Coccinella septempunctata (seven-spot ladybird)

Pictures

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Adult

Coccinella septempunctata (seven-spot ladybird); adult. Ahlen, Germany. April 2011.

©Quartl/via wikipedia - CC BY-SA 3.0

Larva

Coccinella septempunctata (seven-spot ladybird); larva. Saltoun Wood, East Lothian, Scotland, UK. July 2014.

©S. Rae/via wikipedia/flickr - CC BY 2.0

Larva

Coccinella septempunctata (seven-spot ladybird); larva, predating aphids.

©Gilles San Martin/via flickr - CC BY-SA 2.0

Eggs

Coccinella septempunctata (seven-spot ladybird); egg batch, on a thistle (Cirsium spp.).

©Gilles San Martin/via flickr - CC BY-SA 2.0

Adults

Coccinella septempunctata (seven-spot ladybird); aggregation of adults in a conifer (Pinus spp.). Saltoun Forest, East Lothian, Scotlan, UK. October 2010.

©S. Rae/via flickr - CC BY 2.0

Adult

Coccinella septempunctata (seven-spot ladybird); freshly emerged adult, adjacent pupal case. Israel. April 2011.

©Eran Finkle/via flickr - CC BY 2.0

Inter-species predation

Harmonia axyridis (harlequin ladybird); two larvae feeding on a Coccinella septempunctata (7-spot ladybird) larva.

©Mike Majerus/UK Ladybird Survey

Identity

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Preferred Scientific Name

Coccinella septempunctata Linnaeus, 1758

Preferred Common Name

seven-spot ladybird

Other Scientific Names

Coccinella 7-punctata (L.)

International Common Names

English: ladybird, seven spot; ladybird, seven-spotted; seven-dotted ladybird; sevenspotted lady beetle
French: coccinelle à sept points

Local Common Names

Denmark: alm. mariehone; syvprikkt mariehone
Finland: seitsenpistepirkko
France: coccinelle à sept points
Germany: Marienkaefer, 7-punktiger; Siebenpunktmarienkaefer
Norway: 7-prikket marihone
Sweden: sjuprickig nyckelpiga
UK: 7 spot ladybird; seven spot ladybird
USA: seven spotted ladybeetle; sevenspotted ladybeetle; seven-spotted ladybeetle

EPPO code

COCISE (Coccinella septempunctata)

Summary of Invasiveness

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Coccinella septempunctata, the seven-spot ladybird, is a widespread species originally native from Europe, Asia and Northern Africa. Because of its potential as a biological control agent of crop insect pests, several intentional introductions have occurred in the USA. As a result, the species is now widely distributed and established in North America. The broad geographic success of the species, predominant in most habitats of the Palaearctic and a successful invader of the Nearctic Region, may be due to its ecological plasticity, based on genetic and phenotypic polymorphisms. In the USA, C. septempunctata has been reported to compete with and displace several native Coccinellidae species, causing a decrease in their survivorship and abundance.

Taxonomic Tree

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Domain: Eukaryota
Kingdom: Metazoa
Phylum: Arthropoda
Subphylum: Uniramia
Class: Insecta
Order: Coleoptera
Family: Coccinellidae
Genus: Coccinella
Species: Coccinella septempunctata

Notes on Taxonomy and Nomenclature

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There are two subspecies within the species C. septempunctata: Coccinella septempunctata brucki Mulsant, 1866 and Coccinella septempunctata septempunctata Linnaeus, 1758, as recognized by the Integrated Taxonomic Information System (ITIS, 2009).

Rathour and Singh (1991) suggested that ovariole number can be used as a taxonomic character in Coccinellidae and that multiplication, reduction, and stabilization of ovariole number have occurred in the evolution of coccinellids. C. septempunctata has 62 ovarioles per ovary.

Description

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Egg

C. septempunctata passes through three stages: egg, larva and pupa, to develop into an adult. The eggs are elongate, oval and laid on plants, often near to prey. C. septempunctata, like most ladybird species, fix their eggs at one end so they are in an upright position. Eggs take approximately 4 days to hatch, although increasing ambient temperature reduces the length of the egg stage; at 15°C eggs take 10.3 days to hatch, compared to 1.8 days at 35°C (Majerus and Kearns, 1989).

The number of eggs laid per C. septempunctata pair was 284 and 524, respectively, feeding on Hyalopterus pruni and Schizaphis graminum under laboratory conditions, compared with 213 eggs/pair on H. pruni in the field (Varvara et al., 1982).

Larva

The larvae remain on the eggs for approximately 1 day post egg hatch. They eat the egg shells, those of neighbouring larvae and any infertile eggs. The larvae suck body fluid from aphids and as they grow, they will also eat legs and antennae. The fluid from their gut is regurgitated into the aphid allowing some pre-digestion before the body fluid of the aphid is sucked in. There are three moults and four larval instars. Prey density, temperature (Majerus and Kearns, 1989) and prey species (Obrycki and Orr, 1990) can affect the length of the larval stage. Larvae reared at 23±2°C and LD 16:8 on Acyrthosiphon pisum required an average of 13.1 days to complete development, compared to 16 days on Rhopalosiphum maidis (Obrycki and Orr, 1990). One hundred and thirty-four to two hundred and fifty individuals of S. graminum were consumed per larva of C. septempunctata (Varvara et al., 1982).

Rhoades (1996) published a key to the first and second instars that does not rely on colour patterns of live larvae, rather the relative placement and characteristics of the prominent setae on the tergum of the abdomen.

Pre-pupa

The fourth instar larva does not feed for at least 24 hours pre pupation. The tip of the abdomen is attached to the plant substrate; it is immobile and hunched. This is the pre-pupa stage.

Pupa

The final larval skin of the pre-pupa sheds right back to the point of attachment. At 20°C, the pupal stage lasts for 8.4 days. This stage is not completely immobile because it is able to raise and lower the fore region in response to perceived danger (Majerus and Kearns, 1989). The colour of the pupa is variable in some species and C. septempunctata develop into light orange pupae at high temperatures and dark-brown or blackish ones at low temperatures (Majerus and Kearns, 1989).

