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Datasheet

Leucaspius delineatus (sunbleak)

Summary

  • Last modified
  • 23 June 2017
  • Datasheet Type(s)
  • Invasive Species
  • Preferred Scientific Name
  • Leucaspius delineatus
  • Preferred Common Name
  • sunbleak
  • Taxonomic Tree
  • Domain: Eukaryota
  •   Kingdom: Metazoa
  •     Phylum: Chordata
  •       Subphylum: Vertebrata
  •         Class: Actinopterygii
  • Summary of Invasiveness
  • In contrast to its alien and invasive status in the UK and some countries in Northern Europe, L. delineatus is considered rare or vulnerable throughout much of its native range (...

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Pictures

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PictureTitleCaptionCopyright
L. delineatus collected from stillwater in south-eastern England.
TitleAdults
CaptionL. delineatus collected from stillwater in south-eastern England.
CopyrightRobert Britton/Bournemouth University
L. delineatus collected from stillwater in south-eastern England.
AdultsL. delineatus collected from stillwater in south-eastern England.Robert Britton/Bournemouth University
L. delineatus collected from stillwater in south-eastern England. Note scale bar cm.
TitleAdult
CaptionL. delineatus collected from stillwater in south-eastern England. Note scale bar cm.
CopyrightRobert Britton/Bournemouth University
L. delineatus collected from stillwater in south-eastern England. Note scale bar cm.
AdultL. delineatus collected from stillwater in south-eastern England. Note scale bar cm.Robert Britton/Bournemouth University

Identity

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Preferred Scientific Name

  • Leucaspius delineatus (Heckel, 1843)

Preferred Common Name

  • sunbleak

Other Scientific Names

  • Alburnus delineatus Walecki, 1864
  • Aspius ovsianka Czernay, 1851
  • Aspius owsianka Czernay, 1851
  • Cyprinus fischeri Valenciennes, 1844
  • Cyprinus pallasii Arendt, 1860
  • Leucaspius abruptus Heckel & Kner, 1858
  • Leucaspius delineatus caucasicus Berg, 1949
  • Leucaspius delineatus delineatus Heckel, 1843
  • Leucaspius delineatus dimorphus Ruzskii, 1914
  • Leucaspius relictus Varpahowskij, 1889
  • Leuciscus pigmeus Plater, 1861
  • Owsianka czernayi Dybowski, 1862
  • Phoxinellus thracicus Battalgil, 1942
  • Squalius delineatus Heckel, 1843

International Common Names

  • English: belica

Local Common Names

  • Austria: zwerglaube
  • Azerbaijan: ovsianka
  • Belarus: ausianka; malijawka; ovsianka; verkhovka
  • Belgium: ablé de heckel
  • Bulgaria: varlovka; wid dreben
  • China: da yan xiao chi shao yu; xiao chi shao yu
  • Czech Republic: ovsenicka; ovsienka; slunka obecná; slunka stríbritá
  • Denmark: regnloeje; regnløje
  • Estonia: gröplöja; Moderlieschen; mudamaim
  • Finland: allikkosalakka
  • France: ablé de heckel; dos vert; sans mère
  • Germany: Blingo; Malinchen; Moderlieschen; Moderliesken; Moderlisken; Moderrapfen; Modke; Modtke; Mudchen; Mudehen; Mutlose; Mutlosen; Mutterloseken; Rapfenlaube; Schneiderkarpfen; Sonnenfischchen; Weißpiepe
  • Greece: mikrosirko
  • Hungary: kurta baing
  • Iran: mahi riz noqrehai
  • Italy: alburno di Heckel
  • Latvia: ausleja; ovsianka
  • Lithuania: saulažuve
  • Netherlands: vetje
  • Norway: regnlaue
  • Poland: owsianka; slonecznica
  • Romania: albisoara; fufa; pleava; plevusca; sármá; sorean; soret
  • Russian Federation: belaja rybka; beluschka; bil; buza; elez; klin; malijawka; maljuschka; molijawka; moltja; obyknovennaya verkhovka; ovsianka; rogatka; samorodka; verkhovka; werchoplawka
  • Serbia: belka
  • Slovakia: ovsienka
  • Slovenia: belica
  • Sweden: groplöja
  • Switzerland: ablé de heckel; Moderlieschen
  • UK: foy; rain-bleak; white aspe
  • Ukraine: kochoryna; malijavka; mil’ka; ovjes; ovsianka; verkhovka; vivsik; wiwsjanka
  • USA: Moderlieschen

Summary of Invasiveness

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In contrast to its alien and invasive status in the UK and some countries in Northern Europe, L. delineatus is considered rare or vulnerable throughout much of its native range (Lelek, 1987). It is listed as protected under Appendix III of the Bern Convention (Pinder and Gozlan, 2003).

