 1850)

North America

Germany, aquarium trader

Seehausen 2012a

I. senegalensis (Rambur, 1842)

Asia, Africa

Finland, aquarium trader; UK, glasshouse, fish tanks and garden ponds; Germany home aquaria and aquarium trader; Austria, glasshouses

Valtonen 1985; Brooks 1988; Kipping 2006; Parr 2010; Benken & Kommander 2011; Lambertz & Schmied 2011; Seehausen 2012b; this study

Mecistogaster sp.

Central and South America

Poland, first record of an Rudow 1898 exotic sp. in Europe

Pseudagrion austrialasiae Selys, 1876

SE Asia

Austria, glasshouse. First record for Europe

This study

Odonatologica 43(1/2) 2014: 125-135

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G. Laister, G. Lehmann & A. Martens

Taxon

Distribution Remarks

Pseudagrion microcephalum (Rambur, 1842)

SE Asia, Australia

Anax guttatus (Burmeister, 1839)

Asia, UK, glasshouse N Australia, Pacific

Brooks 1988

A. gibbosulus Rambur, 1842

Asia, N Australia

UK, glasshouse

Brooks 1988

Ictinogomphus Asia decoratus (Selys, 1854)

The Netherlands, aquarium

Lieftinck 1978

I. rapax (Rambur, 1842)

Austria, glasshouse. First records for Europe

This study

Epophthalmia vittigera Asia bellicosa Lieftinck, 1948

Austria, glasshouse. First record for Europe

This study

Brachydiplax chalybea Asia Brauer, 1868

Austria, glasshouse. First records for Europe

This study

Celithemis eponina (Drury, 1773)

North America

The Netherlands, glasshouse near Den Haag in 1988

Bos et al. 2007; M. Wasscher in litt.

Crocothemis servilia (Drury, 1773)

Asia

UK, glasshouse and fish tanks; The Netherlands, aquarium; Austria, glasshouses; Germany, aquaria; Poland, aquarium trader

Agassiz 1981; Brooks 1988; Waning Vos 2005; Wasscher & Ruiter 2005; Raab et al. 2007; Parr 2010; Seehausen & von Blanckenhagen 2012; Seehausen 2012b; Buczyński & BielakBielecki 2012; this study

Diplacodes trivialis (Rambur, 1842)

Asia

Austria, glasshouse. First record for Europe

This study

Erythemis simplicicollis North (Say, 1840) America

UK, glasshouse

Brooks 1988

Hydrobasileus croceus SE Asia (Brauer, 1867)

Austria, glasshouse. First records for Europe

This study

Neurothemis fluctuans SE Asia (Fabricius, 1793)

Finland, aquarium trader; The Netherlands, glasshouse and aquarium; Austria, glasshouses

Valtonen 1985; Wasscher & Goutbeek 1998; Waning Vos 2007; this study

Asia

Odonatologica 43(1/2) 2014: 125-135

References

Finland, aquarium trader; Valtonen 1985; Germany, aquaria; Martens & Schiel Austria, glasshouses 2009; Seehausen & von Blanckenhagen 2012; this study

131

Exotic Odonata in Europe

Taxon

Distribution Remarks

Orthetrum chrysis (Selys, 1891) O. japonicum (Uhler, 1858) O. sabina (Drury, 1770) O. testaceum (Burmeister, 1839) Pantala flavescens (Fabricius, 1798)

Asia

Rhodothemis rufa (Rambur, 1842) Rhyothemis p. phyllis (Sulzer, 1776) Sympetrum eroticum (Selys, 1883) S. infuscatum (Selys, 1883) Tholymis tillarga (Fabricius, 1798) Tramea transmarina Brauer, 1867 Trithemis aurora (Burmeister, 1839) Urothemis bisignata Brauer, 1868 U. signata (Rambur, 1842)

E Asia

References

Austria, glasshouse. This study First records for Europe Finland, aquarium trader Valtonen 1985

Asia, UK and Austria, North Africa glasshouses Asia Austria, glasshouse. First record for Europe tropics and UK, imagines on board sub­tropics, of ships; cosmo­poli­ Germany, imago in tan banana box Asia UK, glasshouse SE Asia

Austria, glasshouse. First records for Europe E Asia Lithuania, one young male caught in the wild E Asia Nousty, France, 09-viii-2006; photo taken by J. Deffarges in a garden. First record for Europe Asia, Africa, Austria, glasshouse. N Australia, First records for Europe Pacific SE Asia UK, Austria, glasshouses Asia Asia Asia

Austria, glasshouse. First records for Europe UK, glasshouse Austria, glasshouse. First records for Europe

Brooks 1988; Raab et al. 2007; this study This study Ford 1954; Corbet et al. 1960; Kipping 2006 Agassiz 1981; Brooks 1988 This study Stanionyte 1989; Bernard 2005 J.-P. Boudot in litt.