Prey type affects development time of C. septempunctata. Sattar et al. (2008) reported total larval and pupal duration as 18.3±0.53 and 4.9±0.58 days, respectively, when C. septempunctata were fed Aphis gossypii. Mean percent emergence of males and females was reported as 36.6±2.98 and 56.6±4.21, respectively. The male to female sex ratio was recorded as 1:1.5. When this coccinellid was fed five different aphid species, Arshad and Rizvi (2007) found that overall development time was significantly longer on Lipaphis erysimi.

Adult

The front of the pupal case splits to allow the adult to emerge. When first emerged, the wings and elytra are very soft and barely pigmented. The colouration develops over time and the red colour of the background deepens over the next weeks and months. The dark colours are derived from melanins and the lighter ones from carotenes (Majerus and Kearns, 1989). Although some adults vary considerably in colour pattern, C. septempunctata show little variation, although spot number ranges between 0 and 9, and variation in spot strength is said to be “considerable” (Majerus and Kearns, 1989). Typically, adults are red with seven black spots.

Average longevity was recorded by Kontodimas et al. (2007) as 94.9 days at a constant temperature of 25±1°C, 65±2% RH and 16L:8D. Average total fecundity, net reproductive rate and intrinsic rate of increase were found to be 1996.8 eggs/female, 1004.1 females/female and 0.118 females/female/day, respectively.

For detailed information on all coccinellid life stages, including mating, comparison of sexes and a key to British ladybirds, refer to Majerus and Kearns (1989). Also refer to Marzo (1982), who presented morphological observations of Rhynchota and Coleoptera spermatheca, including C. septempunctata, and Thornham et al. (2007) who studied sexual dimorphism in the distribution and biometrics of the palpal sensilla of C. septempunctata and a description of a new sensillum.

Distribution

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C. septempunctata is native to temperate Europe, North Africa and Asia, but has become established in North America, where it has been found hundreds and thousands of kilometres away from its original release site (Krafsur et al., 2005). Montana and Washington are thought to be the most westerly records in the USA (Rice, 1992).

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.