Taxonomic Tree

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  • Domain: Eukaryota
  •     Kingdom: Metazoa
  •         Phylum: Chordata
  •             Subphylum: Vertebrata
  •                 Class: Actinopterygii
  •                     Order: Cypriniformes
  •                         Family: Cyprinidae
  •                             Genus: Leucaspius
  •                                 Species: Leucaspius delineatus

Notes on Taxonomy and Nomenclature

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The common name of Moderlieschen is of German origin. It is a bowdlerised version of an older name which survives in parts of Germany as Mutterloseken. Literally meaning "the little motherless one", this ultimately refers to the fact that the sticky eggs of the Moderlieschen can withstand exposure to air for a remarkably long time. Deposited on water plants, they sometimes stick to the feet of ducks and similar birds and are carried by these to ephemeral ponds. Large numbers of young moderlieschens are thus sometimes encountered when such ponds dry up, and without adult fish being present this gave rise to the belief that they were "motherless" (World Conservation Monitoring Centre, 1996).

Description

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Dorsal spines (total): 2-3; dorsal soft rays (total): 7-10; anal spines: 3; anal soft rays: 10-14; vertebrae: 36-42; scales in lateral line (vertical scale rows): 39–51 (38); pored lateral line scales: 2-15 (0–19). A small fish, usually less than 9 cm although females can be up to 12 cm (Zhukov, 1965; Movchan and Smirnov, 1981; Arnold and Längert, 1995; Cepkyn, 2002; Environment Agency, 2008), with large silvery scales and an inconspicuous intense silvery band along each side. Lateral line incomplete, sometimes absent, with perforated scales. Anal fin longer than dorsal fin. Mouth turns upwards. The lower edge of the body between the pelvic fins and the vent forms a sharp keel. The scales are very loosely attached and fall away if the fish is handled (Kottelat and Freyhof,  2007). Males tend to be smaller in body size than females. For a detailed description and comprehensive biometric information refer to Beyer et al. (2006).

Distribution

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Natural distribution:

Europe and Asia: From the Lower Rhine and northern Germany eastward to southern Baltic basin; Black Sea basin south to Rioni drainage, northern and western Caspian basin (south to Kura drainage); Aegean Sea basin (from Maritsa to Nestos).

Introduced range:

The species has been widely introduced in the upper Rhine drainage and locally in the UK (primarily southwest England), Switzerland (Kottelat, 1997), and in southern France (Farr-Cox et al., 1996).

Distribution Table

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The distribution in this summary table is based on all the information available. When several references are cited, they may give conflicting information on the status. Further details may be available for individual references in the Distribution Table Details section which can be selected by going to Generate Report.

Continent/Country/RegionDistributionLast ReportedOriginFirst ReportedInvasiveReferenceNotes

Asia

ArmeniaPresentPipoyan, 1996
IranPresentNativeCoad, 1995
KazakhstanPresentNativeBlanc et al., 1971
TurkeyPresentNativeBogutskaya, 1997