This study Agassiz 1981; Brooks 1988; this study This study Agassiz 1981; Brooks 1988 This study

Many of the imported species, such as Anax guttatus, Ceriagrion cerino­ ru­bellum, Diplacodes trivialis, Neurothemis fluctuans, Orthetrum sabina, Pseud­agrion microcephalum, and several others, breed in disturbed, degraded areas in their native range and are common in areas with dense human settlement (Orr 2001). In most cases typical habitats are paddy fields, ditches, canals, and open ponds. In primary forests these species are rare or absent (Orr 2001). Odonatologica 43(1/2) 2014: 125-135

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G. Laister, G. Lehmann & A. Martens

So far in Europe, there is no case of an exotic odonate becoming established in the wild. This is in contrast to other parts of the world. One odonate species, Crocothemis servilia, has successfully invaded the subtropics and tropics outside its native range. So far the species is established in Florida (Paulson 1978; Daigle & Rutter 1984; first record 1975), Cuba (Flint 1996; first record 1994), and O’ahu, Hawaii (Polhemus 1995; first record 1994). The biggest number of established introduced Odonata species have been found in Hawaii, where about eight species have been recorded (Englund 2005). In Japan, the occurrence of Pseudagrion australasiae is thought to be as a result of an accidental introduction (Karube et al. 2011). Some cases are ambiguous. Pantala flavescens is a casual migrant in Europe (Buczyński et al. 2014). Therefore records in the wild are hard to interpret. Another case is Pachydiplax longipennis (Burmeister). In early September 1999, a female of that North American species was found dead on the oil rig Sedco 706 in the North Sea at 60°38’N, 01°39’E (Parr 2000a). It is not clear if the animal reached the oil rig alive. It could have been carried on board a helicopter or on the jet stream. Because there is no evidence that the individual reached Europe alive it has not been included on the European list. Prospect The increase in exotic odonates illustrates the globalization of the aquarium trade and its significance in the dispersal of neobiota. (1) A new trend is aquatic plant breeding in North Africa for the European market. This will reduce the risk of tropical Asian dragonflies being introduced into Europe. However, this increases the risk of an introduction and establishment of tropical Asian dragonfly species in those parts of Africa, which belong to the same thermal area. (2) The picture could change rapidly if South American plants and odonates begin to be imported into Europe. (3) So far none of the introduced species have established populations in the wild. In the temperate part of Europe there is no risk when plants originate from tropical regions. However, the introduction of plant material from other temperate regions, such as North America or China could Odonatologica 43(1/2) 2014: 125-135

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Exotic Odonata in Europe

pose a risk. In Mediterranean Europe, especially on islands, the establishment of a subtropical south-eastern Asian species could be possible. Islands are especially at risk because of their relative small number of species, less diverse genotypes of the inhabitants, and open niches because of isolation effects. (4) Further unidentified species, especially Libellulidae and Coenagrionidae, will probably be found. There are several reports of unidentified Pseud­ agrion specimens (Brooks 1988; Schmidt 1990; Seidenbusch 1996; Seehausen 2012b) which might belong to P. microcephalum. Special requests in aquarist journals and forums (Seehausen 2012b), photo guides from south-eastern Asia (Tang et al. 2010), national checklists (Bos et al. 2007; Parr 2000b; Martens 2014) as well as book chapters (Wildermuth & Martens 2014) will improve knowledge of imported Odonata. Acknowledgements We would like to thank Adolf Niederhumer, Aquarium Linden company, for allowing the first author access to the glasshouses. Martin Schorr provided hard to obtain literature. Many thanks to Sónia Ferreira and Vincent Kalkman, for their request for a review on exotic dragonflies at the 1st European Congress of Odonatology in Vila do Conde, Portugal, which prompted a closer look at this topic and the analysis of unpublished findings. Finally, many thanks to Steve Brooks and Marcel Wasscher, for their very useful comments on the manuscript. References Agassiz D. 1981. Further introduced China mark moths (Lepidoptera: Pyralidae) new to Britain. Entomologist’s Gazette 32: 21-26