Last updated: 17 Feb 2021

Continent/Country/Region	Distribution	Origin	First Reported	Reference	Notes
Africa
Algeria	Present	Native		Lecompte et al. (2016) Dobzhansky (1933)
Egypt	Present			Ali et al. (2005)
Libya	Present			El-Aish et al. (2004)
Morocco	Present			CABI Data Mining (Undated)
Asia
Azerbaijan	Present			CABI (Undated)	Original citation: Siberian Zoological Museum (2018)
Bangladesh	Present			Maula et al. (2010)
China	Present			Pan ShiQing et al. (2002)
-Anhui	Present			Tian et al. (1982)
-Fujian	Present			Pan ShiQing et al. (2002)
-Gansu	Present			Pan ShiQing et al. (2002)
-Guangdong	Present			Pan ShiQing et al. (2002)
-Guangxi	Present			Pan ShiQing et al. (2002)
-Guizhou	Present			Pan ShiQing et al. (2002)
-Hebei	Present			Yan et al. (1981) Yan et al. (1983)
-Heilongjiang	Present			Pan ShiQing et al. (2002)
-Henan	Present			Wu et al. (1981)
-Hubei	Present			Pan ShiQing et al. (2002)
-Hunan	Present			Pan ShiQing et al. (2002)
-Jiangsu	Present			Tian et al. (1982)
-Jiangxi	Present			Pan ShiQing et al. (2002)
-Jilin	Present			Pan ShiQing et al. (2002)
-Liaoning	Present			Pan ShiQing et al. (2002)
-Ningxia	Present			Zhang ZhiKe et al. (2009)
-Qinghai	Present			Pan ShiQing et al. (2002)
-Shaanxi	Present			Yan et al. (1983)
-Shandong	Present	Native		Fan GuangHua et al. (2009)
-Shanxi	Present			Shi GuangLu et al. (2005)
-Sichuan	Present	Native		Lecompte et al. (2016) Pan ShiQing et al. (2002)
-Tibet	Present			Pan ShiQing et al. (2002)
-Xinjiang	Present			Xu JianJun et al. (2005)
-Yunnan	Present			Xu (1985) Chen GuoHua et al. (2009)
-Zhejiang	Present			Pan ShiQing et al. (2002)
Georgia	Present	Native		Lecompte et al. (2016) Barjadze et al. (2009)
India	Present	Native		Kajita et al. (2012) Dobzhansky (1933)
-Andhra Pradesh	Present			Raja et al. (2004)
-Arunachal Pradesh	Present			Singh et al. (2006)
-Assam	Present			Bhattacharyya et al. (2006)
-Bihar	Present			CABI (Undated)	Original citation: Ali et al. (2007)
-Chhattisgarh	Present			Patel and Thakur (2005)	Raipur
-Delhi	Present			Asha Thomas et al. (2008)
-Gujarat	Present			Vekaria and Patel (2005)	North
-Haryana	Present			Narendra Kumar et al. (2005)
-Himachal Pradesh	Present			Ravinder Kumar and Gupta (2006)
-Karnataka	Present			Patil et al. (2008)
-Kerala	Present			Johnson (1983)
-Madhya Pradesh	Present			Bhardwaj et al. (1986)	Chhattishgarh region
-Maharashtra	Present	Native		Vennila et al. (2007)
-Manipur	Present			Bilashini et al. (2007)
-Meghalaya	Present			Doddamani et al. (2017)
-Odisha	Present			Mandal and Patnaik (2008)
-Punjab	Present			Ramesh Arora et al. (1998)
-Rajasthan	Present			CABI Data Mining (Undated)
-Sikkim	Present			Agarwala and Raychaudhuri (1981)
-Tamil Nadu	Present			Sahayaraj and Martin (2003)
-Uttar Pradesh	Present			Gaurav Kumar et al. (2009)
-West Bengal	Present			Nath and Sen (1976)
Iran	Present	Native		Lecompte et al. (2016) Ghahhari and Hatami (2000) Mesbah et al. (2016)
Iraq	Present			Amin and Muhammed (2008)	Erbil City, Kurdistan region
Israel	Present	Native		Kajita et al. (2012)
Japan	Present	Native		Kajita et al. (2012) Dobzhansky (1933)
-Hokkaido	Present			Ohashi et al. (2003)
-Honshu	Present			Ohashi et al. (2003)
-Kyushu	Present			Murakami and Tsubaki (1984) CABI (Undated)
-Ryukyu Islands	Present			CABI (Undated)	Original citation: Takao Museum (2018)
-Shikoku	Present			CABI (Undated)	Original citation: Takao Museum (2018)
Jordan	Present			Capinera (2008) Shannag and Obeidat (2008)
Kazakhstan	Present	Native		Lecompte et al. (2016) CABI (Undated)
Kuwait	Present			CABI Data Mining (Undated)
Kyrgyzstan	Present			CABI (Undated)	Original citation: Siberian Zoological Museum (2018)
Lebanon	Present			Capinera (2008)
Mongolia	Present			Pan ShiQing et al. (2002)
Pakistan	Present			Talpur and Khuhro (2004)
South Korea	Present			Pan ShiQing et al. (2002) Dobzhansky (1933)
Syria	Present	Native		Kajita et al. (2012) Shahadi et al. (2002) Capinera (2008)
Taiwan	Present			Pan ShiQing et al. (2002)
Tajikistan	Present			CABI (Undated) CABI Data Mining (Undated)	Original citation: Siberian Zoological Museum (2018)
Turkey	Present	Native		Lecompte et al. (2016) Kızıltepe et al. (2009) Bașar and Yașar (2011) CABI (Undated)
Turkmenistan	Present			CABI (Undated)	Original citation: Siberian Zoological Museum (2018)
Uzbekistan	Present	Native		Kajita et al. (2012) CABI (Undated)
Europe
Albania	Present			de Jong et al. (2014)
Andorra	Present			de Jong et al. (2014)
Austria	Present			Klausnitzer (2006)
Belarus	Present			de Jong et al. (2014)
Belgium	Present	Native		Jansen and Hautier (2008)
Bosnia and Herzegovina	Present			de Jong et al. (2014)
Bulgaria	Present			Dirimanov and Dimitrov (1975) Natskova (1977) Angelova and Tzolova (2005)
Croatia	Present			de Jong et al. (2014)
Cyprus	Present			CABI (Undated)	Original citation: Özden et al. (2006)
Czechia	Present	Native		Kajita et al. (2012) Leslie et al. (2009) Št'astná and Psota (2013)
Denmark	Present	Native		Lecompte et al. (2016)
Estonia	Present			de Jong et al. (2014)
Finland	Present			de Jong et al. (2014)
France	Present	Native		Kajita et al. (2012) Bourguet et al. (2002) Ferre (2008)
-Corsica	Present			de Jong et al. (2014)
Germany	Present	Native		Lecompte et al. (2016) Freier et al. (2007)
Greece	Present			Zarpas et al. (2007)
Hungary	Present			de Jong et al. (2014)
Ireland	Present			de Jong et al. (2014)
Italy	Present	Native		Kajita et al. (2012) Barbagallo et al. (1982) Tumminelli et al. (2004)
Latvia	Present			de Jong et al. (2014)
Liechtenstein	Present			de Jong et al. (2014)
Lithuania	Present			de Jong et al. (2014)
Luxembourg	Present			de Jong et al. (2014)
Malta	Present			de Jong et al. (2014)
Moldova	Present	Native		Kajita et al. (2012) CABI Data Mining (Undated)
Monaco	Present			de Jong et al. (2014)
Netherlands	Present	Native		Kajita et al. (2012) Blom et al. (1985)
North Macedonia	Present			de Jong et al. (2014)
Norway	Present			de Jong et al. (2014)
Poland	Present	Native		Lecompte et al. (2016) Miszczak (1974) Sądej (2000)
Portugal	Present	Native		Lecompte et al. (2016) Gonçalves et al. (2007)
-Azores	Present			de Jong et al. (2014)
-Madeira	Present			de Jong et al. (2014)
Romania	Present			Voicu et al. (1987) Tălmaciu and Tălmaciu (2005)
Russia	Present	Native		Kajita et al. (2012) Pukinskaya et al. (1981)
-Northern Russia	Present			de Jong et al. (2014)
-Southern Russia	Present			de Jong et al. (2014)
San Marino	Present			de Jong et al. (2014)
Serbia	Present			Tomanović et al. (2008)
Slovakia	Present			Selyemová et al. (2007)
Slovenia	Present			de Jong et al. (2014)
Spain	Present	Native		Kajita et al. (2012) Núñez Pérez et al. (1992)
-Balearic Islands	Present			de Jong et al. (2014)
Sweden	Present	Native		Lecompte et al. (2016)
Switzerland	Present	Native		Kajita et al. (2012) de Jong et al. (2014) Lecompte et al. (2016)
Ukraine	Present	Native		Kajita et al. (2012) Phoofolo and Obrycki (2000)
United Kingdom	Present	Native		Kajita et al. (2012) Majerus and Kearns (1989)
North America
Canada	Present	Introduced	1973	COSEWIC (2012)
-British Columbia	Present			Humble (1991)	Nursery near Prince George
-Manitoba	Present	Introduced		Turnock et al. (1990)
-Nova Scotia	Present	Introduced		Cormier et al. (2000)	Cape Breton Island
-Ontario	Present			CABI Data Mining (Undated)
-Prince Edward Island	Present			Garbary et al. (2004)
-Quebec	Present	Introduced		Hoebeke and Wheeler (1980) Turnock et al. (1990) Lucas et al. (2007)
-Saskatchewan	Present	Introduced		Turnock et al. (1990)
United States	Present	Introduced		CABI (Undated a)	Present based on regional distribution.
-Alaska	Present			Hagerty et al. (2009)
-Arkansas	Present			Cranshaw et al. (2000)
-Colorado	Present			Cranshaw et al. (2000)
-Connecticut	Present	Introduced		Angalet and Jacques (1975) Hoebeke and Wheeler (1980)
-Delaware	Present	Introduced		Angalet and Jacques (1975) Hoebeke and Wheeler (1980)
-Georgia	Present	Introduced		Hoebeke and Wheeler (1980) Tillman et al. (2004)
-Hawaii	Present	Introduced		HEAR (2018)
-Iowa	Present	Introduced		Obrycki et al. (1987) Gardiner et al. (2009) CABI (Undated)
-Kansas	Present			CABI Data Mining (Undated)
-Kentucky	Present			Harwood et al. (2006)
-Maine	Present			Majerus (1994)
-Maryland	Present			Staines et al. (1990)
-Michigan	Present	Introduced		Gardiner et al. (2009)
-Minnesota	Present	Introduced		Gardiner et al. (2009) CABI (Undated)
-Missouri	Present	Introduced		Obrycki et al. (1987)	In 33 of 48 counties surveyed
-Montana	Present	Introduced		Rice (1992)
-Nebraska	Present	Introduced		Kriz et al. (2006)
-New Jersey	Present	Introduced		Angalet and Jacques (1975) Hoebeke and Wheeler (1980) Turnock et al. (1990)
-New York	Present	Introduced		Angalet and Jacques (1975) Hoebeke and Wheeler (1980)
-North Dakota	Present	Introduced		CABI (Undated)	Original citation: Hesler and Petersen (2008)
-Oklahoma	Present	Introduced		Hoebeke and Wheeler (1980)
-Oregon	Present			CABI Data Mining (Undated)
-Pennsylvania	Present	Introduced	1979	Hoebeke and Wheeler (1980)
-Rhode Island	Present			NatureServe (2018)
-South Carolina	Present			CABI Data Mining (Undated)
-South Dakota	Present	Introduced		Elliott et al. (1996) CABI (Undated)
-Texas	Present			Parajulee and Slosser (2003)
-Utah	Present	Introduced		Evans (2000)
-Virginia	Present			Rhoades (1996)
-Washington	Present	Introduced		Rice (1992)
-West Virginia	Present	Introduced		Brown and Miller (1998)	First collected in 1983
-Wisconsin	Present	Introduced		Gardiner et al. (2009)
-Wyoming	Present			Hesler et al. (2014)
South America
Brazil	Present			CABI (Undated a)	Present based on regional distribution.
-Para	Present			Greathead (Undated)
Chile	Present	Introduced		González (2018)