Europe

AustriaPresentNativeFroese and Pauly, 2011
BelarusWidespreadNativeZhukov, 1965; Blanc et al., 1971
BelgiumPresentIntroduced1910Welcomme, 1988Questionable invasiveness. Contradicting statements in the literature exist as to whether the sunbleak is native to Belgium (see Leu et al., 2009 and refs therein)
Bosnia-HercegovinaPresentNativeFroese and Pauly, 2011
BulgariaPresentNativeBlanc et al., 1971
CroatiaPresentNativeFroese and Pauly, 2011
Czech RepublicPresentNativeBlanc et al., 1971; Lusk et al., 2011
DenmarkPresentNativeFroese and Pauly, 2011
EstoniaPresentNativeEstonian Education and Research Network, 1999
FinlandPresentIntroduced1986-1989Froese and Pauly, 2011
FrancePresentFarr-Cox et al., 1996; Keith and Allardi, 2001
GermanyPresentNativeFroese and Pauly, 2011
GreecePresentNativeBobori et al., 2001
HungaryPresentNativeKeresztessy, 1996
LatviaPresentNativeBlanc et al., 1971
LithuaniaPresentNativeBlanc et al., 1971
MoldovaPresentNativeBlanc et al., 1971
NetherlandsPresentNativeBlanc et al., 1971
NorwayPresent, few occurrencesIntroduced1995Simonsen and Matzow, 2000; Froese and Pauly, 2011
PolandPresentNativeFroese and Pauly, 2011
RomaniaPresentNativeBlanc et al., 1971
Russian FederationPresentNativeFroese and Pauly, 2011
SerbiaPresentNativeBlanc et al., 1971
SlovakiaPresentNativeBlanc et al., 1971
SloveniaPresentNativeFroese and Pauly, 2011
SwedenPresentIntroducedBlanc et al., 1971
SwitzerlandPresentIntroducedKottelat and Freyhof, 2007
UKPresent, few occurrencesIntroduced1980sPinder and Gozlan, 2003
UkrainePresentNativeBlanc et al., 1971

History of Introduction and Spread

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England: Initial dispersal of L. delineatus from their original location in Hampshire is believed to have occurred via the River Test, which was used by the fish as a conduit to disperse and subsequently colonise lakes downstream. As these lakes were then used as stock ponds for other fisheries, this resulted in the inadvertent movement of L. delineatus to lakes in other catchments, such as those in the River Itchen valley. The origins of populations located in the Somerset levels are not clear. Their first official recording was in 1990 although the date of entry is assumed to be in the 1980s (Farr-Cox et al., 1996), when the species first appeared in the Kings Sedgmoor Drain. From here they have spread rapidly, and are now extremely common throughout drains, rivers and connected waterbodies of the Levels. They are also present in several lakes in this area and at a complex of fishing lakes near Sherborne in Dorset. The mechanism for dispersal to these stillwaters is unknown, but it is likely that the fish were introduced either unintentionally or intentionally by man (Pinder and Gozlan, 2003). Introduction of eggs transported on anglers keep-nets has been blamed for further translocations to Uckfield, East Sussex (Zieba et al., 2010).

Norway and Sweden: L. delineatus has recently been recorded in a small catchment area in southernmost Norway. This introduction is assumed to have taken place in the early 1990s, with more recent introduction recorded in the Swedish section of the Enningdalselva hydrosystem, which drains into Norway (Hesthagen and Sandlund, 2007).

Risk of Introduction

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As L. delineatus is small in size <8 cm) and bears close resemblance to other species of the Cyprinidae family (mostly common bleak Alburnus alburnus)there is a relatively high risk of accidental introduction due to misidentification. Given the lack of angling and aquaculture interest in the species, all such introductions are likely to be accidental, occurring through the inadvertent movement of the species within batches of other fish being stocked out.

Habitat

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During their first few weeks of development, L. delineatus are found in shallow marginal habitats where water temperatures are generally higher, supporting blooms of zooplankton and facilitating optimum growth conditions. As the larvae become more developed (around 40 days), these marginal habitats are abandoned in favour of deep water away from the banks where shoals are formed, with individuals swimming just below the water surface. The species tends to naturally inhabit lowland river habitats, especially oxbows and other water bodies only connected to rivers during floods. Often encountered in ponds, steppe lakes and small water bodies not connected to rivers. Found between weeds in shallow pools and creeks, shallow lakes, peat and clay excavations and canals (Kottelat and Freyhof, 2007).

Habitat List

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CategoryHabitatPresenceStatus
Freshwater
Lakes Principal habitat
Ponds Principal habitat
Reservoirs Principal habitat
Rivers / streams Principal habitat

Biology and Ecology

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Genetics

Chromosome number: Haploid=25, Diploid=50 (Klinkhardt et al., 1995). Gilles et al. (2001) reports on the use of the mitochondrial control region in investigating the phylogeny of European cyprinids including L. delineatus.

Reproductive Biology

Sexual maturity is usually attained at age one year and standard length about 4 cm (Movchan and Smirnov, 1981). A batch spawner, individual females will lay several batches of eggs in a spawning season (generally between April and July when temperatures reach 15–20ºC; Cepkyn, 2002), guarded by the male until emergence. This combination of early maturity and nest guarding means that a relatively low number of eggs are produced when compared with other species of the Cyprinidae family. However, their parental care ensures higher egg and larval survival rates. In addition, as their reproduction is spread across an extended spawning period, larvae are less vulnerable to episodic changes in environmental conditions, thus enhancing survival rates (Gozlan et al., 2003).