Bos F., Wasscher M. & Reinboud W. 2007. Veldgids Libellen. 5th edition. KMNV Uitgeverij, Zeist

Benken T. & Kommander M. 2011. Die Senegal-Pechlibelle (Ischnura senegalensis) schlüpft in einem Aquarium bei Ulm. Mercuriale 11: 51-52

Brooks S.J. 1988. Exotic dragonflies in north London. Journal of the British Dra­ gonfly Society 4: 9-12

Bernard R. 2005. First record of Aeshna affinis Vander Linden, 1820 in Lithuania (An­ isoptera: Aeshnidae) and corrective notes on the Lithuanian Odonata checklist. Notulae odonatologicae 6: 53-55

Buczyński P. & Bielak-Bielecki P. 2012. Crocothemis servilia (Drury, 1773) (Odonata: Libellulidae) introduced with aquarium plants to Lublin (Poland). Annales Universitatis Mariae Curie-Skłodowska, Lublin, Polonia (C, Biologia) 67: 21-26 Odonatologica 43(1/2) 2014: 125-135

134 Buczyński P., Shapoval A.P. & Buczyńska E. 2014. Pantala flavescens at the coast of the Baltic Sea (Odonata: Libellulidae). Odonatologica 43 (1/2): 3-11 CBD [Convention on Biological Diversity]. 2000. Invasive alien species: Global strategy on invasive alien species. Convention on Biological Diversity, UNEP/CBD/SBTTA/6/ INF/9: i-ix, 1-52. CBD, Montreal. Online on the Internet, URL (10-iv-2014): http://www. cbd.int/doc/meetings/sbstta/sbstta-06/ information/sbstta-06-inf-09-en.pdf Chucholl C. 2013. Invaders for sale: trade and determinants of introduction of ornamental freshwater crayfish. Biological Invasions 15: 125-141 Corbet P.S., Longfield C. & Moore N.W. 1960. Dragonflies. Collins, London Daigle J.J. & Rutter R.P. 1984. New county records for Crocothemis servilia (Dru.). from Florida, United States (Anisoptera: Libellulidae). Notulae odonatologicae 2: 63 Duggan I. 2010. The freshwater aquarium trade as a vector for incidental invertebrate fauna. Biological Invasions 12: 3757-3770 Duggan I., Rixon C.A.M. & MacIsaac H.J. 2006. Popularity and propagule pressure: determinants of introduction and establishment of aquarium fish. Biological Invasions 8: 377-382 Englund R.A. 2005. Threats to native aquatic insect biodiversity in Hawai’i and the Pacific, and challenges in their conservation. PhD thesis, University of Hawai’i Flint O.S. Jr. 1996. The Odonata of Cuba, with a report on a recent collection and checklist of the Cuban species. Cocuyo 5: 17-20 Ford W.K. 1954. Lancashire and Cheshire Odonata (some further notes). The North Western Naturalist 1954: 602-603 Odonatologica 43(1/2) 2014: 125-135

G. Laister, G. Lehmann & A. Martens Gueudre F. & Tison J.-M. 2007. Pathway analysis: aquatic plants imported in France. EPPO Reporting Service No. 1 (2007/016). European and Mediterranean Plant Protection Organization, Paris. Online on the internet, URL (10-iv-2014): http://archives. eppo.int/EPPOReporting/2007/Rse-0701. pdf?utm_source=archives.eppo.org&utm_ medium=int_redirect Karube H., Futahashi R., Odajima T., Odajima A. & Odojima K. 2011. An occurrence of a southeastern Asiatic species, Pseud­agrion australasiae Selys in Japan: a possible case of accidental introduction. Tombo 53: 111-114 Kipping J. 2006. Globalisierung und Libellen: Verschleppung von exotischen Libellenarten nach Deutschland (Odonata: Coenagrionidae, Libellulidae). Libellula 25: 109-116 Lambertz M. & Schmied H. 2011. Records of the exotic damselfly Ischnura senegalensis (Rambur, 1842) from Bonn (Germany). Bonn zoological Bulletin 60: 211-213 Lieftinck M.A. 1978. Over een onopzettelijke kweek van een tropisch-Aziatische libel uit een verwarmd aquarium in Nederland (Odonata, Gomphidae). Entomologische Berichten, Amsterdam 38: 145-150, erratum 163 Martens A. 2014. Exotische Libellenarten in Deutschland. Libellula Supplement 14: [in press] Martens A. & Griese J. 2009. Verschleppung von Agriocnemis pygmaea mit exo­ tischen Wasserpflanzen nach Deutschland (Odonata: Coenagrionidae). Libellula 28: 187-189 Martens A. & Schiel F.-J. 2009. Pseud­agrion microcephalum mit exotischen Wasser­ pflanzen nach Deutschland ver­schleppt. Mercuriale 9: 27-29