History of Introduction and Spread

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Native to temperate Europe and Asia, C. septempunctata is a voracious predator of aphids and as such has been employed in biological control programmes. Between 1956 and 1971 numerous attempts were made to introduce C. septempunctata (originating from France, India, Norway and Sweden) into North America, and although a subsequent generation of the ladybird was recovered later in the year in some areas of New Jersey, Ohio and California, no permanent establishment was found until 1973 (Angalet and Jacques, 1975). In June 1973, several individuals were found in Hackensack Meadowlands, Bergen County, New Jersey; this was thought to be due to accidental introduction (Angalet and Jacques, 1975). Later intentional introductions resulted in establishment in Connecticut, Delaware, Maine, New York, Oklahoma and Pennsylvania, USA (Majerus, 1994). The intentional release of this species into New Brunswick, Canada in 1959 failed to establish, but its occurrence now in Quebec is thought to be the result of accidental introduction or spread from Maine (Larochelle, 1979).

In a 1990 survey, C. septempunctata was found at high altitudes in the Rocky Mountains; records of this species in Montana and Washington are said to be the most westerly records in the USA (Rice, 1992).

Since becoming established in North America, C. septempunctata has been found hundreds and thousands of kilometres from its original release site (Krafsur et al., 2005).

Risk of Introduction

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The European ladybird, C. septempunctata has been intentionally introduced into North America for the biological control of aphid pests and is now established in various areas of the USA and Canada. Climate is described by Hoddle (2004) as a fundamental requirement for the establishment of a species outside its area of origin; the recipient location must be similar in climate to the area of origin. Due to the fact that C. septempunctata is a habitat generalist and can develop on various aphid species, further spread into neighbouring areas with suitable climatic and habitat conditions is probable.

Hodek and Michaud (2008) examined factors that could be responsible for the broad geographic success of C. septempunctata. The species is highly mobile and eurytopic, shows an inhibition in ovipositing in the presence of conspecific larval trails, which represents an adaptive advantage favouring increased egg dispersal and serves to lower the risk of offspring mortality due to cannibalism. C. septempunctata is able to suspend oviposition, and displays heterogenous voltinism and diapause tendencies, therefore some populations feed and reproduce on randomly occurring aphid populations. Other adaptations include pre-hibernating mating, the fact that there is no reproductive diapause in males and the tendency to produce offspring in excess of the carrying capacity of local food resources.

Habitat

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C. septempunctata is a generalist ladybird and as such develops on a wide range of aphids, and thus on a wide range of plants. It is described by Majerus (1994) as “found in most habitats [in Britain]”. Majerus and Kearns (1989) describe the habitat preference of this ladybird as “diverse” and Majerus (1994) stated that early stages have been found on over 250 species of plant native to Britain, and numerous imported and cultivated varieties. The plants listed are: stinging nettles, thistles, bedstraws, umbellifers, knapweeds, vetches, willowherbs, tansy, ragworts, fat hen, goose-foots, chamomiles, bramble, Scots pine, wheat, barley, Brassica spp., beans, peas, sugar beet and rape. In a study to establish the status of exotic and previously common ladybirds in South Dakota, Hesler and Kieckhefer (2008) reported that C. septempunctata was present in a wide range of habitats surveyed. It was the most abundant larval coccinellid in intercropped wheat-alfalfa, where it preyed on Hyalopterus pruni. This species has also been found under rocks in alpine tundra, and on a snow field in the Rocky Mountains, at elevations of 3475 m (Rice, 1992).

Majerus and Kearns (1989) also stated that a regular cycle of movement from one host plant to another throughout the year is displayed by some generalist coccinellids.

Habitat List

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Category	Sub-Category	Habitat	Presence
Terrestrial	Managed	Cultivated / agricultural land	Present, no further details
Terrestrial	Managed	Protected agriculture (e.g. glasshouse production)	Present, no further details
Terrestrial	Managed	Managed forests, plantations and orchards	Present, no further details
Terrestrial	Managed	Managed grasslands (grazing systems)	Present, no further details
Terrestrial	Managed	Urban / peri-urban areas	Present, no further details
Terrestrial	Natural / Semi-natural	Natural forests	Present, no further details
Terrestrial	Natural / Semi-natural	Natural grasslands	Present, no further details
Terrestrial	Natural / Semi-natural	Rocky areas / lava flows	Present, no further details
Terrestrial	Natural / Semi-natural	Scrub / shrublands	Present, no further details

Biology and Ecology

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Genetics

Rozek and Holecová (2002) studied C-banded karyotypes in the males of seven ladybird species, including C. septempunctata and reported that they possess 2n=20, n (males) = 9+Xyp.