During spawning, males take up territories around the stems and underside of the leaves of water-lilies, and other flat-surfaced structures. They then clean any accumulated bio-films from the spawning substrate. The ripe females, which develop swollen, distended ovipositors during spawning, then join the male and deposit strips of up to 80 highly adhesive eggs on the spawning substrate, generally 20–30 eggs (Kryzhanovskij, 1949). An individual female can typically deposit a strip of 40 eggs in less than one second. The maximum fecundity reported is 5000 eggs (Cepkyn, 2002). After the female has spawned, the male guards the eggs and encourages other females to deposit their eggs at the same nest site. The male then guards the eggs until they emergence, which takes approximately four days at 22oC (Pinder and Gozlan, 2003).

Longevity

Although individuals have been aged up to 5 years, few individuals will live longer than 2 to 3 years (Beyer, 2008).

Activity Patterns

In autumn, the shoals of fish tend to keep in the surface layers and are rarely found in deeper waters. With the onset of winter, these shoals tend to disperse (at least from these areas) and so the number of fish per unit area decreases rapidly (Pipoyan, 1996).

Nutrition

The fry of L. delineatus feed on phytoplankton and zooplankton (rotifers and crustaceans). The diet of the fishes at age 10-15 days and older can include larvae of the aquatic insects (Chironomidae, Ephemeroptera and others; Movchan and Smirnov, 1981) and, occasionally, flying insects that fall into the water (Pinder and Gozlan, 2003).

Latitude/Altitude Ranges

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Air Temperature

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Parameter Lower limit Upper limit
Mean annual temperature (ºC) 2 32

Water Tolerances

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ParameterMinimum ValueMaximum ValueTypical ValueStatusLife StageNotes
Dissolved oxygen (mg/l) 3.0 Harmful Hatching ceases (Arnold and Längert, 1995)
Salinity (part per thousand) Optimum 6-10 tolerated
Water temperature (ºC temperature) Optimum 2-32 tolerated

Notes on Natural Enemies

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Due to their small size, L. delineatus is vulnerable to predation by predatory fish. In the UK, these will be pike Esox lucius and perch Perca fluviatilis.

Means of Movement and Dispersal

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L. delineatus occurs in large shoals which are most numerous in autumn. They are found between weed beds in shallow pools and creeks, shallow lakes, peat and clay excavations and canals. If exposed to high flows (in spring, autumn and winter) they will seek refuge in areas of reduced flow. In autumn, the shoals are generally found close to the surface but in winter these disperse and individuals are more likely to be encountered in deeper water (Pipoyan, 1996).

Accidental Introduction

L. delineatus has been introduced as a contaminant of stocked fish, and has been translocated on the keep nets of anglers (Zieba et al., 2010).

Intentional Introduction

It is unlikely that L. delineatus has been introduced intentionally as its small size makes them of limited utility to recreational angling and aquaculture.

Pathway Causes

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CauseNotesLong DistanceLocalReferences
Aquaculture Yes
Hitchhiker Yes
Pet trade Yes

Pathway Vectors

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VectorNotesLong DistanceLocalReferences
Aquaculture stock Yes Yes

Impact Summary

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CategoryImpact
Environment (generally) Negative

Environmental Impact

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There have been few studies on the impacts of invasive populations of this species. Pinder and Gozlan (2003) suggest there is considerable potential for their depredation of the eggs of other fishes. Leu et al. (2009) reported their predation on the eggs of native amphibians. In the UK, sunbleak have been found to host the non-native parasite Ergasilus briani, raising concerns that they could aid its spread (Beyer, 2008).

Risk and Impact Factors

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Invasiveness

  • Capable of securing and ingesting a wide range of food
  • Fast growing
  • Gregarious
  • Has a broad native range
  • Has high reproductive potential
  • Highly adaptable to different environments
  • Is a habitat generalist

Likelihood of entry/control

  • Difficult to identify/detect in the field
  • Difficult/costly to control
  • Highly likely to be transported internationally accidentally
  • Highly likely to be transported internationally deliberately

Uses

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Scales were utilized for production of Essence d'Orient, which was used for coating artificial pearls (Kottelat and Freyhof, 2007). It is occasionally used as an aquarium species.