Exotic Odonata in Europe Orr A.G. 2001. An annotated checklist of the Odonata of Brunei with ecological notes and descriptions of hitherto unknown males and larvae. International Journal of Odonatology 4: 167-220 Parr A. 2000a. Blue Dasher Pachydiplax longipennis (Burmeister) on an oil rig in the North Sea. Atropos 10: 3-5 Parr A. 2000b. An annotated list of the Odonata of Britain and Ireland. Atropos 11: 10-20 Parr A. 2010. Records of exotic Odonata in Britain during 2010. Atropos 41: 39-42 Paulson D.R. 1978. An Asiatic dragonfly, Crocothemis servilia (Drury), established in Florida (Anisoptera: Libellulidae). Notulae odonatologicae 1: 9-10 Polhemus D.A. 1995. New Heteroptera and Odonata (Insecta) records and range extensions in the Hawaiian Islands. Bishop Museum Occasional Papers 42: 42-43 Raab R., Chovanec A. & Pennerstorfer J. 2006. Libellen Österreichs. Springer, Wien Rudow [F]. 1898. Entomologische Notizen. Societas Entomologica, Stuttgart 13: 83 Schmidt E.[G.] 1990. Libellenbeobachtungen in der Stadt: Der Botanische Garten in Bonn. Tier und Museum 2: 42-52 Seehausen M. 2012a. Ischnura ramburi mit Wasserpflanzen nach Europa importiert (Odonata: Coenagrionidae). Libellula 31: 7-13 Seehausen M. 2012b. Exotische Libellenlarven, eingeschleppt mit Wasserpflanzen. DATZ [Deutsche Aquarien- und TerrarienZeitschrift] 65 (11): 52-55 Seehausen M. & von Blanckenhagen B. 2012. Pseudagrion microcephalum (Rambur, 1842) und Crocothemis servilia (Drury, 1773) – zwei exotische Libellenarten in Hessen. Libellen in Hessen 5: 58-62

135 Seidenbusch R. 1996. Schlupf eines unbe­ kannten Pseudagrion-Weibchens. Sulzbach-Rosenberger Libellenrundbrief 3: [with­out pagination, 11-13] Stanionyte A.P. 1989. Sympetrum eroticum Selys new to the Lithuanian SSR Odonata species, found in 1988. In: Jonaitis V (Ed.) New and rare insect species of the Lithuanian SSR; records and descriptions of 1989: 9-11 Tang H.B., Wang L.K. & Hämäläinen M. 2010. A photographic guide to the dragonflies of Singapore. Raffles Museum of Biodiversity Research, Singapore Valtonen P. 1985. Exotic dragonflies imported accidentally with aquarium plants to Finland. Notulae odonatologicae 2: 8788 Waning Vos H. 2005. Het verhaal over Crocothemis servilia. NVL-Nieuwsbrief 4: 8 Waning Vos H. 2007. Neurothemis fluctuans. NVL-Nieuwsbrief 11: 7 Wasscher M.T. & Bos F.G. 2000. The European dragonflies: notes on the checklist and on species diversity. Odonatologica 29: 31-43 Wasscher M.[T.] & Goutbeek E. 1998. Tropische Neurothemis fluctuans (Fabricius) in Nederlandse plantenkas. Brachytron 2: 16-17 Wasscher M.[T.] & Ruiter E. 2005. Crocothemis servilia, een niewe exotische libellensoort voor Nederland. NVL-Nieuwsbrief 4: 9 Wildermuth H. & Martens A. 2014. Taschenlexikon der Libellen Europas. Quelle & Meyer, Wiebelsheim

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