Krafsur et al. (1992) used polyacrylamide gel electrophoresis to reveal the genetic diversity in exotic North American populations of C. septempunctata, recently spread across the northern Nearctic. Sixteen of the 28 putative loci were polymorphic; the average gene diversity for all loci was reported to be 0.1589±0.041. Gene frequencies were estimated at eight polymorphic loci in naturally occurring North American beetles from Arkansas, Delaware, Iowa, Kansas, New York, Oregon and Michigan. F2 beetles from colonies that originated in Eurasia were also studied, along with field-collected beetles from France, Greece and Sicily. Gene diversity among the Nearctic beetles was as great as that among those from the Palaearctic. Only 29% of the variance in gene frequencies was between USDA cultures, Palaearctic and Nearctic beetles; and 71% of the genetic variation was shared by beetles within the 21 subpopulations. There was no evidence of bottlenecks or drift among the Nearctic subpopulations, and gene flow was essentially unrestricted.

In a later study to identify the origins of C. septempunctata and thus help to explain movement and dispersal, and efficacy of the predator as an aphid control agent over large areas, Haubruge et al. (2002) employed random amplified polymorphic DNA (RAPD) analysis. Populations were sampled from Belgium and analysed for RAPD DNA variation; to quantify the genetic diversity within the species and to monitor the spatial foraging. They concluded that RAPD analysis can be a valuable technique for studies of intraspecific genetic variation in C. septempunctata.

Majerus et al. (1998) reported the occurrence of a melanic C. septempunctata (C. septempunctata f. purpuralis) male, collected from the field in the UK. It was mated with F1 females and the conclusion was that C. septempunctata f. purpuralis is controlled by a single recessive allele, with reduced viability.

Reproductive Biology

In general, female ladybirds have the capacity to lay over 1000 eggs each; however, this number is rarely reached (Majerus and Kearns, 1989). Reports for C. septempunctata vary from those such as Sattar et al. (2008) stating that a single female laid 177.0 ± 23.03 eggs during her entire life period, to an average total fecundity of 1996.8 eggs/female in life table studies by Kontodimas et al. (2007). In addition, the latter authors reported average longevity of 94.9 days, a net reproductive rate (Ro) of 1004.1 females/female, and intrinsic rate of increase (rm) as 0.118 females/female/day.

Environmental factors, ease with which a mate is found and the condition of the female affect the number of eggs laid (Majerus and Kearns, 1989). For example, Arshad and Rizvi (2008) observed the survival and fecundity of C. septempunctata at varying temperatures (18±1, 24±1°C and 28±1°C), 65±5% RH and 12 h L:12 h D photoperiod under laboratory conditions for two successive generations. They reported that the highest potential fecundity and net reproductive rate of C. septempunctata were obtained at 24±1°C (165.67 eggs/female and 41.09 females/female/generation, respectively) and the lowest at 28±1°C (146.63 eggs/female and 29.70 females/female/generation, respectively). A temperature of 28±1°C resulted in the maximum finite, intrinsic and annual rate of increase (1.0876, 0.0840 females/female/day and 2.04×10¹³/annum, respectively) and at 18±1°C the minimum values of these factors were observed (1.0794, 0.0764 females/female/day and 1.281×10¹²/annum, respectively). The minimum mean length of generation and doubling time were 40.77 and 8.26 days, respectively at 28±1°C and the maximum values were 48.27 and 9.08 days, respectively at 18±1°C.

Other factors affecting egg hatch include infertility; the condition and age of the parents, and time since the female last mated affect the fertility of the eggs (Majerus and Kearns, 1989). Sattar et al. (2008) reported 98.3±2.79% egg hatch and 82.2±6.20% survival of larvae to the pupal stage in biology experiments of C. septempunctata. Eggs are also vulnerable to predation by other predators (e.g. lacewings) (Majerus and Kearns, 1989).

Phoofolo and Obrycki (2000), studying phenotypic variation in reproductive traits of C. septempunctata and Propylea quatuordecimpunctata, found that 47 to 61% of C. septempunctata females laid their first batch of eggs within the first two weeks of their adult life. Repeatability estimates of daily parity for C. septempunctata populations were reported to be 0.32 for Iowa (USA), 0.35 for Delaware (USA), 0.28 for France, and 0.33 for Ukraine.

When aphid prey (Acyrthosiphon pisum) were removed, Kajita and Evans (2009) showed that C. septempunctata reduces oviposition. This is resumed when prey are again offered. The rapid responses to changes in prey availability shown by C. septempunctata is suggested by the authors as a contributing factor to the greater abundance and reproductive success of this introduced species relative to the native Coccinella transversoguttata in western North American alfalfa fields that exhibit widely varying pea aphid densities.

C. septempunctata undergoes oosorption as a means of reserving resources under poor prey conditions and enhancing future reproductive effort when prey conditions improve (Kajita and Evans, 2009).

C. septempunctata requires diapause prior to the onset of reproduction.

Physiology and Phenology

Hodek et al. (1989) studied the physiological state of C. septempunctata collected in mid-hibernation in Greece following exposure to 23.5±2°C and LD 18:16 (long-day conditions) or LD 12:12 (short-day conditions). It was reported that the delay in oviposition was similar under both short-day and long-day conditions. Oviposition rate and fecundity were higher under short-day conditions. Generally it was found that, the metabolic rate of females increased from an average of about 30 mm³ O₂/h per individual at the onset of exposure to a maximum of about 120 mm³ O₂/h per specimen. It was concluded that in warm greenhouses where the natural photoperiod is not modified, most adults collected in overwintering sites can be induced to reproduce prematurely in early winter.