Uses List

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General

  • Pet/aquarium trade

Similarities to Other Species/Conditions

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Diagnosed from other cyprinids in central and eastern Europe by having an incomplete lateral line with about 2-15 (0–19) pored scales, mouth superior, keel covered by scales between pelvic origin and anus, and 11-13.5 branched anal rays (Zhukov, 1965; Movchan and Smirnov, 1981; Kottelat and Freyhof, 2007). An identification fact sheet has been produced by the Environment Agency (UK) (Environment Agency, 2008).

Prevention and Control

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In England and Wales, the Prohibition of Keeping or Release of Live Fish (Specified Species) Order 1998 made under the Import of Live Fish (England and Wales) Act 1980 covers L. delineatus. This legislation requires any individual wishing to keep any listed species as pets (in either aquarium or garden pond), or angling clubs wishing to stock a listed species, first to obtain a licence under the Act. Full details of the Act, including listed species and application procedures, can be obtained from DEFRA or at the website http://www.defra.gov.uk/aahm. A review of control measures has been carried out by the Angling Trust (Angling Trust, 2011), with chemical control using the piscicide rotenone the preferred option.

Gaps in Knowledge/Research Needs

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There are considerable gaps in knowledge on the ecological consequences of the invasion (i.e. impacts) of L. delineatus and their ability and mechanisms to disperse by natural means.

References

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Angling Trust, 2011. Sunbleak fish - Leucaspius delineatus, Management options. Sunbleak fish - Leucaspius delineatus, Management options. http://www.anglingtrust.net/core/core_picker/download.asp?id=1784

Arnold A; Längert H, 1995. Das Moderlieschen (The sunbleak). Magdeburg, Germany: Westarp Wissenschaften, 128.

Berg LS, 1949. Fishes of freshwaters of the U.S.S.R. and adjacent countries. 4th edition. Vol. 2. Moscow & Leningrad, Russia: Adademia Nauk USSR.

Beyer K, 2008. Ecological implications of introducing Leucaspius delineatus and Pseudorasbora parva into inland waters in England., UK: University of Hull.

Beyer K; Miranda R; Copp GH; Gozlan RE, 2006. Biometric relationships between body and bone size of two invasive non-native fish species in the UK: topmouth gudgeon Pseudorasbora parva and sunbleak Leucaspius delineatus. Folia Zoologica, 55:287-292.

Blanc M; Gaudet JL; Banarescu P; Hureau JC, 1971. European inland water fish: a multilingual catalogue. London: Fishing News (Books) Ltd.

Bobori DC; Economidis PS; Maurakis EG, 2001. Freshwater fish habitat science and management in Greece. Aquatic Ecosystem Health and Management, 4:381-391.

Bogutskaya NG, 1997. Contribution to the knowledge of leuciscine fishes of Asia Minor. An annotated checklist of leuciscine fishes (Leuciscinae, Cyprinidae) of Turkey with descriptions of a new species and two new subspecies. Mitt. Hamb. Zool. Mus. Inst. 94:161-186.

Cepkyn EA, 2002. Leucaspius delineatus (Heckel, 1843). In: Atlas of Russian Freshwater Fishes. Vol. 1 [ed. by Reshetnikov YuS]. Moscow, Russia: Nauka, 266-269.

Coad BW, 1995. Freshwater fishes of Iran. Acta Sci. Nat. Acad. Sci. Brno. 29(1):1-64.

Environment Agency, 2008. SUNBLEAK, Leucaspius delineates fact sheet. SUNBLEAK, Leucaspius delineates fact sheet. http://www.environment-agency.gov.uk/static/documents/Leisure/sunbleak_2160808.pdf

Estonian Education and Research Network, 1999. Systematic list of Estonian fishes. World Wide Web Electronic Publication. Systematic list of Estonian fishes. World Wide Web Electronic Publication. http://www.sunsite.eenet.ee/animals/Kalad/kalalist2.htm

Farr-Cox F; Leonard S; Wheeler A, 1996. The status of the recently introduced fish Leucaspius delineates (Cyprinidae) in Great Britain. Fisheries Management and Ecology, 3:193-199.