Studying the phenology of this coccinellid in Greece, Katsoyannos et al. (1997) found that field-collected specimens underwent four complete and a fifth partial generation per year. Only adults of the first generation reproduced within the year they emerged. Adults of the first and fifth generations died before winter; those of the second to fourth generations overwintered successfully. The greatest numbers of eggs were laid by females of the first and second generations. Visual counts made in the open field at Kopais Plain in central Greece and on the summit of the adjacent Mount Kitheron indicated that all C. septempunctata instars were abundant in the plain between April and June, becoming scarce from July until the end of the warm period of the year following spring. C. septempunctata were not found in the plain in winter. C. septempunctata adults were present all year round on the mountain summit, singly and in aggregations, except in May. The most numerous arrivals of adults were noticed on the mountain in June and emigrations of adults from there were noticed from March until the end of April.

Nutrition

C. septempunctata is able to develop on a wide range of aphids (Majerus and Kearns, 1989) and Hodek and Honek (1996) state that over 20 aphid species are essential prey for this ladybird. Under laboratory conditions, the mean daily consumption of aphids by a pair of C. septempunctata was recorded as 32 individuals of Hyalopterus pruni and 41 of Schizaphis graminum, and 134-250 individuals of S. graminum were consumed per larva (Varvara et al., 1982). Sattar et al. (2008) reported that mean consumption of Aphis gossypii per C. septempunctata adult was 77.8±5.15, and 21.9, 55.9, 107.4 and 227.3 aphids were consumed by a single larva during 1st, 2nd, 3rd and 4th instars, respectively.

However, aphidophagous coccinellids such as C. septempunctata will feed on other food types. Triltsch (1999) found fungal spores (mainly conidia of Alternaria), pollen and thrips, together with other non-aphid arthropods, in the guts of field-collected adults and larvae. There are reports of it feeding on the greenhouse whitefly, Trialeurodes vaporariorum (Ravinder Kumar and Gupta, 2006), nymphs of the citrus psylla, Diaphorina citri (Gupta and Bhatia, 2000), hawthorn mealybug nymphs, Phenacoccus dearnessi (Cranshaw et al., 2000), the Colorado potato beetle (Leptinotarsa decemlineata) (Gusev et al., 1983) and Bemisia tabaci (Zhang et al., 2007). It is also cannibalistic (Varvara et al., 1982; Omkar and Maurice, 2009).

Dixon and Guo (1993) determined the direct and indirect effects of aphid abundance on egg and cluster size in C. septempunctata. It was found that when food supply varied, there was a tendency for cluster size and number of eggs produced per day to vary, but not egg size. Host density has also been shown to significantly affect aphid consumption by all larval instars. Solangi et al. (2007) found that consumption by C. septempunctata larvae significantly increased with increasing density of the mustard aphid, Lipaphis erysimi.

Giles et al. (2002) studied the influence of alfalfa cultivar (Medicago sativa) on suitability of Acyrthosiphon kondoi for the survival and development of Hippodamia convergens and C. septempunctata. It was found that the resistant lucerne cultivar (54H55) would have little to no effect on the nutritional value of A. kondoi for both ladybird species.

Associations

Majerus and Kearns (1989) stated that this species has been found overwintering with Adalia 2-punctata (2 spot), Coccinella 11-punctata (11 spot), Propylea 14-punctata (14 spot), Micraspis 16-punctata (16 spot), Psyllobora 22-punctata (22 spot), Aphidecta obliterate (larch), Calvia 14-guttata (cream-spot) and Exochomus 4-pustulatus (pine) ladybirds.

Environmental Requirements

When conditions such as temperature and food availability are unsuitable for development, the adults overwinter in almost any slightly sheltered position close to the ground (Majerus and Kearns, 1989), in aggregations (Honek et al., 2007). For a general overview of overwintering in ladybirds refer to Majerus and Kearns (1989), for example.

Under laboratory conditions, Hodek (1970) observed the termination of diapause in C. septempunctata, collected at the end of August from overwintering sites in southern Moravia. It was found that the termination of diapause in females exposed to constant temperatures of 0, 5 or 12°C or to fluctuating temperatures (range 5-12°C), for 3, 6 or 9 weeks and then kept with 18 h light at 23°C during the light period and 18°C during the dark one, was more effective at 5 and 12°C compared with 0°C. Moreover at all temperatures, the termination of diapause was greater in the 6-week than in the 3-week samples.

C. septempunctata are only active in daylight. Zotov (2009) showed that sensitivity to light, photopreferendum and locomotory activity are managed by endogenous circadian oscillators. Results indicated that sensitivity to light (100, 1000 or 7000 lux) was maximum in the daytime (periods of activity) and minimum at night (rest period) irrespective of temperature (17 or 26°C).

When studying altitudinal distribution of coccinellids in mountain spruce forests of Pol’ana Mountains, west Carpathians, in Slovakia at altitudes ranging from 600-1300 m.a.s.l., Selyemová et al. (2007) found that C. septempunctata was most abundant at the middle altitudes studied (900-925 m). This beetle has been found at elevations of 3475 m in the Rocky Mountains (Rice, 1992).

Natural enemies

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Natural enemy	Type	Life stages	References	Biological control in	Biological control on
Allothrombium triticium	Parasite
Anisolobus indicus	Pathogen
Araneus diadematus	Predator		Majerus (1994)
Araneus quadratus	Predator		Majerus (1994)
Araniella opisthographa	Predator
Atomus parasiticus	Parasite
Beauveria	Pathogen	Adults	Iperti (1964)
Beauveria bassiana	Pathogen
Beauveria brongniartii	Pathogen		Ghazavi et al. (2005)
Cheilomenes sexmaculata	Predator
Chrysoperla carnea	Predator
Coccipolipus macfarlanei			Hajiqanbar et al. (2007)
Cordyceps memorabilis	Pathogen	Arthropods\|Larvae	Pacioni and Frizzi (1977)
Coturnix coturnix japonicus	Predator
Dinocampus coccinellae	Parasite	Adults	Hodek (1973)
Gregarina dasguptai	Pathogen			India	tea
Hesperomyces virescens	Pathogen		Harwood et al. (2006)
Hexamermis	Parasite		Rubtsov (1971)
Homalotylus eytelweinii	Parasite	Arthropods\|Larvae	Majerus and Kearns (1989)
Homalotylus flaminius	Parasite	Arthropods\|Larvae	Myartseva (1981)
Ischiodon scutellaris	Predator
Medina separata	Parasite
Mesocyclops aspericornis	Predator
Mesocyclops darwini	Predator
Oomyzus scaposus	Parasite	Arthropods\|Larvae	Majerus and Kearns (1989)
Pachyneuron solitarium	Parasite
Paecilomyces farinosus	Pathogen	Adults	Pacioni and Frizzi (1977)
Parus caeruleus	Predator
Parus major	Predator	Adults	Dolenská et al. (2009)
Passer montanus	Predator
Phalacrotophora berolinensis	Parasite	Arthropods\|Pupae	Disney et al. (1994)
Phalacrotophora fasciata	Parasite	Arthropods\|Pupae	Disney et al. (1994)
Philodromus cespitum	Predator
Pica pica	Predator
Trichomalopsis submarginatus	Parasite