Froese R; Pauly D, 2011. FishBase. http://www.fishbase.org

Gilles A; Lecointre G; Miquelis A; Loerstcher M; Chappaz R; Brun G, 2001. Partial combination applied to phylogeny of European cyprinids using the mitochondrial control region. Molecular Phylogenetics and Evolution, 19(1):22-33.

Gozlan RE; Pinder AC; Durand S; Bass J, 2003. Could the small size of sunbleak, Leucaspius delineatus (Pisces, Cyprinidae) be an ecological advantage in invading British waterbodies? Folia Zoologica, 52(1):99-108.

Hesthagen T; Sandlund OT, 2007. Non-native freshwater fishes in Norway: history, consequences and perspectives. Journal of Fish Biology, 71(Suppl. D):173-183. http://www.blackwell-synergy.com/doi/abs/10.1111/j.1095-8649.2007.01676.x

Keith P; Allardi J, 2001. Atlas des Poissons d'Eau Douce de France. Paris, France: Muséum National d'Histoire Naturelle, 387 pp.

Keresztessy K, 1996. Threatened freshwater fish in Hungary. In: Conservation of Endangered Freshwater Fish in Europe [ed. by Kirchhofer, A. \Hefti, D.]. Basel, Switzerland: Birkhäuser Verlag, 73-77.

Klinkhardt M; Tesche M; Greven H, 1995. Database of fish chromosomes. Westarp Wissenschaften, 179 pp.

Kottelat M, 1997. European freshwater fishes. Biologia, 52(Suppl 5):1-271.

Kottelat M; Freyhof J, 2007. Handbook of European Freshwater Fishes. Cornol, Switzerland: Publications Kottelat, 646 pp.

Kryzhanovskii SG, 1949. Ecological and morphological patterns of carp, catfish and loaches fish (Cyprinoideiand Siluroidei). Proceedings of the IM/K., Russia: The Academy of Science of the USSR, 5-332.

Lelek A, 1987. The freshwater fishes of Europe. Threatened fishes of Europe. AULA-Verlag, Wiesbaden.

Leu T; Lüscher B; Zumbach S; Schmidt BR, 2009. Small fish (Leucaspius delineatus) that are often released into garden ponds and amphibian breeding sites prey on eggs and tadpoles of the common frog (Rana temporaria). Amphibia-Reptilia, 30:290-293.

Lusk S; Luskova V; Hanel L; Lojkasek B; Hartvich P, 2011. The Red List of lampreys and fishes of the Czech Republic - version 2010. (Cervený seznam mihulí a ryb Ceské Republiky - verze 2010.) In: Biodiverzita ichtyofauny Ceské republiky VIII. 68-78.

Movchan YuV; Smirnov AI, 1981. Fauna of Ukraine. Vol. 8. Fishes. Part 2/1 [ed. by Movchan YuV. \Smirnov, A. I.]. Kiev, Ukraine: Naukova Dumka.

Pinder AC; Gozlan RE, 2003. Sunbleak and topmouth gudgeon - two new additions to Britain's freshwater fishes. British Wildlife, December:77-83.

Pipoyan SK, 1996. A new species for Armenia, Leucaspius delineatus (Cyprinidae). Journal of Ichthyology, 36(1):128-131.

Simonsen JH; Matzow D, 2000. Sunbleak Leucaspius delineatus: A new fish species for Norway. Fauna Oslo, 53(3):151-153.

Welcomme RL, 1988. International introductions of inland aquatic species. FAO Fisheries Technical Paper, No. 294:x + 318 pp.

World Conservation Monitoring Centre, 1996. Leucaspius delineatus. 2006 IUCN Red List of Threatened Species. http://www.iucnredlist.org/

Zhukov PI, 1965. Fish in Belarus. Minsk, Belarus: Nauka i Technika.

Zieba G; Copp GH; Davies GD; Stebbing P; Wesley KJ; Britton JR, 2010. Recent releases and dispersal of non-native fishes in England and Wales, with emphasis on sunbleak Leucaspius delineatus (Heckel, 1843). Aquatic Invasions, 5(2):155-161. http://www.aquaticinvasions.net/2010/AI_2010_5_2_Zieba_etal.pdf

Contributors

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31/08/11Original text by:

Rob Britton, Centre for Conservation Ecology, Bournemouth University, Poole, Dorset, BH12 5BB, UK

Reviewers' names are available on request.

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