Notes on Natural Enemies

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Various predators and parasitoids have been recorded attacking C. septempunctata; refer to Majerus (1994) for a detailed overview. Due to the relatively large size of a C. septempunctata larva, it may contain up to six Homalotylus eytelweini (Majerus and Kearns, 1989). Another gregarious parasite found in this species is Tetrastichus coccinellae; up to 25 have recorded from a single larva (Majerus and Kearns, 1989). A list of arthropod parasites and hyperparasites was published by Schaefer and Semyanov (1992), which included 14 Hymenoptera from six families, two Diptera from two families, and two ectoparasitic mites.

Rubtsov (1971) described and illustrated the mermithid Hexamermis coccinellae coccinellae n. subsp. found in C. septempunctata. Even though Matchanov et al. (1984) reported the presence of mermithids Hexamermis sp., Hexamermis albicans, Mermis sp. and Gastromermis sp. in C. septempunctata, they were only found in 0.4% of 10,350 insects examined, which included Hypera nigrirostris, Pentatomidae, two species of Tettigonidae and one locust. It was suggested that the climatic conditions of the areas studied (Bukhara and Navoi regions of Uzbekistan, USSR) were contributory to the low prevalence of infection.

Koyama and Majerus (2008) studied interactions between the parasitoid Dinocampus coccinellae and Harmonia axyridis and C. septempunctata. They stated that in Japan, both coccinellids are hosts to the wasp, but a higher proportion of C. septempunctata are successfully parasitized and reported that this was also the case in Britain, to a greater extent.

The eggs of C. septempunctata may come under attack from lacewings and coccinellids, both this and other species (Majerus and Kearns, 1989).

Means of Movement and Dispersal

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Natural Dispersal (Non-Biotic)

The larval and adult stages are mobile and thus able to disperse locally. It has been reported that since its establishment in North America, C. septempunctata has been found hundreds and thousands of kilometres from its original release sites (Krafsur et al., 2005).

Accidental Introduction and Intentional Introduction

C. septempunctata has been widely introduced as a biological control agent for the control of Aphis gossypii. Attempts to introduce it into North America between 1956 and 1971 failed, but it was subsequently found in New Jersey, USA where it is thought to have been accidentally introduced (Angalet and Jacques, 1975; Majerus, 1994).

Pathway Causes

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Cause	Long Distance	Local
Biological control	Yes	Yes
Escape from confinement or garden escape		Yes
Horticulture	Yes	Yes

Pathway Vectors

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Vector	Notes	Long Distance	Local	References
Mail	Adults	Yes	Yes
Plants or parts of plants		Yes	Yes

Plant Trade

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Plant parts liable to carry the pest in trade/transport	Pest stages	Borne internally	Borne externally	Visibility of pest or symptoms
Leaves	adults; eggs; larvae		Yes	Pest or symptoms usually visible to the naked eye

Impact Summary

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Category	Impact
Economic/livelihood	Positive
Environment (generally)	Positive and negative
Human health	Negative

Environmental Impact

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Impact on Biodiversity

Hoddle (2004) states that C. septempunctata has influenced the distribution and abundance of native coccinellid competitors by reducing their survivorship in local habitats, influencing dispersal dynamics and habitat use. For example, Elliott et al. (1996) documented the effect invasion by C. septempunctata had on native coccinellids in South Dakota, USA on lucerne, maize and small grains. Coccinellids were observed 13 years post invasion and 5 years after. It was found that the structure of native coccinellid communities differed significantly for years before compared to years after invasion.

Turnock et al. (2003) studied the abundance of some coccinellids before and after the introduction of C. septempunctata to Manitoba, USA in 1988. The relative abundance of the more common native coccinellids was determined from D-Vac Insect Net® and sweepnet samples in lucerne in 1983-2001, by sweepnet and visual sampling in field crops and other vegetation in 1989-2001, and by transect sampling of aggregations of coccinellines in spring and autumn on the shore of Lake Manitoba from 1989 to 2001. It was found that the relative abundance of Coccinella transverso guttata, Hippodamia convergens, Hippodamia parenthesis, and Coccinella trifasciata has decreased since the establishment of C. septempunctata. It was thought that the decline in abundance of these species was caused by their competitive displacement by C. septempunctata.

Similarly, a study by Brown and Miller (1998) reported that C. septempunctata was dominant in the fauna of the tribe Coccinellini from 1989 to 1994 in eastern West Virginia on apple. However, Harmonia axyridis dominated in 1995 and continued to do so in the guild coccinelline on apple. Furthermore, Brown and Miller (1998) reported a displacement of C. septempunctata with H. axyridis, which provided better control of Aphis spiraecola on apple in the study area.

More recently, in South Dakota, Hesler and Kieckhefer (2008) reported the decline of three previously common ladybirds (Adalia bipunctata, Coccinella transversoguttata richardsoni and Coccinella novemnotata), while the invasives, C. septempunctata and H. axyridis had established there.

Gardiner et al. (2009) carried out a four-USA-state wide survey of native and exotic coccinellids, including C. septempunctata and concluded that grassland dominated landscapes with low structural diversity and low amounts of forested habitat may be resistant to exotic coccinellid build-up, particularly H. axyridis and therefore represent landscape-scale refuges for native coccinellid biodiversity.

Snyder et al. (2004) discuss intraguild predation (IGP) and successful invasion by exotic ladybirds. Their results suggested that larvae of native species face increased IGP following invasion by C. septempunctata and H. axyridis, which may contribute to the speed with which the exotics displace the natives after invasion.

Furthermore, in a study to determine under what conditions an invasion results in displacement or co-existence, and to determine rules that underlay combinations of IGP and intraspecific predation (ISP), Rijn et al. (2005) stated that ISP and IGP seemed to be related to larval size differences, where overall the larger species had a greater overall advantage. However, the authors also surmise that larger larvae require more food thus having a disadvantage in terms of resource competition. It was concluded that the estimated levels of ISP and IGP and competitive ability of interacting species could not fully explain invasion by H. axyridis and C. septempunctata.

Threatened Species

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Threatened Species	Conservation Status	Where Threatened	Mechanism	References
Adalia bipunctata (twospotted lady beetle)	No Details	South Dakota	Predation	Hesler and Kieckhefer (2008)
Coccinella novemnotata (ninespotted lady beetle)	No Details	South Dakota	Predation	Hesler and Kieckhefer (2008)
Coccinella transversoguttata (lady beetle, transverse)	No details	Manitoba	Predation	Turnock et al. (2003)
Coccinella transversoguttata richardsoni	No details	South Dakota	Predation	Hesler and Kieckhefer (2008)
Coccinella trifasciata	No Details	Manitoba	Predation	Turnock et al. (2003)
Hippodamia convergens (lady beetle, convergent)	No Details	Manitoba	Predation	Turnock et al. (2003)
Hippodamia parenthesis	No Details	Manitoba	Predation	Turnock et al. (2003)

Social Impact

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During mass occurrences of C. septempunctata, people have been reportedly bitten by this species (Eichler, 1971). The population explosion of ladybirds, mostly C. septempunctata, in Britain in 1976 is frequently recounted. During this time there were numerous reports of people being attacked by them. The ladybirds were in search of food and biting into anything they landed on to test suitability as a food source. On biting, they inject a small amount of digestive enzymes, which on reacting with the chemical defences of the body produce a stinging sensation (Majerus and Kearns, 1989). Other reports of this species being a nuisance are from the German Baltic Sea coast (Dambeck, 2009) and Albena, Bulgarian Black Sea coast (Krell and Britton, 2009).

Risk and Impact Factors

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Invasiveness

Proved invasive outside its native range
Has a broad native range
Abundant in its native range
Is a habitat generalist
Capable of securing and ingesting a wide range of food
Highly mobile locally
Has high reproductive potential
Gregarious

Impact outcomes

Conflict
Reduced native biodiversity
Threat to/ loss of native species

Impact mechanisms

Competition - monopolizing resources
Predation

Likelihood of entry/control

Highly likely to be transported internationally accidentally
Highly likely to be transported internationally deliberately

Uses

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Economic Value

C. septempunctata has been widely introduced as a biological control agent for the control of aphid pests under glasshouse conditions (e.g. Yarkulov, 1978; Valério et al., 2007). The ability of C. septempunctata to regulate and control aphid pests in the field has been well documented, often as part of a natural enemy complex (e.g. Shanthi et al., 2009). It is often listed as a natural enemy occurring under field conditions in a wide variety of crops, such as pomegranate orchards (Öztürk et al., 2005), sour cherry orchards (Prunus cerasus) (Özkan et al., 2005), mustard fields (Vekaria and Patel, 2005), and ground nut (Sahayaraj and Martin, 2003) and horse bean (Vicia faba minor) (Sadej, 2000) crops, to name but a few.

After studying field and computer generated data, Gosselke et al. (2001) concluded that without antagonists, including C. septempunctata, winter wheat-cereal aphids (Sitobion avenae, Rhopalosiphum padi and Metopolophium dirhodum) would cause economically important yield losses in two out of three cases. As part of a predator complex, C. septempunctata prevented Aphis fabae from exceeding economic damage thresholds in sunflower in Romania (Voicu et al., 1987).

In contrast, Immik et al. (2004) studying the biological control of woolly aphid, Eriosoma lanigerum, stated that the release of C. septempunctata, Chrysoperla carnea and Adalia bipunctata larvae had little effect on the aphid population and could not be economically justified.

The value of C. septempunctata as a pollinator of the endemic and endangered plant, Disanthus cercidifolius, in China (Xiao et al., 2009), and in mango (Singh, 1989) has also been documented.

Uses List

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Environmental

Biological control

Similarities to Other Species/Conditions

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Marzo (1982) studied the spermatheca of Rhynchota and Coleoptera, including C. septempunctata and reported that even among species not systematically related, similarities were observed in the shape of the spermathecae; seven main forms were found.

In a study of reflex bleeding in ladybirds, Jong et al. (1991) discussed the phenomenon in relation to the possibility that Adalia bipunctata is a Batesian mimic of C. septempunctata.

Prevention and Control

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Due to the variable regulations around (de)registration of pesticides, your national list of registered pesticides or relevant authority should be consulted to determine which products are legally allowed for use in your country when considering chemical control. Pesticides should always be used in a lawful manner, consistent with the product's label.

Control

Biological control

Roy et al. (2008) studied the effect of Beauveria bassiana on Harmonia axyridis, C. septempunctata and Adalia bipunctata. Results showed that 80% mortality of C. septempunctata was achieved with a dosage of 109 conidia ml^-1.

Chemical control

The ability of C. septempunctata to reduce pest aphid populations under natural conditions in the field and in protected environments is widely recognised and this species has been extensively introduced as a biological control agent for the control of aphid pests. Therefore the effects of chemicals on this coccinellid are in the context of non-target effects under laboratory conditions (e.g. Ullah, 1977 in part; Ahmad and Ahmad, 2009) and in the field (e.g. Alexandrakis et al., 2005). For example, Ullah (1977) studied the effects of pesticides to control A. gossypii on C. septempunctata and found monocrotophos to be highly toxic to the predator. Similarly, Ahmad and Ahmad (2009) reported that fenazaquin and quinalphos were highly toxic to adults of C. septempunctata.

References

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